Download Biosystematic Studies of Phacelia capitata

AN ABSTRACT OF THE THESIS OF
J.
Shelly
Steph n
Botany
Plant
and
the degree of Master of Science in
for
Pathology
Biosystematic
Title:
presented
Studies
of
on April 11. 1985.
Phacelia
capitata
(Hydrophyllaceae).
a Species Endemic to Serpentine Soils in
Southwestern Oregon
Redacted for privacy
Abstract approved:
Renton L. Chambers
Phacelia
Phacelia
capitata
magellanica
Magellanicae),
perennials
American
this
western
in
secunda
P.
of
North
P.
(=
is
member
a
of
wide-ranging,
the
group
(species
complex
polyploid
group
a
polymorphic
America related to the South
magellanica).
The purpose of
was to determine, to the extent possible, the
project
interrelationships
studies
using
Rruckeberg
of
P. capitata within the Magellanicae,
of
its
distribution, cytology, morphology,
reproductive biology, and population ecology.
Phacelia
and
capitata is endemic to ultramafic (serpentine
peridotite)
outcrops,
substrates,
in
southwestern
Oregon.
generally
native
Field
and
surveys
metamorphic
Jackson
Counties,
revealed
22 extant
The species occurs on open serpentine slopes,
populations.
outcrops,
Douglas,
Coos,
similar
or
and
roadbanks
or
similarly
disturbed
sites,
in areas with a south to southeast exposure.
The
plant community with which P. capitata is frequently
associated is a Pinus ieffreyi/grass savanna.
from five P. capitata sites and two E. corvmbosa
Soils
were
sites
the
chemically analyzed.
soils
were
materials,
and
phosphorus
and
compared
were
from
found
very
serpentinized parent
fully
possess
to
calcium/magnesium
low
levels
low
ratios
of
when
to an eighth sample from a non-serpentine soil.
sample
soil
derived
At six of these locations
metamorphic
chemical
from
one
similar
rock
capitata
P.
site,
A
derived from a
in appearance to serpentine, had
characteristics
intermediate
between
those
of
serpentine and non-serpentine soils.
Chromosome
revealed
counts
from
existence
the
tetraploid
one
of
P.
capitata
populations
diploids (n=11) at 19 of them;
of
were
counts
(n=22)
populations,
20
these
obtained
also
populations
from
two
containing
diploids.
Taximetric
populations
capitata
readily
mostly
capitate
verify
when
heterophylla
studies
of
living plants from 26 Phacelia
the morphological distinctiveness of P.
compared to P. corvmbosa, P. hastata, and P.
ssp.
virgata.
Phacelia
capitata
is
most
distinguished from the other species by its narrow,
entire,
or
silvery-pubescent rosette leaves, its often
sub-capitate
cyme
branches,
and its sparse,
obscure glandulosity.
the
Significant
seed
absence
insect visitors.
collect
of
nectar
from
set in P. capitata does not occur in
the
When natural pollinators
congested
flowers, it is likely
that
self-pollination
inflorescence
predominant
is
cross-pollination
Magellanicae,
between
occurs
to
adjacent
of
an
However,
(geitonogamy).
some extent in members of the
evidenced
as
flowers
hybridization
by
and
introgression at the tetraploid level.
The
origin
problematic.
interbred
with
There
may
no
It
the
is
polyploids
possible
of
capitata
P.
is
P. capitata may have
that
in the past with P. Jeterophvlla ssp. virciata, or
more
a
of
southerly
member
the complex, P. egena.
of
also some morphological evidence that P. capitata
is
have occasionally hybridized with P. hastata.
plants
have
However,
been found that suggest any hybrid contact
between E. capitata and Z. corymbosa.
many
At
numbers
this
of
locations
been
has
increase
an
in
capitata following habitat disturbance, and
P.
appears
there
to
have
from
any
characterized its populations.
The
"released"
species
the
paucity
that
species
evidently possesses poor competitive ability in the
more
formerly
vegetation
closed
and
soils,
also
appears
which
to
exists
on
non-serpentine
be a poor competitor even in
some undisturbed serpentine communities.
Owing
capitata
should
very
to
is
be
narrow
its
not
favorable
endangered
maintained
geographic
response
to disturbance, P.
with extinction.
However, it
on a monitoring list, because of its
distribution
serpentine or sub-serpentine soils.
and
restriction
to
Biosystematic Studies of Phacelia capitata
(Hydrophyllaceae), a Species Endemic to
Serpentine Soils in Southwestern Oregon
by
J. Stephen Shelly
A THESIS
submitted to
Oregon State University
in partial fulfillment of
the requirements for the
degree of
Master of Science
Completed April 11, 1985
Commencement June 1985
APPROVED:
Redacted for privacy
Professor of Botany and Plant Pathology in charge of major
Redacted for privacy
Chairman of department of Botany and Plant Pathology
Redacted for privacy
Dean of Gradua
School
1
Date thesis is presented
Typed by
April 11. 1985
J. Stephen Shelly
ACKNOWLEDGEMENTS
wish
I
Chambers,
thank
to
his
for
major
professor, Dr. Kenton L.
and guidance during all phases of
help
His encouragement, and his frequent sharing
project.
this
my
of knowledge, are greatly appreciated.
wish
also
I
committee,
Robert
Drs.
Petersen,
thank
to
other
the
Frenkel,
members
Wilson,
Mark
of
my
and Roger
the careful reading of the thesis which they
for
did on fairly short notice.
specimens were kindly provided by curators of the
Loan
following
of
Museum of Natural History, University
herbaria:
Oregon
California
(ORE);
Herbarium
(CAS/DS);
Academy
University
of
Sciences/Dudley
Washington
of
(WTU);
Humboldt
State
University (HSC); University of California,
Berkeley
(UC);
and
the Jepson Herbarium of the University
of California (JEPS).
Thomas
Moore
C.
Pathology
and
gratefully
provided
the
of
Oregon
the
acknowledged.
by
and
research,
Support,
the
travel funds, provided by Dr.
Department
of
Botany and Plant
State University Herbarium, are
Funds for computer analysis were
Committee for Unsponsored Research, Milne
Computer Center, Oregon State University.
Insect
Andrew
R.
taxa
were
Moldenke,
identified
by Drs. John D. Lattin,
and William P. Stephen, Department of
Entomology, Oregon State University.
Assistance
with
the computer analysis was provided by
Drs. C. David McIntire, Robert Frenkel, and Steve Broich.
enjoyable day of field work was spent with Mr. Russ
An
Holmes,
Bureau
of Land Management (Roseburg), who provided
information on locations of Phacelia capitata sites.
owe special thanks to LaRea Johnston, for her advice
I
and
expertise
included
completion
assistance
also
go
Bob
Special thanks
befriended me soon after my
who
and
who
and
has taught me much about the
the state; and Mark Nelson, who provided valuable
of
field
Meinke,
work,
photographic
the
of
Oregon,
in
These have especially
respects.
purchasing my pick-up truck!
in
to
arrival
flora
many
in
assistance,
made
and
many
interesting
Phacelia
collections in southwestern Oregon.
Without
of
many
thesis
students, past and present, this
graduate
fellow
advice, and companionship
encouragement,
the
These very special
could never have been completed.
many
among
friends,
Fredricks,
Dan
Joe Antos, Nancy
I am sure, include:
Luoma,
Teresa
Magee,
Bill
Ripple, Glenn
Stewart, and Carolyn Wright.
My
unselfishly
wonderful
thanks are especially for Ginger King, who
sincere
provided
patience
encouragement,
(which
I
fear
I
and
listened
may
have
with
tried on
occasion!) to all my concerns over the last six months.
This
Shelly,
thesis
is dedicated to my parents, John and Lois
for all their love, support, and understanding, and
for
fostering
my interest in the natural world.
am
grateful
for
experienced.
the
gifts
of
all
I
have
To them I
seen
and
TABLE OF CONTENTS
INTRODUCTION
1
TAXONOMIC RELATIONSHIPS
3
ENVIRONMENTAL RELATIONSHIPS
Geology
Soils
Climate
Site locations and characteristics
Associated vegetation
Vegetation history
13
13
16
23
BIOSYSTEMATIC STUDIES
Cytology
Morphology of the species
Taximetric studies
Scanning electron microscopy
Reproductive biology
Flowering and pollination biology
Seed production
Pollen viability
Competitive ability and ruderal response
45
45
48
48
24
32
41
71
78
78
86
90
93
DISCUSSION AND CONCLUSIONS
Speciation and evolutionary relationships
The endemic status of the species
Recommendations for preservation
99
99
110
111
BIBLIOGRAPHY
113
APPENDIX
118
LIST OF FIGURES
Figure
Page
1.
The phacelia maoellanica polyploid complex
(species group Magellanicae).
2.
Distribution of Phacelia capitata in southwestern
Oregon.
15
3.
Cluster analysis of Phacelia species.
69
4.
Seed of Phacelia capitata.
73
5.
Seed of Phacelia corymbosa.
75
6.
Pollen and leaf vestiture of Phacelia capitata.
79
7.
Stem vestiture of Phacelia capitata and P.
corvmbosa.
81
8.
Distribution of Phacelia capitata and E. corymbosa
in southwestern Oregon.
104
9.
Possible interrelationships of phacelia capitata
within the species group Magellanicae.
109
8
LIST OF TABLES
Table
Page
1. Selected soil analyses for eight Phacelia sites in
southwestern Oregon.
22
2. Climatic data for selected weather stations in
southwestern Oregon.
25
3. Reported locations of Phacelia capitata in Oregon.
27
4. Associated tree and shrub vegetation at Beatty
Creek.
37
5. Herbaceous vegetation at Beatty Creek.
38
6. Chromosome numbers for Phacelia species group
Magellanicae in southwestern Oregon.
47
7. Study populations of Phacelia corymbosa, P.
hastata, and P. heterophylla in southwestern
Oregon.
50
8. Means and standard deviations for ten
morphological characteristics measured on living
plants from 26 Phacelia populations.
52
9. Climax leaf lobing in Phacelia capitata.
56
10. Floral branching below the terminal cymes in
Phacelia capitata.
62
11. Comparison of Phacelia capitata, P. corymbosa, P.
hastata, and P. heterophylla ssp. virgata in
southwestern Oregon.
72
12. Insect pollinators of Phacelia capitata and P.
corymbosa.
87
13. Seed production in open-pollinated and enclosed
inflorescences of Phacelia capitata.
89
14. Pollen viability in Phacelia species group
Magellanicae.
91
15. Ruderal response of Phacelia capitata at Lee
Creek.
97
BIOSYSTEMATIC STUDIES OF PHACELIA CAPITATA
(HYDROPHYLLACEAE), A SPECIES ENDEMIC TO
SERPENTINE SOILS IN SOUTHWESTERN OREGON
INTRODUCTION
Phacelia
belonging
group
plants
perennials,
ovary
of species in
group
affinities
having
morphologically
are
subgenus
of
divided
a perennial species
with
species P. secunda (=P. magellanica).
group
the
is
wide-ranging
a
America
North
American
Kruckeberg
the P. magellanica polyploid complex (species
to
Magellanicae),
western
of
capitata
which
and
with
leaves
with
four
The members
defined as being those
which
Phacelia
the South
are
biennials
or
possess entire leaves, or pinnately
divisions or leaflets, and an
entire
ovules,
two
to
each placenta (Heckard,
1960).
The
members
polyploid
species
pillar
occurring
including
of
the species group Maaellanicae form a
complex which has, as its foundation, six
at
the
P. capitata.
however,
Heckard
base chromosome number of x=11,
In his detailed study of the group,
made
(1960)
no attempt to determine the
interrelationships of P. capitata within the complex.
Phacelia
over
to
a
its
capitata
small
rarity,
endangered
is
restricted
to serpentine soils
geographic area in southwestern Oregon.
Due
it has received attention as a potentially
species and has been placed on various state and
federal lists of such species.
2
This
study was undertaken to assess various aspects of
the
distribution,
cytology, ecology, reproductive biology,
and
morphological
characteristics
extent
the
were
possible,
natural
of P. capitata.
populations
To the
of other species in
complex (P. corvmbosa, P. hastata, and P. heterophylla)
considered
interrelationships
In
addition,
be
used
in
of
an
attempt
to
determine
the
P. capitata within the Maciellanicae.
it is hoped that the information obtained can
to evaluate the endangerment status of P. capitata
and to make recommendations for managing the species.
3
TAXONOMIC RELATIONSHIPS
The
Hydrophyllaceae
widespread
family.
regarding
Munz
genera
and
stated
that
and
300
but
genera
and species it contains.
of
that
the
family contains about 25
More recently, Cronquist (1981)
species.
Hydrophyllaceae comprises about 20 genera
the
species.
250
small
differ in their opinions
suggested
(1959)
relatively
a
Taxonomists
number
the
forms
Constance and Chuang (1982) refer to 18
genera and approximately 326 species.
family
The
western
United
by
recognized
to
North
with
Hydrolea
Constance
which
Nama
in
habitats of the
dry
have
Of the 18 genera
(1963), 16 are primarily confined
(exceptions
are
outlying
represented
is
common
States (Cronquist, 1981).
America,
Wigandia,
most
is
Nama,
species
Phacelj.a,
and
in South America,
as well from the Hawaiian Islands).
pantropical,
and Codon is restricted to South
Africa.
The
history,
Hydrophyllaceae
which
was
had
has
reviewed
by
varied
a
Constance
taxonomic
(1963) and is
summarized as follows.
Jussieu
are
now
(1789)
Boraginaceae,
Phacelia,
and
Convolvulaceae.
family
for five of the genera that
assigned to the family.
(Hydrophyllum,
genera
accounted
two
Brown
and
( Hydrolea
(1818)
Of these, he placed three
Ellisia)
and
in
the
family
Nama) in the family
separated the former three
as the order Hydrophylleae and the latter two as the
Hydroleae.
The
Hydroleae,
as
monographed
by
4
Choisy
contained
(1833),
Eriodictyon
Wigandia;
Hvdrolea, Nama, Romanzoffia, and
was added later (1846).
De Candolle
(1845) described the genus Codon.
Finally,
family
and
above
were united into the
genera
by Gray (1875).
He added the genus
later Lemmonia, and divided the family into
Since that time, the familial relationships
tribes.
four
the
Hydrophyllaceae
Tricardia,
of
all
the Hydrophyllaceae have been debated, surely due to the
fact
that,
the
family
Constance
as
(1963) stated, "(t)0 some extent
appears to be a collection of morphological and
geographical
odds-and-ends."
Hydrophyllaceae
classification
under
Baillon (1890) submerged the
the
Boraginaceae,
this
but
Brand (1913) restored the
was not accepted.
Hydrophyllaceae to family status.
More
(1976)
recently,
and
difficulties
relationships
the
family
it
closest
Cronquist
with
(1981)
the
the
illustrate
interpreting
in
of
classifications by Thorne
contrasting
continuing
closest
the
Hydrophyllaceae.
familial
The former aligns
the Boraginaceae, while the latter places
the
to
Polemoniaceae.
Cronquist
(1981)
summarizes his interpretation of the problem:
The
been
Hydrophyllaceae
have
usually
considered to be allied to the Polemoniaceae on
the one hand,
and to the Boraginaceae, on the
other.
I
agree.
For purposes of formal
classification, I find it useful to associate the
Hydrophyllaceae with the Polemoniaceae, and to
put the Boraginaceae in an allied order. This
arrangement helps in the construction of orders
that have a reasonable degree of morphological
coherence.
Despite
its
problematic
relationships,
the
5
Hydrophyllaceae
The
members
shrubs,
and
more
often
sympetalous,
cleft
nearly
variously
in
the
to
many
as
the
flowers
or dissected,
are solitary, or
often helicoid cymes,
modified,
and generally pentamerous; the calyx
or more commonly to the base or
middle,
as
alternate
and
with
the
corolla lobes,
toward the base or well up in the tube, very often
flanked
by
carpels
united
a pair of small scales; the gynoecium is of two
to
terminal,
form
mostly
half-inferior,
a
compound,
unilocular
superior
ovary;
the
or rarely
style
is
usually deeply bifid, sometimes cleft essentially
the base, or undivided; the stigmas are mostly capitate;
placentae
is
compound,
so; the corolla is regular or nearly so; the stamens
attached
to
leaves are
or wholly opposite, simple
cleft,
compound;
the
vestiture;
in
partly
pinnately
to
by the following traits.
seldom (Eriodictvon, Wiciandia)
or
rough-hairy
palmately
perfect,
are
herbs
sometimes
or
entire
rarely
is
are
often
alternate
characterized
is
are
generally
two,
parietal or sometimes axile; the fruit
a capsule, or sometimes irregularly dehiscent
or indehiscent (Cronquist, 1981).
Constance
the
genus
Phacelia
for
the
Phacelia.
and
Eutoca.
genera
Miltitzia.
into
(1963)
the taxonomic history of
Bentham (1837) recognized two genera,
De Candolle (1845) was responsible
Microgenetes
(=Euglypta), Cosmanthus, and
Gray (1875) combined all of these but Miltitzia
Phacelia.
cytological
summarized
and
Further
classifications,
based
on
other evidence, were subsequently produced
6
by
various
suggested,
(1963)
surveys,
with
students
that
Howellanthus
recently,
(=Microcenetes),
classification
three
subgenera Cosmanthus
Phacelia
and
the
of
(x=11)."
by
forth
Eutoca,
genus:
and
Miltitzia,
put
can be achieved
Most
al. (1984) recognize four sections
et
treatment
of extensive cytological
arrangement
the
of
(x=8),
Cronquist
their
in
basis
"...natural
a
recognition
the
(x=9),
the
on
Finally, Constance
genus.
the
of
However, the
Phacelia.
Constance
Euglypta
will
be followed
here.
Phacelia
with
perennial,
borne
helicoid
compound
base;
corollas
shortly
often
the flowers are
aggregated into a
the calyx is divided nearly to the
variously blue or purple to pink,
are
the corolla tube near its base; the styles are
to
to very deeply cleft; the capsules are loculicidal;
leaves
wholly
phakelos,
these
Chuang,
of annual and
array
species;
and
yellow, and tubular or rotate; the filaments are
or
attached
or
cymes,
(Constance
diverse
a
herbaceous
inflorescence;
the
white,
of
taprooted
in
species
200
consists
It
the largest genus in the family,
far
approximately
1982).
the
by
is
entire to pinnately dissected, and chiefly
are
alternate.
meaning
The genus name comes from the Greek
fascicle, and referring to the congested
inflorescences (Hitchcock and Cronquist, 1973).
Phacelia
Magellanicae,
comprises
the
capitata
in
the
is
a
member
of the species group
subgenus Phacelia.
well-known
The Magellanicae
P. magellanica polyploid complex
7
(Figure
This
1).
Heckard
1942,
Kruckeberg,
among
1944,
studies
by
and
(Cave
1959; Heckard, 1956;
1950,
1947,
studied
intensively
was
cytological
after
(1960)
Constance,
complex
1956) that revealed the existence of polyploidy
Superimposed on a series of six diploid
its members.
interfertile complex of tetraploids
(n=11)
species
(n=22)
and
study,
Heckard (1960) stated that each of the basic diploid
an
is
few
a
species,
except
similar
counterpart
intimately
hexaploids
P.
capitata,
on
associates
it
tetraploids."
polymorphic
tetraploid
a
At the time of his
"...morphologically
superstructure
with
the
This
intergradation
1949),
often
indecipherable,
has
which
level
tetraploid
species,
(Anderson,
sometimes
and
has
the
intergrading
introgression
(n=33).
of
between
accompanied
by
led to very complex,
patterns
of
variation.
Heckard (1960) summarized the problem:
For
the most part, as
is
characteristic of
introgressing taxa, entire series of populations,
which sometimes extend throughout large areas,
are
involved
in
the
genic recombinations,
resulting
in
a
variation
pattern that is
extremely
difficult
to
analyze.
This is
especially true in this complex where the initial
differences are not of a magnitude to furnish
characters which can be conveniently measured.
The result of this intergradation is an extensive
assemblage of interrelated plants within which
lines must be drawn somewhat arbitrarily in order
to delimit taxonomic units.
As
has
a
result
had
a
of
such variation patterns, the Magellanicae
protracted
taxonomic
history.
At one time or
another,
the
complex has been said to include five or more
species,
and
as
few
as
three
species.
In
his final
DASYPHYLLA
F RIGIDA
DASYPHYLLA
SIMULANS
-frMOPOLA
ILEPTOSEPALA
HEXAPLOID
C OMPACTA
HASTATA
E GERA
CALIFORNICA U AR GENTEA
TETRAPLMD
PATULA
IMBRICATA
I
I
CORYMBOSA
HET EROPHYLLA
MUTABILIS
0 RE ON EN SIS
---]
CA PIT AT A
PATULA
1
t
I MBR CAT A
CORYMBOSA
IHETEROPHYLL A
17
IBERNARDINA
Figure 1.
MUTABILIS
NE M ORALIS
DIPLOID
VIRGATA]
The Phacelia magellanica polyploid complex (species group
Magellanicae).
From Heckard (1960).
oo
9
taxonomic
treatment,
(1960) recognized 13 species
Heckard
in the United States and Canada.
The
made
Roseburg,
near
annotated
Herbarium,
egena."
In
in
his
mentioned
(1956)
1919, as "Phacelia
description
of
P.
the resemblance of
the new species, but he assigned no name
to
suggestive
features
the
of
P. capitata, it lacks the inflorescence
of
capitata
P.
from
It was subsequently
Co."
Although the silvery, entire leaves of this plant
it.
are
Gray
forma
specimen
to
Douglas
Kruckeberg
capitata,
of
the
at
californica
this
of a P. capitata-like plant was
June 22, 1916 by Peck (6022, WILLU), from a "(d)ry
on
slope
collection
first
species.
The possible interrelationships
with this, and other problematic specimens
Roseburg-Myrtle
the
Creek area, will be discussed in
the final section of this thesis.
Other
collections
made
Douglas
in 1916 by Peck (6033, "(d)ry
and 6035, "Reston, Douglas
bank,
Roseburg,
Co.",
both WILLU), were also discussed by Kruckeberg (1956)
in
relation
seem
Heckard
P. capitata.
to
best
to
Co.",
However, these two specimens
fit the description of P. hastata offered by
The Reston vicinity was searched during my
(1960).
study, but only individuals of P. hastata were located.
The
April
Co.,
13,
first collection of "pure" P. capitata was made on
1934,
Oregon,"
"2
by
specimen
went
capitata
Kruckeberg
miles
Eastwood
unidentified
in
ed.
south of Myrtle Creek, Douglas
and
until
in
Howell (1474, CAS).
it
This
was annotated as P.
1955, by both Heckard and
10
Kruckeberg.
On
Eastwood
Howell
from
and
"Myrtle
originally
the
labeled
its
name
such.
as
capitata,
relocate
date
Douglas
Heckard
"(p)ossibly
Oregon,"
Co.,
P. leucophvlla.
resemblance
similar
as the above collection,
obtained another specimen (1475, CAS)
Creek,
acknowledged
it
same
to
they
Once again, Kruckeberg
P.
capitata but did not
annotated
in 1960 as P.
it
a tetraploid form?"
specimens
which
in the field.
I was unable to
A hybrid origin
for such a plant is possible.
The
next
(20324,
UC),
"stony
slope
went
P.
"good"
collection
was made in 1939 by Peck
presumably from the eventual type locality, a
near
Bridge,
unidentified
until
Coos
Co."
This specimen also
it was annotated by Kruckeberg as
capitata in ed. in 1954, and he eventually designated it
as a topotype.
A
DS)
specimen
in
1942,
labeled
this
collection
of
recognize
apitata
by
commented
genotypes
locate
found
from
by Constance and Rollins (2956,
Mt. Nebo near Roseburg, was originally
leucophvlla.
P.
reminiscent
not
collected
was
Like
Eastwood and Howell 1475,
acknowledged
as
morphologically
E. capitata by Kruckeberg, but again he did
it
as such.
Heckard,
(pers.
It was annotated in 1961 as P.
and although it was a tetraploid, he
comm.)
that
the
plant
of E, capitata and P. corymbosa.
may
combine
I was unable to
individuals similar to this in the field, the plants
on
hastata.
Mt.
Nebo
being
most
readily
referable
to P.
11
The
next
collections
County
Douglas
ORE),
(5314,
ORE).
of
papitata
P.
were
made in
by Detling, first in 1942 from Red Mountain
and
These
then in 1949 from Nickel Mountain (6339,
specimens were also originally labeled as E.
leucophylla,
until
annotated
being
as
capitata
P.
by
Kruckeberg in 1961.
The
1951
collection
holotype
(Kruckeberg
miles
2703, WTU), from a "serpentine roadcut, 2
Coos
42,
illustrated
1951,
years
five
serpentine
has
outcrops
northwestern
later
(Kruckeberg,
1956).
been
sporadically
collected
Counties,
well
as
Since
Douglas
south-central
in
Jackson
It was described and
Oregon."
County,
capitata
P.
capitata was made in
E.
of Bridge, above Coquille River and along State
east
Highway
of
on
and
being
as
re-collected from the type locality in Coos County.
description
his
In
compared
to
it
leucophvlla.
synonymy
under
one
made
of
other
in
close
two
However,
comparisons
and
of the species, Kruckeberg (1956)
relatives, P. corymbosa and P.
P.
leucophylla
was
Heckard
(1960).
placed
in
Further
P.
hastata
P.
capitata with P. corymbosa, P. hastata,
by
member of the complex, P. heterophvlla, are
later section on the morphology of the species,
a
and a description of P. capitata is presented.
his
In
California
made
within
"(n)o
studies,
members
of
Heckard
(1960)
the Magellanicae.
attempt...to
depict
the
emphasized
the
Consequently, he
interrelationships
the complex of Phace ia capitata Kruckeberg..."
The
12
goals
of
my
study were to consider certain aspects of the
biosystematics,
in
order
to
ecology,
ascertain,
relationships
within
hoped
the
that
and
distribution of P. capitata,
to
the
extent
possible,
its
the species group Magellanicae. It is
study
will
contribute
understanding of this complex group.
to
a
better
13
ENVIRONMENTAL RELATIONSHIPS
Geology
The
geology
the Klamath Mountains in southwestern
of
Oregon
and northwestern California is extremely complex and
poorly
understood.
found
the Klamath Mountains, near the Oregon-California
in
The
border.
thought
age
This
these highly metamorphosed rocks is
section presents a review of the terminology
serpentine
Mountains
of
be at least 425 million years (Alt and Hyndman,
to
1978).
of
Some of the oldest rocks in Oregon are
substrates,
province,
and
a
outline
an
brief
the
of
summary
Klamath
of the current
theories as to how the serpentine areas became emplaced.
Traditionally,
define
The
types
of
ultramafic
all
them.
term
which
in
the
silicates
word "serpentine" has been used to
rocks and the soils weathering from
encompasses all those rock
*ultramafic"
the elemental composition largely consists
of
mineral
olivine.
igneous
rocks
iron
The
and
most
from
magnesium,
derived
common
ultramafics
the
are the
peridotite and dunite, and their metamorphic
derivative, serpentinite (Kruckeberg, 1969).
The
along
the
bordered
south
Oregon-California
the
on
by
portion
the
Klamath Mountains, of which the Siskiyou Mountains
the
of
Oregon
north
by
state
line
are
a part, are
the Oregon Coast Range, on the
California Coast Range, and on the east by a
the
Coast
Cascade
Range
Range.
The northern contact with
extends along a line southwest of
14
the
Bandon
the
Roseburg
rocks
vicinity.
The contact with younger volcanic
the Cascades runs south from the Roseburg area to
of
southeast
Medford,
to
area, to the Rogue River, and then northeast to
Redding, California.
Mountains
the mountains near Siskiyou Pass,
over
The southern border of the Klamath
traverses
then
southwest,
northwest,
thence
almost to the California-Oregon state line (McKee,1972).
ultramafic
The
(and
corvmbosa
P.
Curry,
Josephine,
(Figure
2).
found
in
Jackson,
and
Coos,
throughout
scattered
Douglas
Counties
most extensive, contiguous outcrops are
southern
Curry and Josephine Counties (i.e., the
Peridotite
expansive
are
Oregon)
in
The
Josephine
sheet).
To
the
north
these
of
outcrops, the ultramafic areas become more widely
distributed,
in size, and separated by wider areas
smaller
non-ultramafic
of
outcrops occupied by Phacelia capitate
There
substrates.
is
an
overall
southwest-northeast trend in the alignment of the outcrops.
The
of
geology
Oregon,
the
North
began
Era),
and
floor
rest
precursors
the
200
continues
on
it
a
continental
million
today.
years ago (early Mesozoic
Muddy sediments on the ocean
which in turn rest on
basalt
lava
flows,
green
rock
that forms when heavier rocks
absorb
are
and the Pacific Ocean floor
continent
about
serpentinite,
beneath
the result of a prolonged collision between
is
American
which
the Klamath Mountains, and indeed all
of
water.
peridotite.
margin,
These
heavier
serpentinite
As the seafloor slides under
the
muddy
sediments,
and
15
DIPLOID
* TETRAPLOIDS PRESENT
n UNKNOWN
0 TYPE LOCALITY
UNVERIFIED SITE
CUMATE STATION
CALIFORNIA
Figure 2.
Distribution of Phacelia capitata in southwestern Oregon.
Ultramafic formations
outlined (Wells and Peck, 1961).
16
occasionally
peridotite
ranges
slab
slabs
and
ultramafic
soft
along
of
Serpentinite is
and easily deformed that it can be pressed up cold
faults.
Under pressure, it takes on the "consistency
in
The
1972).
(McKee,
bodies
ultramafic
intrusion
of
many
Oregon occurred during
southwestern
Jurassic Period, between 190 and 136 million years ago.
However,
and
the
margin,
continental
the
mass (Alt and Hyndman, 1978).
toothpaste"
the
against
When such a
rocks can be squeezed through fault zones in the
accumulating
so
are added to growing coastal
serpentinite,
pressed
containing
seafloor
as the ancient Klamath Mountains.
such
is
bedrock
of
may be remobilized with further compression,
they
some
location
bodies
as'
may
recently
have
as
been
the
moved
to
their present
Eocene Epoch, approximately
30-38 million years ago (Baldwin, 1976).
Soils
The
effects of serpentine soils on the physiognomy and
composition
physical
review
soil
vegetation
of
and
chemical
have
been
properties
related
to various
such substrates.
of
A
of the properties of these soils, and the results of
sample
characterize
analyses,
the
are
edaphic
presented
in
order
to
environments of P. capitata, and
the more southerly serpentine species, P. corvmbosa.
Understanding
vegetation
numerous
has
the
been
attempts
to
soils on
influence
of
serpentine
problematic
in
the past, because of
find
one
single
property of these
17
soils
which
plant
life.
"(t)his
are
universally
explains
unique impact on
their
the words of Proctor and Woodell (1975),
In
has led to unnecessary controversy, for serpentines
exceedingly
variable."
defeats
variability
This
efforts to generalize about the serpentine phenomenon.
Physically,
rocky
(i.e.
matter
and
serpentine
Woodell,
these
and
soil
nutrient
levels
may
The
properties
should
they
1954;
found
are
rarely
However,
diverse parent
of
soils
of
unfavorable
overriding
for
plant
importance; soil
influence (Walker,
1969; Proctor and Woodell, 1975).
It is
to note that, while serpentine soils have often
equated
no
1954).
has been that, while physical
discriminating
the
Kruckeberg,
soils
reduced
in
considered as part of the complex of
serpentine
provides
interesting
been
be
result
can
low clay
texture,
(Walker,
in
exist
conclusion
making
chemistry
exceedingly
physical characteristics
gravelly
topography)
materials.
growth,
mantle,
steep
conditions
factors
Undesirable
1975).
and
moisture
from
azonal soils)(Walker, 1954; Proctor
(i.e.,
shallow
content,
vary
lithosols), to having a high clay and organic
,
content
(i.e.,
soils
with
xeric soil conditions, Whittaker (1960)
significant
serpentine
and
differences in soil moisture between
non-serpentine
soils
in measurements from
the Siskiyou Mountains.
Proctor
chemical
serpentine
and
Woodell
(1975)
evaluated
at length the
characteristics most often cited as the causes for
soil
infertility
and
toxicity.
These
have
18
included
low
phosphorous,
especially
low
levels
potassium;
and
nickel,
levels
the
of
chromium,
calcium
of
nitrogen,
nutrients
major
high
levels of heavy metals,
and
cobalt; and frequently,
coupled
high
with
levels
of
magnesium.
The
vary
effects
with
major nutrient levels on plant growth
of
In some cases,
the species under consideration.
addition
of
increase
in plant growth, while in others no improvement is
seen
are
variability
subscribe
largely
to
serpentine
plant
demonstrations
of
the
soils and of plant responses to
that
low
major
nutrient
levels
the chemical stringency of serpentine
for
example
One
in which nutrient addition increased
is detailed by Turitzin (1982), in a study of
growth
nutrient
view
the
account
soils.
the
best
the
of
These differences in
1975).
Proctor and Woodell claim that few workers would now
them.
In
one
of
to serpentine soil leads to an
K
Woodell,
and
(Proctor
response
and
P,
N,
limitations
on
a
serpentine area in California.
assessing the effects of adding nutrients to the soil on
growth
of two annual grasses (Bromup mollis and Vulpia
microstachys),
were
the
effect
it
limiting
was
found that nitrogen and phosphorous
nutrients,
while
added calcium had no
on the productivity of these species.
however,
possible
variability
in
the
As mentioned,
responses of other
species in a given area should be kept in mind.
A
large body of literature has developed regarding the
biological
effects
of
high concentrations of heavy metals
19
which
frequently
situation
are
highly
is
due
to
pH
(i.e.,
metal
and
weathering,
and
and
modification
other
plant
same
Woodell,
1975).
plants
of
to heavy
In summarizing the
and Woodell conclude that, 1) there
Proctor
growth,
evidence for many serpentine soils and plants that
nickel
other
response
the
by the
toxicity
metal
the effects of nickel, chromium, and cobalt on
on
good
and
lower uptake of organic
and variability (both inter-
elements,
in
of
characteristics
soil
other
potential),
redox
(Proctor
research
content.
factors, including diversity of parent
intraspecific)
metals
to
several
of
metal
heavy
their
in
the ecological significance of heavy metals
in
compounds,
presence
is
variable
origin
rock
Again the
complex, and soils from different sites
very
is
Variability
characterize serpentine soils.
is an important factor, and equally good evidence in
cases
it is of little or no importance; 2) the
that
considerations
nickel,
chromium
there
but
generally
is
less
evidence
that
important in serpentine soils; and 3) there is
is
frequently
about variability apply to chromium as
little evidence for cobalt being as important as
nickel or chromium in such soils.
The
soils
status
has
also
infertility.
often
been
characteristic
low
of
been
A
calcium
and
magnesium
in serpentine
of major interest in assessing their
low Ca/Mg ratio (i.e., less than 1.0) has
cited
as
influencing
the
plant
most
important
growth;
adverse
in other cases,
Ca levels alone are emphasized, and in yet others, high
20
consideration
major
event, these
any
In
1975).
of the opinion that both Ca and Mg should have
are
authors
Woodell,
and
(Proctor
Mg
account
any
in
soil
serpentine
of
infertility.
The
results of White (1971) adequately demonstrate the
a
series of soil-chemical and vegetation analyses
detailed
in
serpentine
Ca
Ca/Mg
low
a
distributions
levels
the
also
the
found
that
within
the
study
overall
He concludes that earlier
Ca
should
be
modified
are
As emphasized earlier, however,
perhaps
best
restricted
species
to
of plant distributions in the area studied.
importance of high levels of magnesium is
acknowledged by Proctor and Woodell (1975).
observations
to
predominant influence of consistently high
results
interpretations
The
low
and
Fe,
these
even
White
infertility based on low Ca/Mg
serpentine
of
Mg, Ni, and Cr.
of
explain
principally influenced by concentrations of
high
incorporate
the
not
could
species
Cr, in that order.
and
explanations
ratio,
It was also found
observed.
individual
of
were
transects
Ni,
alone
ratio
discontinuities
vegetation
Mg,
In some areas, the highest
(1954).
occurred on serpentine soils.
levels
that
No substantiation was found for the
Kruckeberg
and
(1954)
of
which had earlier been espoused by Walker
theory"
Ca
areas
contact
non-serpentine
and
Oregon.
southwestern
"low
He conducted
of the serpentine soil phenomenon.
complexity
that,
1)
many
They cite
serpentine-endemic plant
have a requirement for high amounts of this element
21
optimum
for
serpentine
growth,
soils
non-serpentine
requirement
and
have
been
species.
might
of magnesium in
levels
the
2)
shown
be
to
to many
toxic
In their words, a "high magnesium
be
of
more
general
importance
in
explaining the restriction of some plants to serpentines."
The
of
most
reasonable conclusion is that the stringency
serpentine
chemical
with
correlated
is
not
with
any single
or physical property of such soils in general, but
suite of factors, mainly chemical in nature.
a
factors
the
soils
may
vary
depending
These
on the composition and age of
parent rocks, topography, climate, and the biota of any
given
area.
extent
of
Chemical
their
factors
may vary, as well, in the
interactions with each other (Proctor and
Woodell, 1975).
Table
samples
State
conducted
by the Soil Testing Laboratory at Oregon
University.
The extraction and analysis methods are
outlined
Hill
1 presents the results of analyses of eight soil
I
by
site
derived
but
capitata
metamorphic
are
darker
and
hastata) appear to be
P.
rocks
Soils at the Boomer
that
are
similar
blackish-orange in color.
to
The
sample is from a non-serpentine area adjacent to the
serpentine
samples
and Gardner (1978).
(E.
from
serpentine
Tiller
Berg
outcrops
are
all
along
soils
Elk
derived
Creek.
from
The remaining six
fully serpentinized
parent materials.
serpentine
The
levels
of
phosphorus
soils
are
all
and
low
Ca/Mg
characterized
ratios.
by low
The sub-
Table 1.
Species and population
Selected soil analyses for eight Phacelia sites in southwestern Oregon.
Soil type
pH
P
(ppm)
K
Ca
(ppm)
meq/100g
Mg
meq/100g
Ca/Mg
ratio
Cu
(ppm)
Fe
(ppm)
% Organic
matter
P. capitata
Beatty Creek
serpentine
7.1
4
35
2.1
12.3
0.17
1.1
74
4.7
2.3
1.8
124
5.4
82
17.6
7.7
2
140
3.1
20.0
0.16
2.1
64
4.0
7.5
1
70
1.7
20.4
0.08
2.4
84
1.9
serpentine
6.7
4
31
1.7
10.4
0.16
0.76
86
5.0
Eight Dollar Mtn II
serpentine
6.9
1
39
2.5
23.3
0.11
1.7
94
3.3
Grave Creek
serpentine
7.2
1
39
2.0
11.5
0.17
1.1
40
2.6
non-serpentine
6.8
34
43
6.3
1.2
5.25
1.5
142
2.4
sub-serpentine
6.2
19
Bridge
serpentine
6.9
Elk Creek
serpentine
Peel
Boomer Hill I
P. corymbosa
P. heterophylla
Elk Creek
23
sample
serpentine
typical
values
has
serpentine
reported
higher Ca/Mg ratio than the more
a
but
soils,
is within the range of
it
by
Proctor
and
large
amounts
of
(1975) for such
Woodell
soils.
Although
serpentine
do
not
The
soils
indicate
highest
iron
are
often found in
and Woodell, 1975), the results
(Proctor
for the samples used in this study.
this
concentrations
iron
of
are
found
in
the
sub-serpentine and non-serpentine samples.
The
do
remaining analyses (pH, K, Cu, and organic matter)
not
indicate
serpentine
any
samples
Serpentine
soils
between the
consistent
differences
and
non-serpentine
from
the
the
P.
capitata
sample.
and P. corvmbosa
sites are quite similar in chemical composition.
Climate
The
climate
Mountains,
effects
western
Oregon,
west of the Cascade
maritime, being influenced by the moderating
is
the Pacific Ocean.
of
Cascades
of
largely
The mountain barrier of the
prevents any continental influence in the
western portion of the state.
Along
Annual
2 °
belt
the
coast,
temperatures
climate
the
is
from
1270
elevation
of
the
Coast
mountains
to
the
ocean.
equable.
from a minimum of approximately
range
C, to a maximum of approximately 20°C.
ranges
quite
mm
to
Range
2540
and
Freezing
Rainfall in this
mm, depending on the
the
nearness
of the
weather and snow are
24
rare.
Inland
from
precipitation
annual
the coast, there is a general decline in
amounts,
variation.
temperatures
and
show
a greater
These trends are influenced by higher
elevations in certain areas.
The
the
inland
Coast
and
generally
which
regions
Cascade
may
from
Normal
uncommon.
ranges
are
483
mm,
is
with
33 ° C,
precipitation
annual
of 38°C not
extremes
the valleys
in
from 305 mm to 940 mm; in the mountains the amounts
greater.
valleys
temperature
in July, normal maximum temperatures
to
28 ° C
minimum
normal
the
approximately -1° C;
range
by mild winters and by summers in
be quite warm and nights moderately cool.
January,
In
a climate which is
have
Ranges,
characterized
days
southwestern Oregon, between
of
the
Some
average
again
occurs each winter; in the
snowfall
annual
amount ranges from 178 mm to
considerably
with
more
in the mountains
(U.S.D.A. Weather Bureau, 1930).
Climatic
southwestern
data
Oregon
for
are
selected
summarized
weather
in
stations
Table
in
The
2.
locations of these stations are indicated in Figure 2.
Site locations and characteristics
Methods.
Phacelia
Oregon
A
capitata
State
Threatened,
compilation
was
University
and
of
13 known locations for
made from herbarium collections at
(OSC), and from the Oregon Rare,
Endangered
Plant
files
housed
there.
Table 2.
Station
Name &
Elev. (m)
Gold Beach
Climatic data for selected weather stations in southwestern Oregon,
1951-80 (National Oceanic and Atmospheric Administration, 1982).
Precipitation Averages
Temperature Averages (°C)
(mm)
Daily Maximum
January July
Daily Minimum
January July
Annual
January
July
January
July
8.3
15.0
12.1
19.6
4.6
10.4
2099.8
369.3
9.3
4.4
22.0
8.3
32.4
0.4
11.5
820.7
164.1
5.3
5.0
20.1
8.9
28.9
1.1
11.3
802.9
151.1
4.1
5.1
20.2
9.1
29.0
1.1
11.4
847.1
151.4
4.8
15
Grants Pass
282
Riddle
202
Roseburg
142
26
Location
information
Management
Land
was also obtained from two Bureau of
botanists, Mr. Russ Holmes, Roseburg, and
Ms. Joan Seevers, Medford, Oregon.
Field
April
were
conducted
April-June
and
1983,
during May-July 1982,
During initial field
1984.
as many locations as possible, in both serpentine
surveys,
and
studies
non-serpentine
surveyed
areas
determine
to
were
Oregon,
southwestern
of
the
location
and
status
of
P.
22
verified
populations
of
P.
capitata populations.
Results.
capitata
sites
which
were
Five
Of
the
were
intensively
studied
(Table
3), 17
relocations of previously observed localities.
new locations for the species were found (Boomer Hill
I, Cow Creek I and II, Doe Creek, and The Drew).
Five
not
previously
located
(Table
Two
Mountain)
were
entering
private
the
cases,
system
and
Butte
(Brushy
inaccessible
due
and
Nickel
restrictions
to
on
land; the latter location is the site of
Corporation
visible
is
Rattlesnake
along
these
mine.
In
both of these
P. capitata is likely to be present, since suitable
habitat
town
of
Nickel
Hanna
sites for P. capitata were
studied during the course of this project
or
3).
reported
Creek
which
of
Peel.
is
from
surrounding
areas.
The
site is part of an extensive population
located
along the Little River near the
Individuals
of
P. capitata can be seen
many portions of the Little River Road in this area,
the
two
sites
selected
for study (Little River and
Table 3.
Site
Reported locations of Phacelia capitata in Oregon. All locations are in
Douglas County except Goolaway Gap (Jackson County) and Bridge (Coos County).
Population
number
T
R
Sec
Location
Elev.,Aspect
Slope
Years
Ownership
(m)
(CLUSTER)
A.
VERIFIED LOCATIONS
Beatty Creek
A090
30S 6W
31
0.4 km N of Cow Creek Rd
Beatty Creek RNA
320
E,SE,W
55-65
1979 Dwan & Balk
1982-4 Shelly
BLM & private
Bilger Creek
A050
29S 5W
3
BLM rd 29-5-11.0, 2.3 km
NW of Bilger Creek Rd
490
S
50
1982 Holmes
1983-4 Shelly
BLM & private
Boomer Hill I
A080
30S 6W
5
BLM rd 30-6-4.2, 8.1 km
SW of Interstate Hwy 5
on Boomer Hill Rd.
760
WSW
35
1983-4 Shelly
BLM
30S 6W
9
BLM rd 30-6-4.1, 0.5 km
from BLM rd 29-6-34.0
850
ESE
40
BLM
1978 Crowder
1984 Shelly & Holmes
Boomer Hill II
Bridge
A210
29S 11W 35
N of St Hwy 42, 3.2 km
E of Bridge
100
S
50
1951 Kruckeberg
1980 Sundberg
1982-4 Shelly
private
Callahan Creek
A180
31S 2W
9
Umpqua NF rd 3230, 0.5
km W of St Hwy 227
490
SE
75
1979 Fosback
1983-4 Shelly
private & USFS
Cow Creek I
A120
30S 7W
35
Cow Ck Rd, 0.5 km W of
Doe Ck Rd
300
WNW
65
1983-4 Shelly
BLM
Cow Creek II
A130
30S 7W
34
NW side of Cow Creek,
1.2 km SW of Doe Ck Rd
300
SE
60
1983-4 Shelly
private
Cow Creek Turn-off
A160
30S 7W
36
Cow Ck Rd, 4.8 km W of
Glenbrook Loop Rd
230
S
5
1979 Dwan & Balk
1984 Shelly
private
Doe Creek
A15A
30S 7W
22
Doe Ck Rd, 1.9 km N of
Cow Ck Rd
400
0
1984 Shelly
private
Table 3.
Site
Population
number
Continued.
T
R
Sec
Location
Elev. Aspect
Slope
Years
Ownership
(m)
(CLUSTER)
Doe/Thompson Ridge
A150
30S 7W
14
BLM rd 30-7-14.0, 3.2
km NW of Doe Ck Rd
640
ESE,SE
40
1982 Holmes
private
1982 Shelly & Holmes
The Drew
A190
31S 2W
21
Umpqua NF rd 3130E,
3.1 km from NF rd 3130
810
ESE
25
1982-4 Shelly
private
Elk Creek
A170
30S 2W
3IS 2W
34
St Hwy 227, 1.6-4.0 km
SE of Tiller
375
W,WSW
60-80
1972 Heckard
1979 Fosback
1983-4 Shelly
private, BLM,
USFS
32S 3W
31
0-0.8 km W of Goolaway
Gap
930
E,SE
20-30
1981 Seevers
1982-4 Shelly
ELM & USFS
Goolaway Gap
3
32
Lee Creek
A030
28S 4W
15
BLM rd 28-4-15.0, 0.5
km NW of Lee Ck Rd
415
S,E
0-70
1977 Fosback
1982-4 Shelly
BLM
Little River
A010
26S 3W
28
Co Rt 17, 5.9 km SE
of Glide
270
SE
35
1976 A & F Parker
1982-4 Shelly
private
Myrtle Creek North
A060
29S 5W
20
Co Rt 14, 1.6 km N of St
Hwy 99, NW of Myrtle Ck
245
SSE
70
1952 Howell
1983-4 Shelly
private
Peel
A020
26S 3W
35
BLM rd 26-3-34.2, 0.5
km E of Peel
295
S
25
1977 Knouse & Moore
1982-4 Shelly
private
32S 3W
27
SW flank of Red Mtn, 0.8
km N of Jackson-Douglas
County line
930
E,SE
0-20
1942 Detling
USFS
1984 Shelly & Nelson
Red Mountain
Rice Creek
A070
29S 7W
35
BLM rd 29-7-36.0, 1.2 km
W of Rice Ck Rd
710
ESE
20-30
1978 Crowder
private
1984 Shelly & Holmes
Salt Creek
A140
30S 7W
36
BLM rd 30-7-36.0, 0.5 km
NE of Cow Ck Rd
340
S
15
1982 Holmes
private
1984 Shelly & Nelson
Weaver Road
AO6A
29S 5W
40
Weaver Rd, 2.9 km SW of
Interstate Hwy 5
210
SE
55
1966 Ornduff
1984 Shelly
private
Table 3.
Site
Population
number
Continued.
T
R
Sec
Location
Elev. Aspect
Slope
Years
Ownership
%
(m)
(CLUSTER)
B.
UNVERIFIED LOCATIONS
Brushy Butte
Myrtle Creek South
28S 4W
18
29S? 5W
?
Ca 9 km S of Dixonville
N,NW
?
1979 Miles
private, BLM?
"2 mi S of Myrtle Creek" 180?
?
?
1934 Eastwood &
Howell
private?
SE?
1949 Detling
private
50
1982 Holmes
private
?
1983 Norris
USFS
610
Nickel. Mountain
-
30S 6W
21
Nickel Mountain, ca 6.5
km W of Riddle
Rattlesnake Creek
-
26S 3W
34
Little River Rd, 8 km
from St Hwy 138
320
Red Mt Lookout
-
32S 3W
23
Red Mt lookout, and N
along ridgetop
1460
365?
S,SW
?
30
Peel)
are
on
each
end of this series of populations.
I
was
not aware of the Red Mountain Lookout site until after
the
1984
field
Eastwood
miles
and
season.
Howell
The
(1474,
historical
CAS)
is labeled as being "2
of Myrtle Creek, Douglas County."
S.
collection by
This vicinity
was surveyed, but no plants were located.
The
entire
capitata
occurs
southern
Coos,
southwestern
the
for
southeast
of
Phace ia
and
western
Douglas,
Jackson
Counties,
in
The northernmost location
occurs along the Little River in Douglas
Cedar
this
of
range
southernmost location is near Goolaway Gap,
studied
easternmost
northwestern
and
westernmost
1956);
south-central
in
species
The
sites
geographic
Oregon (Figure 2).
County.
The
known
Springs
location
and
is
Mountain, in Jackson County.
the most isolated of the 22
is also the type locality (Rruckeberg,
site
is
near
Bridge,
in Coos County.
The
location is along Elk Creek, just southeast of
Tiller in Douglas County.
are
The
serpentine
the
northeasternmost
Oregon
(Figure
located
level
2).
on
slopes
areas,
or
outcrops
In
where
P. capitata is found
such substrates in southwestern
general,
the sites studied are
with southerly exposures; a few are in
on
slopes with west, northwest, or east
exposures.
The steepness of these sites is quite varied,
with
ranging
at
slopes
which
Bridge,
P.
where
from 0 to 80%.
The lowest elevation
capitata occurs is at the type locality near
a
portion
of the site is at approximately
31
100
The
m.
Goolaway
site,
highest
elevation
site is 930 m at
study
Gap, although the unverified Red Mountain Lookout
km northeast of Goolaway Gap, apparently occurs
7.1
at an elevation of 1460 m.
is of great interest to field inventory studies of
It
P.
capitata
response
and
that
at
18
have
the
of
been
The ecological
It is sufficient here to state that,
Mountain,
partially
least
activities
Only
railroads.
of this will be discussed
sites studied, the serpentine habitats
22
at
quarrying
construction
and
the
at
disturbed,
Beatty
of
Creek,
mainly
by
roads
and
Bridge,
Red
and Rice Creek sites are the habitats largely or
completely
undisturbed;
there
is
at the Bridge site.
livestock
in
ramifications
later section.
a
a strong favorable
shows
to disturbance at many locations.
evolutionary
in
species
the
abundance
so
some
light grazing by
Phacelia capitata increases
greatly in disturbed areas, that many of
its
populations have been "released" from any paucity that
may
have
characterized them in earlier times.
tendency
shared
is
with
most
of
This weedy
the other taxa of the
species group Magellanicae (Heckard, 1960).
the south of its range, P. capitata is replaced on
To
serpentine
soils
corymbosa.
The
in
by
the
closely
related
species
P.
latter species, in addition to occurring
the Siskiyou Mountains region of Oregon and California,
grows
in
location
the
in
north Coast Ranges of California and in one
Butte
County,
California,
in
the
Sierra
32
The
Nevada.
approach
most
Oregon,
Here,
closely
the
in
km
the
gap.
northwestern
in
two
species
Jackson
County,
these
of
area southwest of Cedar Springs Mountain.
stretch of the West Fork Evans Creek Road, west of
the
Phacelia corymbosa is found approximately 8.9 km
Nowhere
(see
These two locations
southwest, along Grave Creek.
separated
are
ranges
the Goolaway Gap site for E. capitata occurs along a
0.8
to
geographic
were
by
an
these
of non-serpentine substrates.
area
species found in biotic sympatry
two
Figure 8, page 104).
The Grave Creek location for P.
corvmbosa
represents
locations
for this species; the other is on Iron Mountain,
in
southern
Coos
one
and
the
of
northern
two northernmost known
Counties
Curry
(Baker,
1956).
Associated vegetation
The
have
spectacular influences that ultramafic substrates
on
great
plant
the
interest
taxonomic
dwarfed,
is
often
the
vegetations
some
one,
effects
unique,
words
and
of
and
are of physiognomic and
is common.
Species composition
endemism and range disjunction are
serpentine
of
ecologists
Serpentine vegetation is frequently
xerophytism
characteristic
In
These
importance.
and
community
both
to
biosystematists.
which inhabits them has been of
life
floras
Whittaker
(Kruckeberg, 1969).
(1954),
most
serpentine
throughout the world appear to be dominated by
or
some
combination,
of
three growth forms:
33
coniferous
trees,
plants."
He
Siskiyou
in
states that the stunted, open appearance of
serpentine
and
size
exclusion
broad-leaved
trees.
status
as
menziesii
Pseudotsuga
of
reduction
or
opening,
vegetation results from the reduction
numbers
such
shrubs, and "grass-like
sclerophyllous
equivalents
shrubby
to
and the
the
of
In soils of less favorable nutrient
there
serpentines,
lowering,
trend towards
a
of the plant community
reduction
and
is
toward the lower strata (Whittaker, 1954).
in
the
(1954,
In
a
associations occurring on ultramafic soils
plant
The
Siskiyou
Mountains have been studied by Whittaker
White (1971), and Atzet and Wheeler (1984).
1960),
his gradient analysis studies, Whittaker found that, on
gradient
xeric,
fairly
from
more
mesic, low elevation sites to more
higher elevation sites, the vegetation changes from
open
Chamaecyparis
Pseudotsuga
menziesii
intermediate
forest-shrub
monticola-P.
to
very
through
stands,
stands
of
monticoladistinctive
Pseudotsuga-Pinus
lambertiana-P. ieffrevi-Calocedrus decurrens,
open
decurrens.
lawsoniana-Pinus
stands
This
of
jeffreyi
P.
latter
and occasional g.
community is termed a woodland,
"temperate savannah," or pine steppe (Whittaker, 1954).
White
serpentine
(1971)
substrates
characteristics
successional
are
observed
being
status.
in
plant
five
southwestern
related
Gentle,
to
associations
Oregon,
elevation
on
their
and
rocky slopes above 1200 m
distinguished by the open character of the vegetation.
34
Important
species
sandbergii,
Ceanothus
include
Allium
species,
Poa
idahoensis,
Aspidotis
falcifolium,
cuneatus.
vegetation
Festuca
and
densa,
At middle elevations (860-980 m) the
retains
its
including
open
Ceanothus
character,
but
cuneatus,
increase
shrubby
in
abundance.
Below
860
according
to
intermediate,
Vulpia
or climax.
young,
i.e.,
status,
Within the youngest successional
species are predominant (Epilobium minutum,
microstachys,
heterophylla,
These
vegetation transects were grouped
successional
annual
areas,
the
m,
Aira
Githopsis
species
are
Agoseris
caryophyllea,
speculariodes, and Madia exiqua).
considered
to
indicators of open
be
mineral soils.
successionally
the
In
cuneatgs
becomes
perennial
intermediate areas, Ceanothus
important,
grasses
and
with
along
forbs
an
($tipa
increase in
lemmonii
and
Eriophyllum lanatum) and the fern species Aspidotis densa.
The
climax
described
Pinus
as
an
open
at
elevations
grassland
below 860 m is
savannah with scattered
ieffrevi and occasional Calocedrug decurrens.
was
notable
a
herbaceous
($tipa
and
community
stratum
lemmonii,
Festuca
Monardella
lanulosa).
absence
is
dominated
Danthonia
spp.)
villosa,
Notable
and
the
by
cuneatus.
perennial
The
grasses
californica, Roeleria nitida,
forbs
Polvgala
in
Ceanothus
of
There
(Ranunculus
californica,
climax
occidentalis,
and
situation
Achillea
was
the
35
absence
of
species
indicative
Aspidotis densa.
There was a low abundance of
mineral
open
of
due
soil,
the
to
limited amount of such habitat (White, 1971).
Atzet
and
ieffrevi
Wheeler (1984) further described the Pinus
Series
ieffreyi/grass
Association
approximately
with
Siskiyou
the
in
1390 m.
approximately
Mountains.
The
P.
defined as occurring below
is
The climax dominant is P. ieffreyi,
55%
average
of herb and grass
cover
species.
Vegetation
collected
plant
community
adjacent
center
type
occurs.
to,
Table
data
Quantitative
were
order to describe the associated species and
in
generally
see
sampling.
This
with
which
capitata
Pbacelia
study was conducted within, and
the Beatty Creek Research Natural Area (RNA;
The site is located approximately in the
3).
of the known geographical range of the species, and
the plant community is undisturbed.
Methods.
circular
The
data
macroplots.
east-facing
slope
within
RNA.
the
west-facing
Two
along
were
collected
macroplots
the
west
in five 1000 m2
were situated on an
side of Beatty Creek,
The other macroplots were located on the
slope
above
the
east
macroplot
the
side
of
the
creek,
outside the RNA.
Within
collected:
and
of
following
data
were
m2 canopy coverage of tree and shrub species,
Pinus
for
number
each
ieffreyi
and
Pseudotsuga
menziesii, the
trees greater than 3 m in height and the number
36
of
trees
and
less
The slope
than or equal to 3 m in height.
The average
aspect of each macroplot were also noted.
elevation of the area is approximately 320 m.
Within
microplots
were
each
total),
(40
positioned
angles
macroplot
at
eight
placed
were
The radii
on each of 4 radii.
2
angles to each other, and at 45°
90°
to the up-downhill axis of each macroplot.
radius,
the
microplots
0.25
On each
were placed on the uphill side, 6
and 12 m from the center of the macroplot.
Within
plants
of
each
present
microplot,
were
all
species
of
herbaceous
Approximate canopy coverage
listed.
each species was then recorded, according to the canopy
coverage classes of Daubenmire (1959):
Cover class
canopy coverage
%
Results.
trees
and
Midpoint %
2.5
15.0
37.5
62.5
85.0
97.5
0-5
5-25
25-50
50-75
1
2
3
4
5
6
75-95
95-100
sampling
The
shrubs
are
results
for the associated
given in Table 4.
The results for
the herbaceous vegetation are given in Table 5.
The
associated
in
the
scattered,
"savanna"
plant
at
tree
community
with
which
P.
capitata
is
Beatty Creek is dominated by Pinus jeffreyi
Individuals
layer.
giving
appearance.
the
community
Notable
of
the
the
latter
are
well-documented
shrubs include Ceanothus
cuneatus and Rhus, diversiloba.
Thirty-six
herbaceous species were encountered in the
37
Table 4.
Associated tree and shrub vegetation at Beatty Creek. Data
were collected in five 1000 m2 circular plots, and are
presented as percent canopy cover; for Pinus jeffreyi and
Pseudotsuga menziesii, figures in parentheses indicate the
number of stems greater than 3 m, and the number less than or
equal to 3 m, in height.
Study dates: 25-26 April 1984.
*
Plot
2
3
Slope position
low
low
low
Slope (%)
56
63
60
E
SE
W
Aspect
4
5
high
middle
50
49
WSW
SW
Trees
Pinus jeffreyi
30(18;6)
15(18;19)
Pseudotsuga
menziesii
1.6(1;0)
2.0(1;0)
7.5(10;6) 7.5(18;7) 30(30;35)
0
0
0.15(0;1)
Umbellularia
californica
0
3.5
0.25
0
0.4
Arbutus menziesii
0
1.8
2.5
0
0
Ceanothus cuneatus
0.1
1.1
0
Holodiscus discolor
1.8
0
0
Shrubs
Rhus diversiloba
0.15
0.6
0.05
0.05
0
0
*-Plots located within the Beatty Creek Research Natural Area.
0.05
0
0.15
38
Table 5.
Herbaceous vegetation at Beatty Creek. Data are averages of
percent cover (using canopy cover class midpoints), and percent frequency, in 40 0.25 m2 plots; study dates: 25-26 April
1984.
Species
Average % cover
% Frequency
Grasses and graminoids
Bromus carinatus
Luzula campestris
Melica geyeri
Poa scabrella
Vulpia microstachys
1.0
6.5
0.1
16.3
1.5
15.0
40.0
5.0
45.0
47.5
0.9
37.5
42.5
5.0
37.5
5.0
10.0
5.0
Annual forbs
Collinsia parviflora
Cryptantha intermedia
Crocidium multicaule
Galium aparine
Githopsis specularioides
Plectritis congesta
Thysanocarpus curvipes
1.1
0.1
0.9
0.1
0.6
0.1
Perennial forbs (including ferns)
Achillea millefolium
Aspidotis densa
Brodiaea pulchella
Castilleja pruinosa
Cerastium arvense
Delphinium menziesii
Dodecatheon hendersonii
Eriophyllum lanatum
Eriogonum nudum
Erythronium oregonum
Hieracium scouleri
Iris chrysophylla
Lithophragma parviflora
Lomatium nudicaule
Lupinus albifrons
Microseris sp.
Monardella villosa
Phacelia capitata
Ranunculus occidentalis
Saxifraga occidentalis
Sedum sp.
Silene hookeri
Thlaspi montanum var. montanum
Viola hallii
0.4
12.6
0.2
0.4
0.9
0.1
0.1
1.1
0.5
0.7
6.5
6.4
0.4
1.6
0.1
0.1
2.3
0.4
2.9
0.1
0.5
1.9
3.8
0.1
2.5
82.5
7.5
15.0
10.0
5.0
5.0
32.5
20.0
27.5
35.0
20.0
15.0
27.5
5.0
5.0
43.5
2.5
40.0
5.0
20.0
50.0
77.5
5.0
39
microplots.
Aspidotis
Prominent
densa,
species
Luzula
Iris
chrysophylla,
other
important
and
include
montanum
Thlaspi
Ranunculus
are
cabrella,
Hieracium scouleri,
campestris,
species
Poa
var. montanum;
occidentalis,
Monardella villosa, and Silene hookeri.
Within
serpentine
at
successional
the
observed by White (1971), the vegetation
sites
Beatty Creek appears to be between the intermediate and
climax
two
stages.
species
status,
ieffreyt
savanna
grasses
and
herbaceous
good
cuneatus and Aspidotis densa,
Ceanothus
considered
successional
a
pattern for lower elevation
are
the
intermediate
However,
well-developed
forbs
flora.
present.
are
is
of
indicative
here,
and perennial
components
major
the Pinus
of
the
Overall, the Beatty Creek community is
example
of
Pinus
a
ieffrevi/grass
savanna
association.
Many
in
communities with habitat features and vegetation fairly
similar
at
of the other 21 populations of P. capitata occur
Red
Mountain
highest
Table
the one studied above.
to
elevations
and
3),
different
from
quantitative
partial
of
lists
interest.
include
and
Pinus
where
the
the
P.
was
capitata
plant
described
were
These represent the
Gap.
associated
one
studies
of
Goolaway
Notable exceptions are
conducted
at
studied (see
communities
Beatty Creek.
are
No
at these sites, but
associated species at each location are
At the Red Mountain site, unique associates
lambertiang,
Garrya buxifolia, Amelanchier
40
pallida,
and
Silene,
Goolaway
Gap
site,
locations
campanulata ssp. glandulosa.
associated species not noted at other
Whipplea modesta, and
chrysophylla,
Castanopsis
Ceanothus
viscida,
Arctostaphylos
include
pumilus,
At the
Vicia californica.
The
Hill
and
I
II sites appeared unique as
occurring at higher elevations than Beatty Creek and
well,
in
Boomer
more
mesic
Boomer
Hill
site
I
Lathyrus
ursinus,
perforatum,
thapsus,
As
perfoliata.
appear
to
nature
than
capitata
The
formosa,
americana,
dubium.
be
species
hastata,
at the
Rubus
Bromus ;igidus, Hypericum
polvohyllus,
munitum,
Verbascum
Phacelia
include
Cvnosurus
Polystichum
Associated
situations.
echinatus,
Torilis
arvensis,
sativa,
Vicia
Lonicera
hispidula,
Collomia
beteroPhylla,
and
Montia
noted previously, the soils at this site
of
different,
a
less
stringent chemical
the more typical serpentine soils of other P.
sites.
Boomer Hill II site includes Salix sp., Aauilegia
Petasites
frigidus,
Chrysanthemum
This
site
is
Eauisetum arvense, Veronica
leucanthemum,
unusual
in
the
and
Trifolium
high amount of
seepage which occurs from the gentle slopes.
There
Pinus
were
jeffreyi
Otherwise,
the
only
was
two
Phacelia
not noted:
capitata sites where
Boomer Hill I and Bridge.
species is associated with P. ieffrevi (in
varying amounts) throughout its range.
41
Vegetation history
Due
to
topography,
gradient
from
to
factors
have
its
Klamath
of
in
mixed
and
and
summarized by
America; 2) forest
North
evergreen
communities) occur
vegetation
The
offered
is
a center of floristic diversity and
is
characteristics
summary
are
to most closely resemble Arcto-Tertiary
endemism.
these
The most striking
3) the region has an exceedingly rich flora for
latitude,
narrow
vegetation
patterns
thought
and
These
1) the area has one of the most highly
(1961):
(redwood
there;
climatic
very wide range of habitats.
a
to exist in the region.
vegetation
forests
steep
a
had a major influence on the vegetation types
come
formations
and
the coast inland, the Klamath Mountains have
characteristics
complex
geologic history, dissected
diversity,
contain
have
Whittaker
complex
edaphic
evolved
that
long,
a
history giving rise to
must have been extremely complex.
A
to aid in postulating about the
here,
evolutionary history of Phacelia capitata.
The
been
vegetation
considered
region
particular
in
According
sources
evolution
to
the
of
Detling
(1968);
the Pacific Northwest has
discussed
was
that
of the Klamath
by Whittaker (1961).
there have been three principal
former,
plant associations in the Pacific Northwest: 1)
of
in situ; 2) the northern Arcto-Tertiary Geoflora;
and
3)
the
southwestern
Geofloras
by
history
the
Madro-Tertiary
United
Geoflora of western Mexico and
States.
The movements of these
began during the Oligocene epoch, as a drying and
42
cooling
trend allowed the replacement of Eocene subtropical
forests
by
more
Metasequoia
forests
temperate
much
across
comprised
forests
the Pacific Northwest.
of
Arcto-Tertiary
the
and
Sequoia
of
These
Geoflora, and their
spread
across
During
the late Miocene, continued cooling and increasingly
dry
to
conditions affected the Arcto-Tertiary forests, leading
a
decrease
forests
A
the region continued into the Miocene epoch.
the more humid coastal strip (Whittaker, 1961).
to
general
increase
dry-adapted
(Detling,
in deciduous forms and restriction of these
herbaceous
1968).
vegetation
probable
centers
southwest,
was
Northwest;
these
Geoflora.
in
the
there
thought
the
which
of
origin
very
these
changes
spread of these latter dry-adapted
many
of
and
Pinaceae
the
of
accompanied
types
The
types,
members
in
moved
scattered
in
from
northward
areas
the
of
important to the flora of the Pacific
comprised the Madro-Tertiary
assemblages
This flora reached southwestern Oregon by early
Pliocene epoch, and reached its maximum development
by
the
that
Klamath
middle
of the epoch (Detling, 1968).
during
Pliocene epoch the vegetation of
the
Mountains
It is
took on essentially its present-day
character (Whittaker, 1961).
During
culminated
the
late
eventually
epoch.
At
retreat
occurred,
higher
Pliocene,
least
elevations
in
four
the
mixed
cooling trend
ice ages of the Pleistocene
periods
leaving
further
a
of
glacial
northern
with
more
relict
advance and
species
xeric,
at
southern
43
vegetation
Pacific
types (i.e., the Pine-oak Forest) throughout the
Northwest
xerothermic
period
Hypsithermal
northward
More
1968).
recently,
a
and warmer than the present, the
dryer
interval,
occurred
4-8,000
years
ago.
The
spread of the Madro-Tertiary Geoflora resumed and
continued.
the
(Detling,
This trend brought new genera and species over
Klamath
chaparral
Mountains
vegetation
from
northern
California,
and
became established in the Rogue River
basin (Detling, 1968).
The
events
of
last 4,000 years, under a cooler,
the
moister climatic trend, are summarized by Detling (1968).
With possible temporary unrecorded reversals the
Mesic Coniferous Forest has again become dominant
over much of the western portion of the region.
It has
largely replaced the xeric Rogue flora
except for numerous refugia where soil conditions
have probably hindered its
(the xeric flora's)
competitors.
Serpentine
areas
of
southwestern
Oregon are perhaps good
examples of such soil-influenced refugia.
The
end
results
these
of
historical
events
are
presented by Whittaker (1961).
The central relation of the Klamath Region is
regarded primarily not as one of a center of
origin for forests of other parts of the West,
but as a center toward which mesophytic forests
of the past have shrunk, and as a center of
accumulation of species of varied evolutionary
history in the diverse habitats of ancient land
surfaces.
In
summary,
of
53
considered
members
of
the
out
Pacific
it is interesting to note that 28 genera,
by
Detling
Madro-Tertiary
Northwest.
Phacelia
(1968) to be important
Geoflora,
capitata
extend into the
is
found
in
44
association
has
geographic
seems
grass
as
these
of
28 genera:
Ceanothus,
Umbellularia.
and
Rhps,
ieffreyi
It
nine
Arctostaphvlos,
Arbutus,
pinus,
with
In
Garrya,
Amelanchier,
Holodiscus,
the genus Pinus, P.
its area of northernmost distribution the
region to which Phacelia capitata is restricted.
to postulate that the Pinus ieffreyi/
reasonable
woodland,
associated,
may
communities
of
with which Phacelia capitata is frequently
be
a
good
example of one of the diverse
Madro-Tertiary derivation which entered the
Klamath Region from the south.
45
BIOSYSTEMATIC STUDIES
Cytology
Materials
microsporocytes,
collecting
These
and
methods.
and
young
inflorescences
fixing
acetic
chromosome counts from
buds were obtained in the field by
floral
several
For
inflorescences
from the plants.
immediately placed in a killing
were
solution of 3 parts 95% ethanol: 1 part glacial
which
acid,
was
mixed in the field, for 24 hours.
The
material
was
then
70%
ethanol,
and
stored in a freezer in a third change of
washed in two overnight changes of
the same.
buds
the
In
progression
therefore
carmine
stain
or
from
45%
acetic
and
the
the alcoholic hydrochloric acidby
buds
hours
Snow
(1963),
to ensure that
Inflorescences were warmed in
in
a 55-60°C oven, were then
in 70% ethanol in a freezer.
Anthers were
the buds and placed on a slide in a drop of
acid.
cells
inflorescences were
in distilled water and studied immediately,
stored
dissected
several
24-48
briefly
were
using
described
for
entire
The
buds were obtained.
stain
rinsed
May).
stained,
suitable
the helicoid cymes there is a distal
meiosis throughout the early growing season
of
April-late
(late
the
of
A cover slip was placed on the material
were squashed by applying pressure.
from
one
inflorescence
had
Often,
to be prepared
before an appropriate meiotic stage was observed.
Results.
Twenty-eight chromosome counts were made in
46
the
course
for
P.
of
corvmbosa,
hastata.
20 for Phacelia capitata, five
this study:
These
two for E. heterophvlla, and one for P.
previous
plus
results,
counts
for
E.
capitata and P. corvmbosa, are shown in Table 6.
The
base
chromosome
Magellanicae
is
counted,
capitata
P.
particular
Callahan
interest
may
reveal
all
six
At 19 of 20 sites
two
populations,
Elk
is
mixture
a
counts
from
of
now
are
Of
Creek and
diploids
The
and
the Callahan Creek site
the presence of diploids there, as well.
in
group
occurs at this chromosome level.
are
Further
species
diploids
species
the
1960).
(Heckard,
population
tetraploids.
in
which contain tetraploid individuals.
Creek,
Creek
Elk
x=11
number
Thus,
Magellanicae that occur as basic
the
known to have morphological counterparts
at the tetraploid level.
Two
in
previous tetraploid counts (n=22) for E. corvmbosa
Oregon
additional
E.
Coast
Sierra
These, plus five
counts obtained during this study, indicate that
corvmbosa
Diploids
cited by Heckard (1960).
were
occurs
of
this
Ranges
of
Nevada
primarily
species
this
level in Oregon.
are known to occur in the North
California,
(Heckard,
at
and
1960).
in
one locality in the
counts may yet
Further
reveal the existence of diploids of E. corymbosa in Oregon.
Three
Oregon,
1960).
that
all
The
this
counts
for
P.
n=11,
had
previously been recorded (Heckard,
two
heterophvlla
in
southwestern
additional counts reported here indicate
species
(as
ssp.
viraata)
is
primarily
47
Table 6.
Species
Chromosome numbers for Phacelia species group Magellanicae
in southwestern Oregon.
Chromosome
Collection
Population
number (n)
P. capitata
P. corymbosa
P. hastata
Shelly 419
Shelly & Nelson
Shelly & Holmes
Shelly & Holmes
Kruckeberg 2703
Shelly 440
Shelly 405
Shelly 411
Shelly 681
Shelly 706
Shelly & Nelson
Shelly & Nelson
Heckard 2930;
Shelly 433
Shelly & Nelson
Shelly 399
Shelly 690*
Shelly 414
Shelly 354*
Shelly & Nelson
Shelly & Nelson
Shelly 689
Beatty Creek
Bilger Creek
Boomer Hill I
Boomer Hill II
Bridge
Callahan Creek
Cow Creek I
Cow Creek II
Cow Creek Turn-off
Doe Creek
The Drew
Elk Creek
Elk Creek
11
11
11
11
11
Goolaway Gap
Lee Creek
Little River
Myrtle Creek North
Peel
Red Mountain
Salt Creek
Weaver Road
11
11
Eight
Eight
Grave
Rough
Waldo
Wimer
22
22
22
22
22
22
Shelly 700
Shelly 702
Shelly 441
Shelly 698
Shelly 697
Constance &
Bacigalupi 3394
22
Shelly & Holmes 767
11
11
Shelly 714
Shelly & Nelson 728
Dollar Mtn I
Dollar Mtn II
Creek
& Ready
Road
Boomer Hill I
P. heterophylla Cow Creek
Elk Creek
22
11
11
11
11
11
11
22
11
11
11
11
11
11
724
766
781
734*
726
738
736
725*
*-Representative voucher from the same population, the actual count
being obtained from a separate, unvouchered individual.
48
diploid
that
in
tetraploid
although
area,
elsewhere
level
it
its
in
does
occur
at
the
range, mainly as ssp.
heterophylla (Heckard, 1960).
The
the
reported
count
first
here for E. hastata is apparently
from southwestern Oregon.
one
Only tetraploids
of this widespread species have been found (Heckard, 1960).
Morphology of the species
Taximetric
The morphological characteristics
studies.
of
Phacelia
capitata
were studied, to describe the degree
of
diploid
variation
within the species and the variation
pattern
populations
the
in
comparative
purposes,
containing
tetraploids.
For
studies of P. corvmbosa, P. hastata,
and P. heterophvlla ssp. virgata were also made.
Methods.
studies,
due
material
of
corolla
to
capitata
P.
obscured
capitata
no
were
of
and
used in the numerical
representative
herbarium
because the features of the
pressed specimens.
It was hoped
could be conducted to augment the
However, survival of transplants (8/29) of
and
E.
overwatering.
capitata)
in
studies
studies.
field
plants
paucity
a
greenhouse
that
P.
are
Living
Of
the
flowered.
pre-treatment,
corymbosa
was
survivors,
poor, possibly due to
only
one
plant
(P.
Seed germination of P. capitata, with
was good (80%), but none of these plants
flowered.
A
was
total
studied.
of
530
For
individuals from natural populations
Phacelia
capitata, 420 plants from 20
49
populations
were
were examined.
omitted,
present
Creek
plants
sites.
plants)
and
measurements
population,
With
number
individuals
of
the exception of the Elk
Bridge (27 plants) populations, 20
location
each
at
small
the
to
these
at
(33
due
Boomer Hill II and Red Mountain
measured.
were
were
not
taken
because
the
plants
Goolaway
the
for
Corolla
Gap
were not flowering at the
times the site was visited.
comparison
For
studied
of
with P. capitata, 110 individuals were
representing
other species of the Maciellanicae: 80
P. corvmbosa (4 populations), 20 of P. beterophylla ssp.
virgata
population),
(1
population).
Site
and
of
10
hastata
P.
(1
locations and characteristics for these
populations are listed in Table 7.
From
each
morphological
leaf
(mm),
corolla
number
individual
data
studied,
length
(mm),
(mm),
width
of
corolla
10
blade length
characteristics were collected:
width
for
opening (mm),
calyx length (mm), stem height (cm),
of leaf lobes, type of leaf lobing, number of floral
branches below the terminal cymes, and glandulosity.
The
five leaf and flower characteristics were measured
with
a
Stem
height was measured with a ruler, from the base of the
Zeiss
10x
handlens
which contains a 10 mm scale.
flowering stem to the tip of the terminal cymes.
Qualitative
the
degree
manner:
of
assessments of the type of leaf lobing and
glandulosity
were
made
in
the following
Table 7.
Study populations of Phacelia corymbosa, P. hastata, and P. heterophylla in
southwestern Oregon.
Site
Population
number
T R
Sec
Location
County
Elev. Aspect
Slope
(m)
(CLUSTER)
P. corymbosa
0030
38S 8W
19
Siskiyou NF rd 4201 at
Illinois River
Josephine
390
SW
0-15
Eight Dollar Mtn II 0040
38S 8W
8
Siskiyou NF rd 4103, 5
km W of Selma
Josephine
450
S
10
Grave Creek
33S 4W
21
Grave Ck Rd, 3.5 km N
of Slate Ck Rd
Jackson
850
SE
5
Rough E. Ready
40S 8W
18
Rough & Ready Botanical
Wayside, 8 km S of Cave
Junction
Josephine
425
Co Rd 5560, 4.2 km E of
O'Brien
Josephine
475
Eight Dollar Mtn
I
0
Waldo
0020
40S 8W
28
Wimer Road
0010
41S 9W
4
P. hastata
H010
Siskiyou NF rd 4402
Josephine
490 SE
(Wimer Rd), 6.5 km SW
of O'Brien
Same as P. capitate Boomer Hill I population (see Table 3)
E010
30S 7W
35
N side of Cow Ck, 0.1
km W of Doe Ck Rd
Douglas
300
S
0-30
30S 2W
34
St Hwy 227, 0.8 km E of
Tiller
Douglas
350
S
10
SW
0-5
0-20
P. -heterophylla
Cow Creek
Elk Creek
51
Tyve of leaf lobing
0-leaves entire.
1-leaves pinnatifid (lobes with a broad base
at the midvein).
2-leaves pinnately compound (lobes reaching
the
midvein,
with
entire
leaflets
distinguishable).
Glandulosity
(when
viewed
dissecting microscope)
with
Zeiss
40x
0-eglandular.
1-a few scattered glandular trichomes among
the longer tapering trichomes, mainly in
and near the inflorescence.
2-densely
and
conspicuously
glandular,
especially
on
intermediate-length
trichomes in and near the inflorescence,
on the pedicels, and on calyces.
Results.
ten
The
means and standard deviations for the
morphological characteristics were calculated for each
population, and these are given in Table 8.
basic
The
Magellanicae
degrees
which
consists
of
axis
bears
flowering
stem.
consists
of
branches
bearing
1960)
plants
in
the
a tap root, modified by varying
of
rosette
a
and
gives rise to an
stem pattern fundamentally
The
one
to several helicoid cymes (Heckard,
.
characters
Heckard
The
size
and
outline
of
leaves
are
fundamental importance in the Magellanicae.
of
(1960)
emphasized
the
importance of the "climax
type" (the mature rosette leaves produced just before
formation
due
the
an axis, usually foliate (or bracteate), with
Leaves.
leaf
of
branching, and a simple or branched caudex, of
each
annual
organization
to
of
the
the
flowering
stem) in making comparisons,
large amount of variability possible within a
Table 8.
Means and standard deviations for ten morphological characteristics measured on
living plants from 26 Phacelia populations. Left figure in each population is
the mean, right figure is the standard deviation. lki20 except as noted.
Phacelia capitate
Characteristic
Beatty
Creek
Bilger
Creek
Bridge
Boomer
Hill I
(n=27)
Callahan
Creek
Cow
Creek I
Cow
Creek II
Cow Creek
Turn-off
36.9
7.0
46.2
6.2
45.2
9.2
37.7
7.0
57.9
11.4
37.7
9.7
56.3
14.1
36.5
9.0
3.4
0.6
4.1
0.8
4.1
0.8
4.7
0.9
5.4
1.1
3.5
0.6
4.5
1.0
3.4
0.8
COrolla opening (mm) 5.3
0.9
4.2
0.5
3.4
0.6
4.9
0.7
5.8
0.8
3.9
0.7
4.4
0.9
3.8
0.4
Corolla length (mm)
5.8
0.5
5.7
0.4
5.8
0.4
5.3
0.4
6.6
0.7
5.6
0.5
5.6
0.5
5.6
0.4
Calyx length (mm)
5.0
0.6
5.3
0.6
5.4
0.4
4,0
0.5
5.3
0.6
5.2
0.7
5.2
0.9
5.1
0.5
29.6
6.5
35.1
11.8
25.6
5.4
28.6
6.2
42.4
10.5
29.9
5.8
30.7
8.0
34.7
7.1
No. of leaf lobes
0
0
0
0
0
0
0.04
0.2
0
0
0
0
0.1
0.3
0
Type of lobingl
0
0
0
0
0
0
0.04 0.19
0
0
0
0
0.1
0.3
0
0
0.4
0.8
2.0
1.6
2.2
2.1
0.9
1.0
2.7
2.0
1.4
1.4
1.6
1.1
1.1
1.1
1.0
0
1.0
0
1.0
0
1.0
0
1.0
0
1.0
0
1.1
0.2
1.1
0.2
Blade length (mm)
Leaf width (mm)
Stem height (cm)
No. of floral
branches
Clandulosity2
1-Type of lobing:
0-leaves entire
1-leaves pinnatifid
2-leaves pinnately compound
2-Glandulosity:
0-eglandular
1-a few scattered glandular trichomes
among the longer tapering trichomes,
mainly in and near the inflorescence
2-densely and conspicuously glandular,
especially on intermediate-length
trichomes in and near the inflorescence,
on the pedicels, and on calyces
Table 8.
Continued.
Phacelia capitata (continued)
Characteristic
Blade length (mm)
Doe
Creek
Doe/
Thompson
Ridge
The Drew
Elk
Creek
Goolaway
Gap*
Lee
Creek
Little
River
Myrtle
Creek
North
(n=33)
Peel
44.9
10.6
50.2
9.3
47.4
12.4
60.7
15.0
33.4
9.9
40.4
7.9
40.6
8.7
48.0
12.3
38.2
10.4
3.8
0.7
4.4
0.8
4.2
0.9
6.8
2.1
3.3
0.8
3.7
0.6
3.9
0.7
4.5
1.3
4.0
1.0
Corolla opening (mm) 4.5
0.6
5.3
1.0
4.9
0.7
7.0
0.8
-
-
4.1
0.7
4.6
0.5
3.9
0.4
4.7
0.8
Corolla length (mm)
6.1
0.4
6.6
0.6
6.1
0.4
6.8
0.5
-
-
5.7
0.4
6.1
0.6
5.5
0.4
6.2
0.6
Calyx length (mm)
6.0
0.7
6.1
0.6
6.0
0.7
5.6
0.7
4.7
0.6
4.8
0.6
5.0
0.5
5.2
0.6
4.8
0.5
33.8
9.6
34.2
6.2
32.2
8.4
39.5
9.3
18.4
5.8
26.3
5.3
31.2
6.1
38.2
11.4
32.1
5.0
No. of leaf lobes
0.5
1.1
0.1
0.5
0.1
0.5
0.2
0.8
0
0
0
0
0
0
0
0
0.2
0.9
Type of lobing
0.3
0.6
0.1
0.2
0.1
0.2
0.2
0.6
0
0
0
0
0
0
0
0
0.1
0.5
No. of floral
branches
Glandulosity
3.2
2.5
3.2
2.5
1.8
1.4
3.0
1.9
1.1
1.7
1.7
2.4
1.3
1.0
3.3
4.5
1.0
0.7
0.9
0.3
1.0
0
1.0
0
1.0
0
1.0
0
1.0
0
1.0
0
1.0
0
1.0
0
Leaf width (mm)
Stem height (cm)
*-Data are pre-flowering.
Table 8.
Characteristic
Blade length (mm)
Continued.
Phacelia capitata (continued)
P. corymbosa
Rice
Creek
Eight
Dollar
Mtn. I
Salt
Creek
Weaver
Road
P. hastata
Eight
Dollar
Mtn. II
Waldo
8.7
47.8
7.4
44.1
14.2
37.8
6.3
36.1
5.8
29.8 5.6
3.5
0.6
3.9
0.8
4.1
1.1
10.0
2.1
8.9
1.2
7.9 1.7
10.0
Corolla opening (mm) 5.0
0.8
4.0
0.7
4.1
0.7
7.3
1.0
7.1
1.4
7.0 1.3
Corolla length (mm)
6.4
0.5
5.4
0.6
6.0
0.5
7.3
0.6
6.8
0.6
Calyx length (mm)
5.4
0.6
4.9
0.9
5.4
0.9
5.1
0.6
4.8
26.8
6.0
33.5
8.5
29.3
6.2
18.1
3.4
No. of leaf lobes
0
0
0
0
0.2
0.6
1.1
Type of lobing
0
0
0
0
0.1
0.3
1.3
1.7
2.6
3.1
1.9
1.0
0
1.0
0
1.0
Stem height (cm)
No. of floral
branche
Glandulosity
Cow Creek
(n=10)
35.5
Leaf width (mm)
Boomer
Hill I
Wiener
Road
P. heterophylla
ssp. virgata
36.2 12.0
58.8
10.2
59.6
11.4
2.7
9.8
2.2
12.0
2.8
6.8
0.8
7.2
0.7
5.2
0.8
7.6 0.8
7.1
0.9
6.8
0.5
5.4
0.4
0.7
5.5 0.5
5.1
0.7
6.2
0.8
4.5
0.5
14.3
4.2
9.9 2.5
17.7
8.9
32.5
7.9
50.9 12.0
1.6
3.0
1.7
1.6 1.4
1.7
2.1
0.3
1.0
4.1
1.7
0.7
0.9
1.7
0.7
1.2 1.0
0.9
0.9
0.2
0.6
1.8
0.4
1.7
1.8
0.4
1.5
0.5
1.0 0.5
1.6
0.5
3.9
2.5
17.4
6.9
0
2.0
0
2.0
0
2.0
2.0
0.2
1.0
0
2.0
0
0
55
single
plant.
living
plants these climax leaves were frequently withered
anthesis.
at
living
For this reason, measurements were made on
from sterile rosettes, which appeared to be
leaves
representative
as
method
was found in P. capitata that even on
It
provides
It is believed that this
possible.
as
an adequate comparison of the distinctive
leaves of P. capitata with those of related species.
The
sterile rosette leaves of diploid E. capitata are
generally
were
entire, although in six populations lobed leaves
observed.
lobed
In these six cases combined, the number of
leaves,
the
number
populations
two
On these 12
leaves was only 12 out of 147 observed.
sterile
such
rosette
leaves
(Peel,
Doe
leaves
were
compound
pinnately
Creek),
Thus, lobing of
observed.
infrequent
of
is
lobes varied from one to four; in
of
occurrence
in diploid P.
capitata.
It
should
populations
populations
whether
or
from
the
of
leaf
of the 18 diploid
studied,
capitata
E.
13
and
in
both
containing tetraploids, it could be determined
climax
leaves of fertile rosettes were lobed
lobing
populations,
total.
However,
in
Percent of
the 13 diploid populations ranged
in
(Bridge) to 85% (Boomer Hill I); in nine of these
0%
degree
that
These results are shown in Table 9.
not.
climax
13
the
noted
be
of
lobed
Thus,
lobing
climax
the
climax
compared
leaf
leaves were 50% or less of
climax
lobing
leaves do show a greater
sterile
to
is
rosette
sporadic
leaves.
in occurrence
56
Table 9.
Climax leaf lobing in Phacelia capitata.
Population
Number of stems observed
with lobed climax leaves
Beatty Creek
6/20
30
Bilger Creek
4/20
20
14/17
82
0/27
0
13/20
65
Cow Creek II
7/20
35
Cow Creek Turn-off
2/20
10
14/20
70
Doe/Thompson Ridge
6/16
38
The Drew
7/20
35
Elk Creek
18/33
55
Goolaway Gap
13/20
65
Rice Creek
10/20
50
Salt Creek
7/20
35
11/20
55
Boomer Hill I
Bridge
Callahan Creek
Doe Creek
Weaver Road
57
since
and,
these
leaves
withered
frequently
are
at
anthesis, is of limited use in comparative studies.
The
degree
leaf
of
lobing
is
greater
in both E.
corymbosa
and
E. heterovhylla than in P. capitata.
corvmbosa
the
number
six,
with entire leaves being infrequent.
of
but
lobing
ranges
sometimes
heterophylla,
eight;
of
leaf
(Heckard,
1960). In E.
number of leaf lobes ranged from one to
the
the
single
a
The nature
pinnatifid to pinnately compound,
from
within
lobes ranged from zero to
leaf
of
In P.
individuals observed, 16 had pinnately
20
compound leaves.
Heckard
entire
being
The
lobes.
leaves
may
leucophylla
E.
hastata
describes the leaves of E. hastata as
(1960)
or
occasionally
tendency
have
with
one
for this species to possess entire
led to some of the early confusion of E.
(=E. hastata) with E. capitata.
are
to two pairs of
The leaves of
usually longer and wider than those of E.
capitata.
Leaf
blade;
width
terminal
diploid
and
lobed
in
was
measured at the widest point of each
leaves,
lobe was measured.
E.
the
the
widest point on the large,
The narrow width of leaves in
capitata is distinctive.
predominantly
This characteristic,
simple blades of sterile rosettes,
are major distinguishing features of P. capitata.
Blade
lowermost
the
length
lobes
(as
measured
from
the
base
of
the
in lobed leaves, and from the junction of
blade with the petiole in entire leaves) appears quite
58
variable
useful
diploid
in
to
It is taxonomically
capitata.
P.
some extent, but not to the same degree as leaf
outline and width.
Throughout
Flowers.
biserial
are
circinate
and
crowded
peduncle,
Maaellanicae, the flowers
the
along
forming
the
upper
helicoid
a
cyme
side
of
a
(Heckard,
1960).
Corolla
in
is significant for species comparisons
Magellanicae.
the
distinct
P.
shape
a broad, open-campanulate corolla (i.e.,
types:
egena);
margins
of
extremes in shape are of two
The
and
a
the
lobes
narrowly
(i.e.,
P. imbricata).
types,
though
not
cylindrical corolla with the
incurved and embracing the stamens
Between these extremes, the corolla
sharply
defined, are stable enough to
delimit certain taxa (Heckard, 1960).
this
In
(measured
length
of
the
lobes
incurved.
spreading
face
of
the
corolla opening
expanded flowers) and the
of
corolla (measured from the base to the tip
used
capitata,
P.
width
the
the
were
lobe)
diploid
the
across
of
a
study,
being
to
quantify
corolla
shape.
In
corollas are mostly cylindric,
the
neither
nor
spreading-campanulate
Rarely, a few individuals showed some degree of
in
the
lobes
(Boomer
Hill I, Doe Creek, Doe/
Thompson Ridge).
The
correlated
Creek
size
of
with
populations
the
corolla
ploidy level.
opening
appears
to
be
The Elk Creek and Callahan
of P. capitata, containing tetraploids,
59
have
the
highest
largest
mean
openings
on
average,
of
The corolla
corymbosa and P. Jastata also are larger,
P.
populations
The
values for diploid populations.
of
the
average corolla openings, exceeding the
than
those of diploid P. capitata.
The
the former two species are all tetraploid.
corolla openings of diploid E. heterophvlla are within
the range of diploid P. capitata, however.
Differences
width
for
in corolla length show the same trends as
the corolla opening.
of
tetraploid
the
Again, there is a tendency
populations
have longer corollas
to
than diploids.
The
result
increased
of
the
individuals
increased
effects,
of
cell
corolla
"gigas"
effect (Stebbins, 1971), in which
higher
and
of tetraploids may be a
size
ploidy
organ size.
levels show a tendency for
Because of such apparent
corolla shape should be considered carefully when
comparing these species.
The
were
the
at
calyx
length
full
anthesis.
species
was also measured in flowers which
The range of variation among all
and populations overlapped considerably; this
characteristic
thus
is
probably
of
little
use
taxonomically.
Stem
height.
Magellanicae,
species
dm)
show
overlap
as
In considering height of stems in the
Heckard
P.
(1960)
noted
that,
while
such
corymbosa (1-3 dm) and E. nemoralis (10-15
striking differences, the ranges of most species
considerably,
"...limiting
the utility of height
60
as
taxonomic
a
general,
He also noted that, "...in
criterion."
there
was
difference in height
significant
no
between diploid and tetraploid forms of the same species."
The
results
difference
consistent
E.
capitata
(9.9-18.1
ieterophylla
plants
in mean stem height between diploid
(25.6-38.2
cm).
The
studied,
cm)
tall
also
is
study indicate that there is a
this
of
tetraploid E. corvmbosa
and
nature
similar
are
heights
biennial
E.
In the few P. hastata
distinctive.
stem
the
of
to
those
encountered in diploid P. capitata.
There
is
populations
from
a
tendency for P. capitata plants from the
containing tetraploids to be larger than those
diploid populations, but individual observations from
these two groups do overlap.
Floral
majority
could
The
considered
be
(1960)
stated that the
Notable
paniculate.
exceptions
those species having virgate inflorescences (i.e.,
heterophylla
floral
Heckard
of inflorescence patterns within the Magellanicae
include
E.
branching.
virgata),
ssp.
in
which numerous short
branches overlap to form a densely branched system.
term
condition
"capitate"
in
which
is
defined
several
Heckard
by
congested
as "...the
cymes are more or
less separate from other cymes and form a head."
his
In
states
that
scorpioid
that
is
description of P. capitata, Kruckeberg (1956)
the
cymes
usually
inflorescence
consists
of two to three
in a congested, subcapitate erect cluster
borne
singly
and
terminally.
In this
61
study,
number
the
cymes
was
counted,
feature.
The
branching
is
variable
allows
from
variability
results
(Table
10)
indicate
terminal
in
highly
Even if one
that
many
of
the
capitata show branching
P.
Exceptional individuals
capitulum.
populations
diploid
apparent
still
observed
stems
is
"capitate" terminology used by
the
is
such
a single cyme below the terminal
of
fit
populations.
of this
that
and
capitata
P.
in
between
and
it
the
plants with 13 and 17
included
Creek North), 14 (Salt Creek), 10 (Lee Creek), and
(Myrtle
9
the
to
Kruckeberg,
below
assess
presence
inflorescence
flowering
to
frequent
within
the
floral branches below the terminal
of
Hill
(Boomer
Doe/Thompson Ridge) cyme branches
and
I
below the terminal cluster.
the
Of
zero,
one,
stems
had
stems
80
two
or
two
studied
branched
the
virgate
to
corvmbosa studied, all had
below the terminal cluster; 38
cymes.
Thus,
the tetraploid
for
is
not
same extent as diploid P. capitata.
The
inflorescence
distinctive.
P.
cymes
such
populations
of
in
of
Further
Oregon,
this
species
P. heterophvlla ssp. virciata is
studies
of
P.
hastata
would be
necessary to fully assess cyme branching in that species.
is interesting to speculate on the extent to which
It
the
variable,
branching
habitat
whose
in
sometimes
P.
high,
degree
of
inflorescence
capitata may be a phenotypic response to
disturbance.
Of
the
three populations studied
habitats are largely undisturbed (Rice Creek, Beatty
62
Table 10.
Floral branching below the terminal cymes in Phacelia
capitata.
Population
Occurrence of floral
brinches below the
terminal cymes
Occurrence of more
than one floral branch
below the terminal cymes
Number
Number
Beatty Creek
6/20
30
1/20
5
Bilger Creek
17/20
85
12/20
60
Boomer Hill I
16/20
80
14/20
70
Bridge
15/27
56
6/27
22
Callahan Creek
19/20
95
12/20
60
15/20
75
7/20
35
Cow Creek II
17/20
85
10/20
50
Cow Creek Turn-off
12/20
60
7/20
35
Cow Creek I
Doe Creek
18/20
90
14/20
70
Doe/Thompson Ridge
20/20
100
14/20
70
The Drew
16/20
80
10/20
50
Elk Creek
32/33
97
28/33
85
Goolaway Gap*
11/20
55
4/20
20
Lee Creek
16/20
80
5/20
25
Little River
15/20
75
9/20
45
Myrtle Creek North
17/20
85
9/20
45
Peel
15/20
75
5/20
25
Salt Creek
19/20
95
11/20
55
Rice Creek
12/20
60
5/20
25
Weaver Road
15/20
75
11/20
55
*-Data from pre - flowering individuals.
63
Creek,
Bridge), the latter two contained the smallest
and
proportions
below
terminal
the
diploid
individuals with two or more cyme branches
of
The majority of other stems of
ones.
P. capitata came from highly disturbed areas, such
Further studies would
exposed roadbanks and rock pits.
as
needed
be
determine
to
relationship
and
define
to
nature
the
possible
this
of
what factors, e.g., improved
soil texture, might be influencing it.
Vestiture
the
two
common
tapering
wide
trichome and the glandular trichome.
trichome
may be erect or appressed.
The
It exhibits
range of sizes, the extremes of which may give the
appearance
large
Heckard (1960) described
basic types of trichomes in the Magellanicae, the
tapering
a
and glandulosity.
of
two
trichomes
smaller
distinct
reaching
trichomes
types of tapering trichomes:
lengths of four to five mm, and
which
often
form
velutinous
a
undersurface beneath the large ones.
The
part
of
They
are
septate
the
because
when present, are
trichomes,
covering
shorter trichomes.
of
most prevalent on upper parts of the stem, calyx
leaves.
the
velutinous
pedicels,
lobes,
glandular
The
peduncles,
glandular
and
petioles,
trichomes
are
the veins of
often
obscured
of their small size and relative infrequency among
tapering
small,
observable
on
trichomes.
They
are
readily
the upper portions of the stem, etc., using
a 40x dissecting microscope.
Kruckeberg
(1956)
stressed
the
eglandulosity of P.
64
capitata.
that
The
species
the
throughout
individuals
the
visible
stem
one
individual (Doe
Cow Creek Turn-off) appeared
II,
glandular.
It
is
important to emphasize
trichomes
of
P.
microscopically.
the
Only
discernible glandular trichomes; two
Creek
glandular
near
study indicate, however,
does possess short, glandular trichomes
no
(Cow
conspicuously
my
of
populations.
its
possessed
Creek)
that
results
capitata
are only
are most abundant on the
They
inflorescence, on the pedicels, and on the
calyx lobes; they are less frequent on the leaves.
Phacelia
capitata
the
by
corymbosa is readily distinguishable from P.
its
short- to medium-length trichomes are all gland-tipped
SEM
(see
section).
glandulosity
hastata shows patterns of
Phacelia
similar
heterophvlla
P.
conspicuous dense glandulosity, in which
to
Phacelia
capitata.
E.
was seen to be quite glandular, and resembled
corymbosa in this respect, at least on the stems in the
inflorescences.
The
vestiture
leaf
of E. capitata is illustrated in
the SEM section.
Species
Phacelia
capitata,
(1956),
is
cespitose
stems
erect,
presented.
arising
sterile
unbranched
A
modified
perennials
each
numerous
description.
with
from
leaf
description
revised
from
that
Plants
are
1-many
thin,
of
of
Kruckeberg
deeply
taprooted
wiry
flowering
a rosulate tuft of leaves, with
rosettes
often present; stems are
or occasionally branched, (11)20-45(63)
65
cm
finely
tall,
consisting
sericeous,
herbage
silvery-gray
the
a hyaline-shiny pubescence made up of three
of
trichome
types:
longer,
shorter,
matted
trichomes, and with scattered, very short
glandular
trichomes, especially near the inflorescence, on
pedicels,
the
the
on
basal
calyx lobes, and leaf veins, rare or absent
lower
pairs
are
basal
of
long;
climax
basal
lobe
helicoid
or
borne
more
pedicels
lobes
in
are
0.5-0.8
surface
singly
2-3
capitate or subcapitate
branches below the terminal cymes;
fruit
are
with
1-3 mm long, hispid; calyx
mm
(3.0)4.2-6.6(7.7)
long-hispid
corollas
long,
margins, the abaxial
are
white,
rotate,
mm
long,
(2.3)3.0-6.0(8.4)
mm
broad,
entire, obtuse-rounded lobes; appendages are attached
barely
the
of
or occasionally slightly campanulate-spreading,
(4.1)5.0-6.9(8.0)
with
consist
or terminally, or frequently with 2
cyme
early
inflorescences
congested,
a
short-hirsute;
cylindric,
long,
cauline leaves are gradually but not
in
wide,
cm
to petioles 1-2 cm
tapering
linear-oblong,
mm
with 1-2
(1.7)2.5-6.5(8.8)
wide,
upwards;
lateral
compound
are entire, or occasionally have 1-2
pairs;
cymes
cluster
mm
leaves
reduced
pinnately
or
lobes,
(2.3)2.9-6.0(13.0)
wholly
linear-lanceolate, simple and entire,
pinnatifid
rarely
sterile leaves of the
portion of the stem;
rosettes
very
bristles underlain by
appressed
a
free
distally;
millimeter
portions
stamens
above
forming
and
style
the base of the corolla tube,
a
long (3 mm) and narrow "V"
are
exserted
5-7
mm, the
66
filaments
glabrous
mid-length
along
ovoid,
mm long, densely clothed with stout bristles 2
mm
2-3
the
mature
long;
specimens),
with
or
few scattered hairs about
a
immature
filament;
seeds
are
are
capsules
mm long (in diploid
1.9-3.0
brown, with a reticulate-pitted surface; n=11,
22.
Cluster
analysis.
taximetric
studies,
morphological
were
cluster
to
CLUSTER.
capitata
mean
the
values
for
the
10
attributes measured for 25 study populations
subjected
program
summarize the results of the
To
was
analysis
using
the computer
The Goolaway Gap population of Phacelia
omitted
the analysis because data for
from
this site were collected prior to flowering.
The
Methods.
agglomerative,
dendrograms
(Keniston,
many
different
This
introduces
options
i.e.,
until
Nonetheless,
degrees
of
fusion
analysis,
of
1978).
the
strategies
subjectivity
one
nature;
in
thus,
process
are
CLUSTER is interactive, as
exist
data
hierarchical,
is
clustering
the
possibilities
standardization
CLUSTER
combinatorial
and
summarizing
obtainable
well;
program
set,
and
into
for
and
transformation
or
for selection of
dissimilarity measures.
the
process
of data
must actually choose between various
the most satisfactory results are obtained.
CLUSTER
allows
for
a useful summary of the
difference between the species and populations
studied.
In
this analysis, the Bray-Curtis dissimilarity index
67
been
has
utilized.
attributes
the
with
measure,
relatively
whereas
values
largely
determine
because
four
for
(Boesch,
leaf
the
number
and
this
of
attributes
index
glandulosity)
low
is that
of
scores
Thus,
in
are
this
stem height would
and
Due to this, and
results.
(type
cyme
of
with
1977).
length
cluster
attributes
lobes,
cluster,
feature
high scores largely determine the value of
unimportant
instance,
leaf
A
leaf lobing, number of
branches
below the terminal
were assessed in a qualitative
or
non-metric way, the data require standardization (Dr. C.
D.
McIntire, pers. comm.).
was
division
for
a
the
by the attribute maximum, in which all values
given
maximum
morphological
value
"high
The standardization method used
observed
score
attribute
for
are
divided by the
This removes
that attribute.
bias," and allows use of quantitative and
qualitative values together.
The
fusion
individual
thus
as
the
group
result,
mean
In the
strategy, inter-group resemblance is defined
of
resemblances
all
members
to
of
another.
between members of one
Thus,
group
average
has no tendency toward "space distortion."
it
introduces
relationships
measure
the way in which group/
final clusters which are formed.
the
average
clustering
defines
and group/group resemblances are calculated, and
defines
group
strategy
produces
only moderately sharp clustering, but
relatively
originally
(Boesch,
As a
1977).
little
expressed
distortion
by
the
to
the
dissimilarity
The group average fusion strategy
68
best
summarizes
exist between the
which
differences
the
Phacelia species and populations studied.
The
Results.
shown
results
in Figure 3.
seen
at
cluster analysis are
the
of
The four major clusters of interest are
They
a dissimilarity level of approximately 0.12.
include,
from
left
populations),
tetraploids
2)
two
and
P.
1) diploid P. capitata (17
right,
P.
populations containing
capitata
hastata
populations),
(four
corymbosa.
to
population),
(one
3)
E.
and 4) P. Jieterophvlla ssp.
virciata (one population).
The
mean
results
values,
variation
at least on the basis of
that,
populations
diploid
the
uniform
fairly
indicate
morphologically.
within
the
species,
of P. capitata are
The
actual
degree
of
especially with respect to
size and outline, and floral branching, were discussed
leaf
above.
similarity
The
containing
tetraploid
population
is
these
are
the
Creek
and
Callahan
to
capitata
interest,
of
intermediate
these
actual
E.
between
intergrading
Creek).
but
and
the
the
populations
E. hastata
it does not "prove" that
taxa at the former sites (Elk
The similar tendencies toward
entire leaves, larger corolla openings and lengths,
longer,
and
expressed
three
glandulosity have most likely contributed
populations being clustered together.
The
source of morphological variation in the E. capitata
populations
containing
tetraploids,
is considered in the final section.
which is problematic,
69
Figure 3.
Cluster
species.
analysis
of
Phacelia
010-Little River; 020 (phacelia capitata=A:
Peel;
030-Lee Creek;
050-Bilger Creek; 060 Myrtle Creek North; 06A-Weaver Road; 070-Rice
Creek;
080-Boomer Hill I; 090-Beatty Creek;
120-Cow Creek I; 130-Cow Creek II; 140-Salt
Creek;
150-Doe/Thompson Ridge; 15A-Doe Creek;
160-Cow Creek Turn-off; 170-Elk Creek; 180 Callahan Creek; 190-The Drew; 210-Bridge; E.
corymbosa=0:
Road; 020-Waldo; 030 010 -Wiener
Eight Dollar Mtn. I; 040-Eight Dollar Mtn. II;
E.
hastata=H:
010-same site as Boomer Hill I
above; P. heterophvlla ssp. viruata=E:
010-Cow
Creek.
See Tables 3 and 7 for population
locations.).
r-{
0
OL
r
1-4
zo
orn
x 1-1 01-
mDD
-0-1
2>
73-0)0
m3)-0
z
(r)
-1-103>
z
1-41_,
0
cn cul
rsm. im .2
1-1 1-4
rri C) X
rn-irrip
71
Z
3> CO
2>
o
<73--1
0 C rn
C-<71U1
OD-1C
7:173-1 r
ocoDo
L,J
F-6
°C)
D
___
E010
A180
0010
0030
0020
0040
H010
A.170
A15A
A190 11.
A080
A130
A210
A150
AOGA
AOGO
A020
A070
A090
A120
A160
A030
A050
A140
A010
N
-4
LT1
01
.1 CO
DISSIMILARITY
71
Cluster
the
major
analysis
appears to be useful in illustrating
differences
Magellanicae,
corymbosa,
which
exist
the species group
in
at least when well-defined entities (i.e., P.
heterophvlla
P.
virgata,
ssp.
and diploid E.
capitata) are compared.
The
major
capitata,
morphological
corymbosa,
P.
differences between Phacelia
Jastata,
P.
and E. heterophvlla
ssp. virgata are summarized in Table 11.
Scanning
Phacelia
leaf
E.
electron
capitata
vestiture
capitata
electron
Seeds of
Methods.
E. corvmbosa, pollen morphology and
and
of E. capitata, and upper stem vestiture of
and
corymbosa were studied using scanning
E.
microphotographs.
specimens.
taken
microscopy.
All
pollen,
The seeds used were dry, mature
stem
segments,
and leaves were
from herbarium specimens and hydrated; they were then
critical-point
dried
Al.
All
(1968).
studs
following
the procedure of Cohen, et
specimens were then mounted on aluminum
using Duco adhesive (Microstick adhesive for pollen),
coated
with
approximately
gold-paladium
alloy
photographed.
University
in
angstroms
180
of
60-40
a vacuum evaporator, examined, and
The photography was done by the Oregon State
Scanning Electron Micrography Lab, Department of
Botany and Plant Pathology.
Results.
is
similar
(Figure
5A);
Magellanicae
The reticulately ridged and pitted seed coat
in
P.
this
capitata
(Figure
4A) and E. corymbosa
feature is shared by all species of the
(Heckard,
1960).
Of particular interest is
Table
Species
capitata
n
11,22
II.
Comparison of Phacelia capitata, P. corymbosa, P. hastata, and P. heterophylla ssp. virgata in
southwestern Oregon.
Habit
Stem
height (cm)
perennial,
mat-forming
20-45
7.5-25
corymbosa
22
perennial,
mat-forming
hastata
22
perennial, with
1-many stems,
rarely mat-forming
20-50
biennial
30-75
heterophylla
"P. virgata
11
Leaf outline
Leaf size
length(cm) width(mm)
Inflorescence
Corolla
shape
Clandulosity
linear-lanceolate,
2.5-6.5
entire; rarely, with
1-2 pairs of leaflets
at base of lamina
2.9-6.0
Cymes often terminal,
or sub-terminal,or
with I-many lateral
inflorescences below
the terminal cymes
linear to lanceoblong, entire, or
often with 1-2 pairs
of leaflets at base
of lamina
2.4-5.2
6.0-14.0
cymes terminal, or
cylindricoften with 1-2 subcampanulate
terminal cyme branches
conspicuosly glandular overall,
with short-medium glandular
trichomes
lanceolate, entire,
or rarely with 1-3
leaflets
3.5-7.5
7.0-13.0
cymes terminal or
sub-terminal, often
with 1-many lateral
inflorescences below
the terminal cymes
cylindric
scattered, short glandular
trichomes, especially near
the inflorescences, on the
pedicels, calyx lobes, and
leaf veins
pinnately dissected,
with 1-4 pairs of
lobes
3.8-8.0
virgate, with 10many overlapping
lateral cymes
cylindricslightly
flaring
8.0-17.0
cylindric
(campanulate
in tetraploida)
campanu late
scattered, very short
glandular trichomes, especially
near the inflorescences, on the
pedicels, calyx lobes, and leaf
veins
medium-length glandular
trichomes frequent in the
upper portions of the
inflorescence
73
Figure 4.
Seed of Phacelia capitata.
A Dorsal surface (50X)
B
Portion of dorsal surface (150X)
C
Dorsal surface pit detail (340X)
D
Detail of a double "pore" complex (700X)
74
75
Figure 5.
Seed of Phacelia corymbosa.
A
Dorsal surface (40X)
B
Portion of dorsal surface (160X)
C
Dorsal surface pit detail (280X)
D
Detail of "pore" (1600X)
Figure 5.
77
the
presence,
distributed
junctions
or
and
more
coat.
the
of
they
not
the
in
previously
of
genus
Phacelia,
documented.
concerning
supplying
seed
of
or
study
for
improved
shows
cell-wall
(Figure 5D) channels through the seed
been
include
regulation
These
presence of such seed "pores" in other species
possibilities
"pores"
hardened
by
the
with
may provide seemingly open (Figure 4D)
Mactellanicae,
apparently
bordered
be
to
constricted
The
associated
("pores")
pit margins (Figures 4B-D, 5B-D).
the
appear
material,
species, of a set of distinctively
structures
of
"pores"
both
in
has
Intriguing
the function of these
oxygen
to
the
embryos,
dormancy (i.e., in P. corvmbosa, which
germination after cold stratification
seed
(Quick, 1947)), and/or facilitation of water imbibition.
Another
prominent
circular
This
is
this
area
gone,
a
second
do
(Figures
is
material
(Figures 5C, D).
frequently cracked, and when one layer
deeper
lining is revealed in the pit bottoms;
surface
often
has
a
small
crack
near its
"cracking membrane" system may also provide
study
in
determining
seed germination ecology.
to
appear
4B,
morphology
is
interesting
not
coat
In the bottom of each pit is a
thin-looking
of
This
relationship
seed
of
P. corvmbosa.
in
"membrane"
center.
an
peculiarity
to
C),
be
its
possible
These membranes
as clearly developed in E. capitata
seeds
of
which
germinated well in the
greenhouse with no cold stratification treatment.
An
SEM
study
of
pollen
morphology
in
the
78
Hydrophyllaceae,
species
two
The
imbricata.
described
uniformly
baculate;
grains
extending
pole,
the
equally
pseudocolpi
pollen
these
of
exine
the
foveo-reticulate;
colpi
The
pollen
of
nearly
capitata
P.
but
toward
and almost
are isopolar, the
grains
the
colpi
the
finely
is
are
species
tricolpate-tripseudocolpate,
as
conspicuously
P. heterophylla and P.
Magellanicae,
the
in
and Chuang (1982), included
Constance
by
reaching either
not
long as the true colpi.
as
fits this description closely
(Figure 6A, B).
A
surface
illustrates
and
of
of the long, tapering trichomes,
trichomes
shorter
undersurface
the leaf vestiture in E. capitata
of
presence
the
the
presence
view
(Figure
form
velutinous
a
reveals
Magnification
6C).
glandular
small
which
the
hairs, in this case near the
leaf margin, in the species (Figure 6D).
The
P.
conspicuous,
corvmbosa
(Figure
distinguishing
E.
capitata
possess
very
medium-length glandular trichomes of
from
it
is
not
small
7C)
are
very
important
P. capitata (Figure 7A).
in
However,
eglandular (Figure 7B), since it does
glandular
trichomes
similar to those
which are found in P. corvmbosa (Figure 7D).
Reproductive bioloav
Flowering
and
Phacelia
capitata
elevation
sites
pollination
begins
in
Flowering
biology.
early
May
at
the
in
lower-
and continues until early to mid-June.
At
79
Figure 6.
Pollen and leaf vestiture of Phacelia capitata.
A Pollen grain (4500X)
B
Detail of pollen grain colpus (18000X)
C
Leaf vestiture (30X)
D
Detail of vestiture at leaf margin (300X)
80
81
Figure 7.
Stem vestiture of Phacelia, capitata and P.
corymbosa; stem segments taken from just below
the terminal cyme branches.
A
Phacelia capitata stem vestiture (150X)
B
Short glandular trichome of E. capitata
(600X)
C
Phacelia corymbosa stem vestiture (40X)
D
Short glandular trichome of P. corymbosa
(500X)
Figure 7.
83
higher-elevation
the
until
May
late
first
flowers
the
lower
portions
that
the
flowering is delayed
initial
early June and lasts until early July.
or
The
noted
sites,
open within an inflorescence are on
to
the cyme branches, although it was
of
lowermost buds are not always the first to
bloom.
As flowering proceeds, the initially helicoid cyme
branches
uncoil until, at the time of seed maturation, they
are nearly or completely ascending.
flowers
In
the
stamens
5-7
mm
with
of
and
beyond
the
capitata which are fully expanded,
P.
are well exserted, extending from
styles
the corolla tube.
ends
style
the
of
The anthers are subequal
branches.
The
stigmatic
surfaces
are
very small, the stigmas being confined to the
tips
the
style
of
branches and consisting of the ends of
only a few cells (Heckard, 1960).
The
of
quite
capitata are congested, forming a
P.
many-flowered
tight,
be
flowers
inflorescence
attractive
which, as a whole, may
pollinators.
to
In cases where the
in an inflorescence were counted, the number ranges
flowers
53-303; these inflorescences varied from approximately
from
2.5 to 5.0 cm in diameter.
Throughout
protandrous,
stigma
of
the
MaQellanicae,
flowers
are
the pollen being ripe and available before the
is receptive (Heckard, 1960).
dichogamy,
the
or
temporal
This is one mechanism
separation of sex functions in
flowers,
whereby
Protandry
is only effective, however, if it is total, i.e.,
self-pollination
may
be
inhibited.
84
if
all
of
stigma
is
Another
receptive
as
situation
seen
protracted
and
may
likelihood
be
(Faegri
may
which
factor
protandry
ages
pollen is removed from a flower before the
the
Pijl,
1971).
effectiveness of
self-pollination
is
the
the Magellanicae, in which flowering is
in
many adjacent flowers of slightly differing
open at one time on a single individual.
of
having
ripe
simultaneously
within
one
single
within
of
der
the
decrease
inhibitor
an
van
and
pollen
and
inflorescence
The
receptive stigmas
opposed
(as
to
is undoubtedly increased in such a
flowers)
The congested arrangement of the flowers may make
system.
pollen transfer to such receptive stigmas likely.
Owing
in P. capitata, pollen must apparently be transferred
occur
from
the
anthers
to
geitonogamy.
equivalent
to
flower),
flower to the stigma of another
same individual.
that
on
as
one
of
flower
one
factors, for self-pollination to
above
the
to
This process is referred
In the genetic sense, geitonogamy is
autogamy
except
(self-pollination occurring within
that the services of a pollinator are
required (Faegri and van der Pijl, 1971).
The
process of geitonogamy in Phacelias with congested
inflorescences
is
well-described
by
Willis
(1895), in a
discussion of pollination in E. tanacetifolia.
dichogamy of the flower gives it some chance
especially as it is
cross-fertilization,
largely visited for honey and pollen; but it
seems probable that self-fertilization is more
common, even when it is visited by insects. The
are closely packed together...Insects
flowers
crawl over the whole mass of flowers, touching
stamens and styles indiscriminately, and probably
often knock pollen on to the stigmas from the
The
of
85
surrounding anthers...Every flower on the plants
examined set a full complement of seed.
Heckard
concurred
(1960)
observing
that,
common
the
in
with the observations of Willis,
although
self-pollination
probably
is
Magellanicae, "...it does not seem to occur
to any extent when bees are lacking."
During
visitors
insect
behavior
was
landing
the
on
Macior,
just
"floral
this
opportunity
for
the
same
moving
(1895). Upon
It could be seen
stamens
styles
are
inter-plant
visitation
observed.
In
as
many
of
giving
the
Even autogamy is possible if
Insects were often observed
mature.
plants
cross-pollination
and
undoubtedly
about,
another
to
close
often
so
the
same
by
way,
is
if
pollen
same
of
together
opportunity
an
P.
that
was also
pollinator
there
well,
the
on
populations
dense
particularly
this
(Dr. L.
the flowers.
inflorescence
In
nectar
on anthers long enough for stigmas of
to
one
the
Willis
by
collecting
geitonogamy.
flower
individual.
the
pushed
remains
from
capitata
from
process
are
pollen
surface,"
comm.)
flowers
some
described
as
inflorescence, the insects would crawl over
pers.
in
the action of
studies,
field
of
E. capitata plants was observed. Their
on
an
entire
that
course
the
for
is retained on the
body of the insect.
Cross-fertilization
evidenced
by
interspecific
observed
by
the
does
instances
intergradation
Heckard
occur
(1960).
of
at
in
hybrid
the
these
plants, as
swarms
and
by
tetraploid level, as
The suspected allopolyploid
86
origin
tetraploid species, such as E. hastata,
certain
of
would require cross-pollination events as well.
general, the pollination syndrome of E. capitata is
In
described
best
geitonogamy
facultatively
as
probably
is
cross-pollination
Though
xenogamous.
frequent, opportunities for
most
No evidence of apomixis in the
do exist.
Magellanicae has been observed (Heckard, 1960).
visiting
Insects
and
by
potential
most
and
of Entomology, Oregon State University.
pollinators
frequently
spp.)
J. Lattin, A. Moldenke, and W. Stephen
Drs.
Department
the
These
of E. capitata
corvmbosa were collected during May, 1984, and were
P.
identified
of
populations
several
listed in Table 12.
The
visitors were bumblebees (Bombus
observed
honey-bee,
the
are
Avis mellifera.
The variety of
this limited sample indicates a
insect
visitors
found
rather
general,
"open" pollination system for E. capitata.
In
of pollination ecology in California, Moldenke
study
a
both
that Phacelia species are heavily visited by
stated
(1976)
"specialist"
and
visiting
P.
species
while
generalists,
"generalist"
one
ordinarily
members
the
family
The Bombus
E.
corymbosa
instance,
an
individual
genus
a
vectors.
and
capitata
in
Melissodes,
of
in
of
specialists
are
of
visiting
Asteraceae (Dr. A. Moldenke, pers.
comm.), was found collecting nectar from P. capitata.
Seed
which
production.
Phacelia
pollinators
Methods.
To compare the extent to
capitata sets seed with and without natural
available,
five
exclosures
were established.
Table 12.
Insect pollinators of Phacelia capitata and P. corymbosa.
Phacelia species
Order
Family
Genus and species
P. capitata
Hymenoptera
Apidae
Apis mellifera L.
Bombus californicus Smith
B. caliginosus Frison
B. edwardsii Cresson
B. huntii Greene
B. vosnesenskii Rad.
P. corymbosa
Anthophoridae
Melissodes sp.
Megachilidae
Osmia sp.
Lepidoptera
Lycaenidae
Everes comyntas Godart
Hymenoptera
Apidae
Apis mellifera L.
Bombus vosnesenskii Rad.
88
These
exclosures
Area
Natural
set up in the Beatty Creek Research
were
on
April
fruiting
The
1984.
25,
inflorescences were retrieved on June 30, 1984.
The
which
exclosures consisted of small milk carton sections
were placed in a short segment of nylon netting.
The
below
the
netting
tied
was
inflorescence;
supported
tied
was
it
entire
the
stem
the
around
off
the top to a stake, which
on
were
exclosures
All
exciosure.
established before the first flowers had opened.
retrieving
Upon
from
natural
and
same
the
the exclosures, paired inflorescences
The number of flowers
pollination, were collected.
produced
seeds
by
been left open to
had
which
individuals,
five inflorescence pairs were
the
then counted.
Results.
enclosed
was
were
inflorescences
significant
no
excluded
agreement
noted
production
Seed
with
that
is
open-pollinated
in
summarized in Table 13.
and
There
seed production in inflorescences that
from
pollinators.
the
observations
These
of
results
Heckard
are in
(1960), who
caged plants of P. californica produced only an
occasional seed per cyme.
Although
for
the data indicate that pollinators are needed
appreciable
provide
information
self-pollinated
however,
seed
or
set
in
P.
capitata,
do not
on the extent to which the flowers are
out-crossed;
as
selfing is probably prevalent.
self-pollination
they
previously
stated,
Careful artificial
would be needed to determine the extent of
Table 13.
Inflorescence*
1-1 open
Seed production in open-pollinated and enclosed inflorescences of
Phacelia capitata.
Number of
flowers
158
Number of flowers
producing seed
Percent flowers
producing seed
61
38.6
Number of flowers
with evidence of
seed predation
Number of flowers
producing:
1 seed 2 seeds 3 seeds
18
55
6
0
0
0
0
0
1-1 closed
130
0
0.0
1-2 open
108
40
37.0
4
40
0
0
1-2 closed
136
0
0.0
0
0
0
0
2-1 open
247
175
70.8
20
85
87
3
2-1 closed
219
1
0
1
0
0
2-2 open
154
82
53.2
30
55
27
0
2-2 closed
154
5
3.3
0
4
1
0
3-1 open
59
49
83.1
0
49
0
0
3-1 closed
53
0
0.0
0
0
0
0
0.46
*-First number indicates individual plant; second number indicates inflorescence pair from each plant.
90
self-compatibility.
in
the
stigmatic
small
breaking
the
of
difficulties
complex.
a
owing
field,
hand pollinations are difficult
to
the congested inflorescences and
surfaces;
several
attempts
stems.
Heckard
(1960)
ended in the
alludes to the
in artificially self-pollinating plants in the
With few exceptions, artificial pollinations gave
percentage
low
Such
of seed set compared to that obtained by
natural pollinations.
Some
capitata
no
seeds
occurs
inflorescences of E.
in
In the cases where this was observed,
(Table 13).
holes
small
predation
seed
were present in the sides of the capsules, and
had
No insects were found that could
developed.
be implicated in this seed predation.
Pollen
viability.
A
Methods.
survey
of
pollen
viability
in
corvmbosa,
P. hastata, and P. heterophvlla ssp. viraata was
populations
Phacelia
of
capitata,
P.
The pollen grains were sampled from dried herbarium
made.
specimens.
One
single
flower,
cotton
blue
in
to
three
were
used.
dehisced
The
anthers,
all from a
pollen was stained with
and counts were made using a
lactophenol,
Zeiss light microscope.
Resul s.
The
pollen
viability counts are listed in
of
pollen
viability is often useful as a
Table 14.
The
degree
conservative
For
example,
sterile
indicator
reduced
hybrids
of
the
pollen
resulting
fertility
of individuals.
viabilities may characterize
from
interspecific
crosses.
91
Table 14.
Pollen viability in Phacelia species group Magellanicae.
Number of
pollen grains
examined
Species
Population
n
P. capitata
Bilger Creek
Boomer Hill I
Boomer Hill II
Bridge
Callahan Creek
Cow Creek I
Cow Creek II
Cow Creek Turn-off
Doe Creek
Doe/Thompson Ridge
The Drew
Elk Creek
Goolaway Gap
Little River
Myrtle Creek North
Rice Creek
Salt Creek
Weaver Road
11
P. corymbosa
P. hastata
Eight
Eight
Rough
Waldo
Wimer
11*
11
11*
22*
11*
11*
11*
11
?
11*
11
11*
11*
11*
?
11*
11*
Dollar Mtn I 22
Dollar Mtn II 22
& Ready
22
Road
Boomer Hill I
P. heterophylla Elk Creek
22
22*
22
11*
Percent
viable
pollen
Collection
65
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
Shelly
& Nelson
& Holmes
& Holmes
& Nelson
& Nelson
709
412
719
706
& Holmes
& Nelson
& Nelson
& Nelson
349
721
& Holmes
& Nelson
720
447
579
552
418
568
45
96
97
81
36
Shelly
Shelly
Shelly
Shelly
Shelly
700
702
698
697
691
1703
49
Shelly & Holmes 767
538
96
Shelly & Nelson 727
480
758
572
593
583
645
710
802
817
526
516
710
865
198
761
813
223
687
95
93
58
95
98
97
92
85
86
99
98
71
98
93
96
80
71
*-Pollen viability obtained from a specimen other than the chromosome count
voucher.
724
769
781
754
733
783
734
726
737
792
725b
92
viability
Thus,
suspected
degree
may be of use in studying known or
counts
intergradient
interfertility
of
In
this
project,
be
used
to
in assessing the
and
populations
species or populations.
between
it was hoped that pollen viability could
study
capitata
populations
Callahan
Creek),
hybrid nature of the P.
potential
the
containing tetraploids (Elk Creek and
and to describe the range of variation in
pollen viability among diploid populations of the species.
The
results
percentage
of
Although plants from
of E. capitata generally show the high
populations
diploid
inconclusive.
are
pollen expected in nonhybrids, there
viable
are
four exceptions (Boomer Hill II, Elk Creek, Salt Creek,
and
Weaver
severely
high
lowered
and
low
tetraploid
pollen viabilities were found in plants from
of
E.
diploid
viability
pollen
Due
corymbosa.
nonhybrid
in
populations,
In addition, both
(i.e., less than 75%).
localities
variability
an
in which pollen viability of diploids is
Road)
and
to
this
tetraploid
is of limited usefulness as
indicator of hybrid origin in the tetraploid E. capitata
populations.
Environmental
Magellanicae,
the
observed,
be
but
attributed
involving
found
may
strongly
influence pollen
Heckard (1960) found that in nonhybrid species
viability.
of
factors
pollen
viability
was
always
that "...a variation of about 10 percent can
to
studies
that
high
the
of
pollen
environment."
E.
In an extreme case
californica and E. imbricate, he
viability
varied
within
the
same
93
populations
concluded
through
depending
that
decrease
observed
abortion
period was "...largely a result of a
blooming
the
pollen
increasing
the
Heckard
when they were sampled.
on
in available water and an accompanying decrease of
Such
vigor."
temperature
environmental variations, perhaps including
may
fluctuations,
to
contributed
have
the
variability in pollen viability observed in this study.
Competitive ability and ruderal response
The
hypothesis
serpentine
soils,
physiological
because
by
restrictions
of
several
as
result
a
possible
of
but
habitats
such
to
ability
competitive
poor
of
communities
only
not
are limited to
species
certain
that
more
in
also
closed
non-serpentine substrates, has been offered
botanists.
(1951),
Kruckeberg
in
found
studies
and
off
comparing
growth
serpentine
soils, concluded that the superior growth of all
the
tested
species
serpentine
uniquely
merely
serpentine
non-serpentine
aside
endemics
of
restricted
In
later
a
to
requirement
specific
some
serpentine."
plants
than
chemical
from
serpentine
by
soil suggests that
not
are
on
report
1954), he again suggested that, because growth
(Kruckeberg,
of
endemics
because
provided
species
non-serpentine
on
serpentine
"...the
plant
of
unable
found
to
be
better
on
on serpentine soils, some other factor
tolerance
restriction.
are
was
to
must
be sought to explain
He postulated that the serpentine
maintain
themselves
against the
94
competition
rigorous
Stebbins
Major
and
ability
grow
to
that
from extinction by
preserved
are
stating
agreed,
(1965)
endemics
"...serpentine
the
non-serpentine plant communities.
of
on serpentine where more competitive
plants cannot grow."
Phacelia
example
of
A
and
grow
However,
a
on
E.
possibly
the
Boomer
species
the
apparently
provide
poor
with
endemic
excellent
an
competitive
the species has the ability to germinate
non-ultramafic
soils (80% germination rate).
capitata is not found off serpentine soils (or
metamorphic derivative thereof, i.e., at
close
Hill
observations
to
greenhouse germination test on non-serpentine
that
showed
appears
serpentine
a
ability.
soil
capitata
indicate
may
poor
be
the
that
result,
a
The following field
in nature.
site)
I
competitive
serpentine restriction of
in
part, of the species'
evident even in its
ability,
own native, undisturbed habitat.
often
serpentine communities, P. capitata is
undisturbed
In
found
in
serpentine
slopes
frequently
along
channels
in
very specific types of microsites.
at
the
Peel
intermittent,
which
cover
of
site,
individuals
vernally
other
On the
scoured
species
occur
stream
is low.
The
plants
which occur in these shallow channels are often very
large,
as well.
of
annual
In the surrounding vegetation, where cover
grasses and other forbs is higher, the frequency
of E. capitata is noticeably less.
The
apparently
poor
competitive
ability
of
the
95
species,
is
again within its own undisturbed native community,
also
evidenced
Creek.
Here,
infrequent
throughout
outcrops
interesting
are
However,
layer
instances
in
of the Pinus
large
where
E. capitata plants
bedrock occur,
In addition, an
the cracks of the rocks.
in
and
scattered
widely
herbaceous
the
serpentine
of
frequent
are
individuals
savanna.
jeffreyi
serpentine slopes along Beatty
the
on
example which suggests the need for a very open
seed germination was observed here.
"safe
site"
In
location where a dead Pinus jeffreyi trunk had fallen
a
over,
root
hole
large
a
capitata
left in the soil by the upturned
was
hole contained seven large, floriferous
This
mass.
Phacelia
good
for
individuals,
while
the surrounding
in
layer, very few plants, and no other such concentrated
herb
"clumps," were found.
Owing
to
apparent
the
competition,
perhaps
in
establishment
phases,
one
invade
the
grow
and
case--at
serpentine
abundant
rock
quarries,
might
all
sites
18
has
on
for
lowered
germination
seed
the
expect
well in disturbed areas.
soil
more
requirement
P.
capitata
and
to
Indeed, such is
where some disturbance of the
occurred,
P.
capitata is noticeably
the open, barren areas (i.e., roadbanks,
etc.)
than in the surrounding, undisturbed
serpentine habitat.
Quantitative
were
collected
traversed
data
at
Lee
to illustrate this ruderal response
Creek,
by a logging road.
on
a
serpentine hillside
Seven 4x4 m square plots were
96
set
three
up:
plots
adjacent
bedrock
The
of
compared
however,
the
on
cover
of
fact
that
to adjacent undisturbed areas where cover
The plants are most abundant,
serpentine
raw
species is lower.
other
the
cotyledons
an
still
on the exposed
a higher percent cover
species is higher.
other
on
The results are given in Table
outcrop.
have
and
plot
one
and
show that P. capitata individuals are more
data
numerous,
outcrop
vegetation,
undisturbed
in
15.
meters above the uphill edges of the roadbank
five
located
plots on the barren roadbank, three plots
only
Also of interest is the
observed
seedlings
were
apparent)
where percent
roadbanks
in
with
(i.e.,
of the roadbank
two
plots.
would
It
be
naive to conclude, on the basis of these
observations alone, that competitive exclusion is the
field
or even the most important, cause of the serpentine
single,
restriction
The situation may be much more
of E. capitata.
and more controlled experiments would be necessary
complex,
to fully understand the autecology of the species.
As
example,
an
which
growth
rates
of E. capitata on
and normal soils could be compared, to determine
serpentine
whether
the
the
have
species
grows
ordinarily
toxic
elements,
such as magnesium.
evidence
indicating
associated
with
serpentine
soils
optimally
that,
of
concentrations
certain
Also, Grime (1979) summarizes
in
instances
nutrient-deficient
were
on ultramafic soils
where
habitats
plants
such
as
supplied with high rates of mineral
Table
Plot
15. Ruderal response of Phacelia capitata at Lee Creek.
Data collected from seven 4x4 m plots.
Number of plants
(non-flowering; flowering)
Percent cover
Number of
seedlings
Percent cover of
other vegetation
Herbs
Shrubs
Trees
0.0
0.0
A. Roadbank plots
8.0
5
1.5
14; 31
5.0
1
2.0
1.0
10.0
3; 24
25.0
0
1.0
0.0
0.0
1
0; 28
2
3
B. Undisturbed plots
4
4; 4
1.5
0
55.0
0.0
15.0
5
0; 0
0.0
0
95.0
0.0
75.0
6
1; 0
0.1
0
90.0
0.0
70.0
5; 9
2.0
0
8.0
0.0
0.0
C. Rock outcrop plot
7
98
nutrients
(i.e.,
in quantities detrimental to the growth of such
accumulated
plants.
which
these nutrients appeared to be
calcium),
P. capitata provides such an example, in
Perhaps
there
a "requirement" for the low concentrations
is
of
phosphorous and calcium which probably characterize many
of
its sites.
and
requirements
case
on
capitata
example
of
naturally
If
a
soil
generally
in
serpentine
a
may
endemic
lower
the
such as P.
Lastly, E. capitata may be a good
habitats characterized by high stress
status) and light disturbance.
nutrient
characteristic
competitive
the
substrates,
the intrinsically slower growth rate which is
then
so,
operate
"stress-tolerator" (Grime, 1979), in that it
exists
poor
(i.e.,
of
such soils.
on
may
Lack of required soil properties, in
non-serpentine
ability
competitive
ability
competitive
poor
also exists.
together
this
Of course, the possibility that special soil
of such species may also decrease
ability
of
this
and
other
serpentine
endemics.
Thus,
(1975)
as
state,
hypothesis,
ecology
unknown.
Kruckeberg
and
Proctor and Woodell
the full nature of the competitive exclusion
which
and
(1954)
may be so important when considering the
evolution
of
serpentine
endemics,
remains
Clearly this is a fertile area for research, and
Phacelia capitata may provide a very revealing subject.
99
DISCUSSION AND CONCLUSIONS
$peciation and evolutionary relationships
provides
4agellanicae)
both the morphological and
complexes,
such
a mature polyploid
of
example
an
group
(species
alliance
magellanica
Phacelia
The
complex.
In
ecological
extremes
species.
However, the diploids are often less common than
are usually represented by the diploid
polyploids.
the
restricted,
and
amounts
the
populations
their
geographic
Their
more
are
genetic variation within
of
usually
is
ranges
other,
each
with
the
addition,
In
less.
or
diploids
are
usually
sympatric
only
in restricted areas, so that the likelihood
hybridization
of
of
doubling
and
different
involving
allopatric
diploids
much
is
number
chromosome
reduced (Stebbins,
1971) .
the
Of
species
six
exclusively
almost
considered
in
"morphological
in
and
ecological
attributes
occur
is
its
the
is
in
P. capitata and P.
characteristics
the
form
serpentine soils.
of
example
good
a
extreme,"
ecological
morphological
to
study:
this
these
of
distinct
restriction
Maaellanicae that occur
the
or partially as diploids, two have been
detail
Each
corymbosa.
of
having
and
a
very
special
of virtually complete
Of all six species which
as diploids in the Magellanicae, however, P. capitata
most
restricted
morphological
in its geographical range.
distinctiveness,
restriction
Thus,
to
100
distribution
pattern,
diploid
the
(or a close derivative), narrowly endemic
soils
serpentine
level
almost
and
make
exclusive occurrence at
an
capitata
E.
unusual, unique
entity in the Mapellanicae.
It
of
difficult to elucidate the evolutionary history
is
such
diploid
a
speculation
of
in
a
mature
complex, but
In considering possible modes
interesting.
is
polyploid
speciation for E. capitata, it is convenient to consider
types
two
origins:
insular
endemics
are
diverged
from
ancestral
those
upon
depleted
and
which
their
in
Insular
common,
but which
were
never
differ
endemics.
following
the
were
more
once
of these
establishment
a small, isolated area.
which
those
which
species,
a small group of individuals of a widespread
species,
individuals
are
endemic
of
Depleted species
common, widespread, and
richer
in biotypes (genetically different populations), but
whose
present
variability
1942a)
two
endemic
to
depletion
genetic
of
of restricted species may be further
types
one another by two criteria.
from
insular
living
Secondly,
specialized
the
form
continental
species;
form,
if
First, if
closely related to widespread species which
is
surrounding
in
strictly
other
due
subsequent loss of biotypes (Stebbins,
through
distinguished
occur
is
.
These
an
rarity
it
if
areas, it is probably a
it is closely related to no
is more likely a depleted species.
endemic
is
morphologically
a
highly
compared to its relatives, it is probably
101
an
insular
descendent
if
it
is
less
it may be a depleted ancestor (Stebbins, 1971).
specialized
Based
competitive
distribution,
and morphological characteristics of Phacelia
theories regarding the origins of modern-day
and
vegetation
geographic
restriction,
edaphic
ability,
cytology,
the
regarding
observations
on
capitata,
while
these,
of
Klamath
the
in
can
one
Mountains,
support,
much difficulty, an insular mode of origin for this
without
species.
polyploid
In
with
complexes
lowest
the
number
in nature, the members
found
chromosomes arc the primitive
of
while the polyploids are derived from them (Stebbins,
ones,
1942b).
addition,
In
irreversible
1971).
at
least
such
as
the
divergence
and
the
can
Magellanicae
an
be
evolution in
known,
and
the diploid level will have
at
mode
primary
of
direction
the
speciation
initial,
predominantly
is
lower to higher levels (Stebbins,
from
Thus,
complexes
formed
trend
polyploidy
of
evolution for such
groups.
Because P. capitata occurs almost exclusively as a
diploid,
it
leading
to
chromosome
can
from
be concluded that the evolutionary events
can
the
origin
level.
described
be
the
This
as
ancestral
occurred at this
species
the
of
being the case, then P. capitata
one
of the evolutionary "offshoots"
stock
diploid
which
formed
the
foundation of the Magellanicae.
The
Tertiary
northward
Geoflora
migrations
probably
of
brought
elements
with
of the Madrothem,
sometime
102
since
beginning
the
Phacelia
species
correlates
which
from
mature
Pleistocene
or
500,000
10,000,000
and
Pliocene
and
selection
for
between
approximately
Upon
the ancestral
substrates
serpentine
the
perhaps
few,
a
may
capitata
ago.
the
in
southwestern Oregon, the process of natural
serpentine-tolerant
divergence
epochs,
years
with
during
originated
probably
contact
California
E.
This
polyploid complexes that may presently be
of
considered
species'
evolved.
capitata
E.
well with the conclusion of Stebbins (1971) that
majority
the
the Pliocene epoch, the ancestral
of
partially
preadapted,
could have begun.
individuals
Further
ended with the "insular" isolation of
have
the serpentine outcrops which it presently
on
occupies.
The
ability
fact
helps
only provides one possible explanation for the
not
species'
present
explain
species.
If
serpentine
reproduce
this
rise
to
capitata
thus
may
imbalanced
to serpentine soils, it also
restriction
the hypothetical evolutionary history of the
the
few
tolerance
individuals
only
were
which were developing
able
to
survive
and
in the more open communities on serpentine soils,
could
tolerance,
E. capitata possesses poor competitive
that
reinforce
selection
the
for
serpentine
and further divergence of the individuals giving
the
species
through
time,
could
occur.
despite
The persistence of E.
poor competitive ability,
be owed to its ability to tolerate the chemically
soils,
and
to
its
consequent "escape" of the
103
of competition. The morphological distinctness of Z.
rigors
capitata
from other widespread members of the Magellanicae,
well
as its restricted geographic distribution, provide
as
further evidence that the species is an insular endemic.
The
cytogeography
the
origin
of
species
in
Oregon.
because
both
It
logical
is
corymbosa
Two problems make
previously.
location
for
location
species
First, Z.
P.
of
diploid
corvmbosa
is
in
Siskiyou
County,
175 km south of the southernmost
capitata (Heckard, 1960).
Z.
both
are
known
northernmost
The
approximately
California,
substantiate.
to
hypothesis
not known to occur as a diploid in Oregon, as
is
discussed
speculate that,
are largely restricted to serpentine
species
difficult
8
pattern of these
to
they may share a common ancestor.
a
Figure
species.
distribution
allopatric
the
this
latter
the
for
illustrates
soils,
a related
is also of interest when considering a possible
south,
mode
corymbosa,
P. capitata on ultramafic soils to
replaces
which
species
phacelia
of
distinct
they
morphological
very
Second, the two
entities in the
different
from
one
Magellanicae,
and
another.
a common diploid ancestor was in fact shared
If
are
two species, it must surely have been long extinct.
by
the
No
evidence
for any present genetic interaction between Z.
capitata and Z. corymbosa was found.
Even
restricted
the
the study of a morphologically distinct, narrowly
endemic
difficulties
such as Z. capitata has not been spared
of
polyploidy
and
interspecific
104
capitata 2n
* capitata 411
0 capitata ?n
N
* corymbosa 4n
corymbosa ?n
CALIFORNIA
Figure 8.
Distribution of Phacelia capitata and P.
corymbosa in southwestern Oregon. Ultramafic
formations outlined (Wells and Peck, 1961).
Phacelia corymbosa locations are plotted from
herbarium specimens, except for the tetraploid
location at Sexton Mountain, Josephine County
(Heckard, 1960).
105
with
Further,
diploids
factors
...confounding
as
due
introgression
to
Stebbins
tetraploids."
due
to
fact
the
taxonomists
origin
limitations
necessary
is
in
other
or
(1971) concluded that this may be
guides
tetraploid
such
diploids
local
from
inadequate
many
of
caution
diploids."
in
the morphological criteria used by
that
are
encountered
that
its later
and
variation of tetraploids in one place
the
"...interpreting
taxon
a
the
states,
(1976)
other polyploids may cause
and
not
states
he
by
polyploidy
of
"...attainment
interaction
Jackson
As
hybridization.
evolutionary
the
to
With these
populations.
mind, the interrelationships of E. capitata
in
within the Magellanicae may be considered.
Fhacelias
the early collections of Magellanicae -type
of
Certain
in
County,
Oregon
(see
which
Kruckeberg
(1956)
likened
Relationships),
capitata,
are
P.
at
Although
genomes
P.
genetic
between
These
collections may
these two species, but most
this time best referred to the widespread, variable
The
Heckard
of
which
P.
specimen
collected
by
Constance
and
DS) is the most intermediate in appearance.
(2956,
conspicuous,
latter
hastata.
E.
intergrades
hastata.
Rollins
to
of the species with other members of the
especially
represent
former
of
suggestive
are
interrelationships
complex,
Taxonomic
Douglas
(pers. comm.) felt this plant may combine
capitata
and
P.
corymbosa,
it lacks the
intermediate-length glandular trichomes of the
would
be expected in such a hybrid.
On this
106
basis,
because
and
was
specimen
easily
found
Nebo
Mt.
where this
not on serpentine soil, it is most
is
to E. hastata, with a note as to possible
referable
with
affinities
near
area
the
capitata.
P.
found the former
have
I
species to be extant in the Mt. Nebo area.
A
location
only
with
sympatry
examples
768,
their
possession
buds
capitata
and
interest
at
P.
perhaps
predominantly
chromosome counts on
On the basis of other features,
between
P.
at least be postulated.
Of
hybridization
of
can
site is the less stringent nature of the
between
intermediate
is
habitats
to
which
E.
capitata
the
disturbed
the
factors
nature
which
of
the
highly
is largely
and a more normal soil chemical profile.
restricted,
the
hastata
this
which
serpentinized
plus
occurrence
the
however,
soil,
meiosis.
past
of
specimens were unsuccessful, as all the
intermediate
were
possibility
narrow,
Attempted
leaves.
the
be intermediate, however, in
to
relatively
of
as a tetraploid, a
Some collections (Shelly &
species.
appear
771)
silvery
entire,
limits
ordinarily
Holmes
latter
the
"good"
that the former species
showed
and
between
hybridization
representing
individuals
species
which
situation
capitata was found in biotic
E.
of
diploid
a
as
This is
member of the complex (E. hastata).
another
each
of
occurs
where
counts
Chromosome
these
is the Boomer Hill I site.
intergradation
species
the
which is potentially revealing with respect to
site
This,
roadside habitat, are
have allowed P. hastata and E.
107
capitata to become sympatric here.
populations
The
origins
determining
population
at
Creek population, only a tetraploid count
It may be best, in seeking to describe the
obtained.
was
significance
in
a mixture of diploids and tetraploids, while
is
Callahan
the
km
presence of tetraploid
The
other.
approximately
at both of these sites may therefore be related
(i.e., hybridization
event
polyploidization
single
a
each
within
occurring
sympatry,
of
individuals
to
two sites, to realize that they are
these
of
neighboring
0.8-1.0
in
The Elk Creek
individuals.
such
of
involved
problems
the
of
examples
further
are
of P. capitata containing tetraploids
followed by chromosome doubling) in the past.
the
On
basis
diploid
other
appears
This
heterophvlla.
P.
with
species
is
E. capitata, occurring along
highway both north and south of the serpentine outcrops
In surveys throughout the vicinity, no
Creek.
above
Elk
other
morphologically
were
found.
individuals
undisturbed
suggests
Highly
of
E.
at
possibly
variable,
capitata
are
this
not
tetraploid
restricted
location,
but
to
occur
areas above the road as well.
serpentine
the
in
This
that the possible hybridization event which led to
polyploidization
habitat
distinct members of the Magellanicae
roadbank
disturbed
the
these intergradient populations
for
sympatric
neighboringly
the
parent
be
to
field observations, the most likely
of
disturbance
of
in
E.
the
capitata
area,
occurred
prior
to
the tetraploids having
108
since colonized the roadbank.
With
respect
E, capitata
of
species
conclusions
have
from hybridization in the
resulted
either a species similar to P. egena or with the
with
Long-term introgression
sympatric E. beterophvlla.
involving
have
It is feasible that the
lines.
and leaf lobing patterns of E. capitata at
could
Creek
locally
these
along
corollas
larger
past
its morphological characteristics
are too poorly known at this time to allow any strong
here,
Elk
The occurrence of this
egena.
P.
and
Oregon,
in
population, Heckard
have interacted with a Californian member
may
Magellanicae,
the
Creek
Elk
collection (2930, JEPS) the possibility that
his
on
noted
the
to
tetraploids
the
and
diploids
at this site may
so obscured the hybrid nature of the plants that exact
determination
lengthy
field
origin
their
of
and
artificial
is
not
possible.
Only
hybridization studies could
perhaps offer answers to the origin of these populations.
Figure
P.
capitata
complications
the
9 summarizes the possible interrelationships of
within
patterns
of
cytological
Magellanicae.
In the face of the
discussed above, it is appealing to interpret
occurrence
"evolutionary
the
of
limited
noise,"
when
interspecific gene exchange as
one
considers the distinctive
morphological, ecological, and almost complete
isolation
southwestern Oregon.
of E. capitata within the complex in
109
Figure 9.
TETRAPLOID
Possible interrelationships of Phacelia capitata
within the species group Magellanicae.
1 egena 1
(n=22)
DIPLOID
(n=11)
heterophylla
ssp. virgata I--
110
The endemic status o
Although
and
discussions
of
area
which
taxa
which
Neoendemics
evolved in a particular
have
Paleoendemics
Major, 1965; Stott, 1981).
former range (Stebbins and
third
described,
category,
which
is
above. extremes.
area,
(Richardson,
the
is
holoendemic,
has
also
been
intermediate in character between the
Such
species, even when covering its
a
still narrowly restricted, most often by
physical
of
or physiological (ecological) barriers
characteristics
The
1978).
categories
endemic
formed the basis for
have not yet spread.
they
types,
extreme
confined to a very limited portion of their
now
are
virtue
two
which once possessed much wider distributions but
taxa
maximum
have
species has
endemic
endemism since the late 1800s.
new
A
of
complex,
paleoendemics,
represent
which
very
been
neoendemics
are
classification
the
traditionally
from
the species
with
of
three
the
respect to various attributes are
summarized by Richardson as follows:
Endemic status
Neo- Holo- Paleo-
Attribute
taxonomically isolated
geographically isolated
polymorphic
derived characters
environment stable
ploidy level high
potential to expand area
age (recent +, old -)
As
he
makes
attribute
clear,
+
+/+/-
+/+/+/are
-
-
+?
+
+
+
there
E. capitata
+
+
-
+
+
+
-
many
(+)/-
(+)/-
exceptions
+
-
+?
+
to
the
states, and there is no clear distinction between
the three types.
111
is geographically and ecologically
Since
E.
capitata
restricted
to
a small area, and surely did not once occupy
a
much
larger
a
neo-
former
its
seems
factors which might prevent
other
or
south of its present
areas
unless there are unknown soil
logical,
climatic,
The
seems to most closely fit the
it
serpentine
the
to
distribution
this.
It is,
The possibility of E. capitata expanding
category.
range
a paleoendemic.
not
is
holoendemic;
or
chemical,
it
clear if the species is best characterized as
not
however,
area,
soils occupied by E. capitata and E.
analyzed
corymbosa are quite similar in chemical nature.
According
localized
generalized
must
specialized
in
North
western
be
Stebbins
to
America,
distribution
recent
of
habitats
This,
origin.
such
as
vegetation,
surrounding
the
in
regions
in
are
close relatives of more
have
and
which
endemics
(1971),
too,
supports
the
classification of E. capitata as a neoendemic.
Thus,
the
most
reasonable interpretation of Phacelia
capitata may be to classify it as an insular neoendemic.
Recommendations for preservation
Endemic
taxa
such
as
Phacelia capitata are of great
As a consequence of their
scientific
interest
restricted
geographical distributions and often specialized
ecological
requirements,
become
endangered
destruction.
and
as
a
value.
they
are
also
most
likely
to
result of habitat modification or
It is imperative that such species be studied
112
and documented before they are lost to extinction.
'hacelia
serpentine
increase
at
time it is not appropriate to consider the species
with
was
placed
and
endangered
of
Natural
been
Thus, at least
abundance of the species.
the
"endangered"
list
paradoxical case for a
a
endemic, since habitat disturbance has led to an
in
this
presents
capitata
Most recently, the species
extinction.
on the "watch list" of Oregon rare, threatened,
taxa
plant
threatened
Heritage
after being removed from a
species,
Data
throughout
Base,
range (Oregon
their
In addition, it has
1985).
recommended for removal from candidacy as a threatened
species under the Federal Endangered Species Act of 1973.
The
the
geographically
species
capitata
can
on
provide
does
a
narrow
a
"watch list."
restriction of
serpentine
basis
for
maintaining
E.
In this way, its populations
be monitored to detect any decline in numbers which may
occur
with
increased
further
mining
tendency
in
disturbed
for
still
important,
population
the serpentine substrates.
of
allowed
populations
destruction of its habitat, such as by
areas
has,
for
the
Its weedy
time
the persistence of this unique species.
of
such
ecology,
however,
species,
to
so
protect
that
being,
It is
undisturbed
features of their
habitat requirements, and evolutionary
uniqueness may be studied in natural settings.
113
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APPENDIX
118
APPENDIX
cited
Specimens
addition
(in
to
those
discussed in the
Taxonomic Relationships section):
Hwy.
isotype),
m,
OREGON:
Coos Co.: N of St.
km E of Bridge, 100 m, Kruckeberq 2702 (WTU,
3.2
42,
Douglas
Kruckeberg.
capitata
Fhacelia
$undberg
1031
(OSC), Shelly k Nelson 754 (OSC).
Beatty Creek, 0.4 km N of Cow Creek Road, 320
Co.:
Shelly 419 (OSC); BLM rd. 29-5-11.0, 2.3 km NW of Bilger
Creek
Road,
Road
(BLM
m,
Shelly
0.5
km
from
8.1 km SW of Interstate Hwy. 5,
k Holmes 766, al (OSC); BLM rd. 30-6-4.1,
BLM rd. 29-6-34.0, 850 m, Shelly £ Holmes 781
Creek, Umpqua NF rd. 3230, 0.5 km W of St.
Callahan
227,
Hwy.
30-6-4.2),
rd.
760
(OSC);
m, $helly k Nelson 724 (OSC); Boomer Hill
490
490
Fosback
m,
s.n,.
(OSC), Shelly 440 (OSC),
& Nelson 733 (OSC); Cow Creek Road, 0.5 km W of Doe
$helly
Road,
Creek
300
m,
Shelly 405, 70n (OSC); NW side of Cow
km SW of Doe Creek Road, 300 m, $helly 411, 412
Creek,
1.2
(OSC);
Cow Creek Road, 4.8 km W of Glenbrook Loop Road, 230
m,
$helly
of Doe Creek Road, 640 m, Shelly k Holmes 783 (OSC);
NW
The
Drew,
2930
m,
$helly
(JEPS),
Creek,
(OSC),
270
810
Shelly
k Nelson 734 (OSC); Elk Creek,
St. Hwy. 227, 1.6-4.0 km SE of Tiller, 375 m, Beckard
along
Lee
719 (OSC); Doe Creek Road, 1.9 km N of Cow
Road, 400 m, Shelly 706 (OSC); BLM rd. 30-7-14.0, 3.2
Creek
km
681,
m,
BLM
Shelly
parker
rd.
433 (OSC), Shelly k Nelson /21 (OSC);
28-4-15.0, 415 m, Fosback It al. s.n.
399 (OSC); Little River, 5.9 km SE of Glide,
k Parker s.n. (OSC), Shelly 111, 1Q. (OSC);
119
14, 1.6 km NW of Myrtle Creek, 245 m, Howell 28807
Rt.
Co.
721 (OSC); Nickel Mountain, ca. 365 m,
Shelly
414,
Detling
6339
(ORE);
295
Knouse
(CAS),
m,
flank
736
Shelly
(OSC), Shelly 354 (OSC); SW
p.n.
BLM rd. 29-7-36.0, 710 m,
Creek,
Rice
(OSC);
792 (OSC); Salt Creek, BLM rd. 30-7-36.0,
Holmes
m, Shelly & Nelson 725, 725b (OSC); Weaver Road, 2.9 km
340
SW
k Moore
Red Mountain, 930 m, Detling 5314 (ORE), Shelly k
of
Nelson
rd. 26-3-34.2, 0.5 km E of Peel,
BLM
of Interstate Hwy. 5, 210 m, Ornduff s.n. (JEPS), Shelly
Goolaway Gap, 930 m, Shelly
Jackson Co.:
(OSC).
720
689,
& Nelson 737, 738 (OSC).
P.
Road,
Jackson Co.:
OREGON:
Creek
Slate
of
Co.:
Grave Creek
Road, 850 m, Shelly 441,
Eight Dollar Mountain, Siskiyou NF
at Illinois River, 390 m, Shelly 700 (OSC); Eight
4201
Mountain, Deer Creek, 5 km W of Selma, 450 m, Shelly
Rough
(OSC);
Cave
N
Josephine
Dollar
702
km
3.5
(OSC).
rd.
Jepson.
corvmbosa
and
Ready
Botanical Wayside, 8 km S of
425 m, Shelly 698 (OSC); Waldo, 4.2 km E of
Junction,
O'Brien,
475
m, Shelly 697 (OSC); Wimer Road, 6.5 km SW of
O'Brien,
490
m,
Constance & Bacicialupi 3394 (WTU), Shelly
691 (OSC).
E.
Douglas
hastata
Boomer
Hill
Interstate
(OSC).
Hwy.
Road
5,
ex
Lehmann.
(BLM
760
rd.
m,
Douglas Co.:
OREGON:
30-6-4.2),
8.1
km
SW
of
Shelly & Holmes 767, 768, 771
120
P.
heterophylla
Douglas
OREGON:
Road,
227,
(OSC).
300
0.8
Co.:
Shelly
m,
km
Pursh
E
ssp.
yin:rata
(Greene)
Heckard.
Cow Creek, 0.1 km W of Doe Creek
714 (OSC); Elk Creek, along St. Hwy.
of Tiller, 350 m, Shelly k Nelson 727, 728