POLLEN-OVULE RATIOS: A CONSERVATIVE INDICATOR OF
BREEDING SYSTEMS IN FLOWERING PLANTS
ROBERT WILLIAM: CRUDEN
Department of Botany, University of Iowa, Iowa City, Iowa 52242
Received August 8, 1975. Revised March 26, 1976
The evolutionary shift from xenogamy
(outcrossing) to autogamy (selfing) has
been mediated through decreased flower
size and alterations in floral morphology
(Ornduff, 1969) which reduce the ener-
getic cost per flower (Cruden, unpubI.)
and facilitate self-pollination, respectively.
Recently, a number of workers have shown
that the flowers of self-incompatible and
other xenogamous taxa produce more pollen
grains than closely related self-compatible
and/ or autogamous taxa (Arroyo, 1973;
Baker, 1967; Cruden, 1973; Gibbs et aI.,
1975; Lloyd, 1965; Vries, 1974). In con-
trast to other floral characteristics the
number of ovules per ovary in the taxa
studied did not decrease. In these taxa
the evolutionary shift from xenogamy to
autogamy has been accompanied by a de-
crease in the ratio of pollen grains to
ovules (P /0).
It is generally accepted that differences
in breeding systems tend to be correlated
with successional stages. Colonizers of
disturbed habitats and plants of early
successional stages tend to be self-com-
patible and/or autogamous (Baker, 1955,
1959, 1965; Stebbins, 1958) whereas
plants of later successional stages, includ-
ing tropical forest trees, tend to be
xenogamous (Baker, 1959; Bawa, 1974;
Bawa and Opler, 1975; Stebbins, 1950,
1958). Since, as I will show, Pl O's are an
integral part of a plant's breeding system,
they also should be correlated with habitat
or successional stage.
My overall objective is to show that
P/ O's reflect the likelihood of sufficient
pollen grains reaching each stigma to re-
sult in maximum seed set. The more ef-
ficient the transfer of pollen, the lower
the P /0 should be. It follows logically
EVOLUTION 31:32-46. March 1977
that cleistogamous flowers should have the
lowest P/ O's, and that autogamous flowers
will have lower P/ O's than xenogamous
flowers, i.e., that P/ O's are correlated with
breeding systems. My data show that this
is the case and suggest that P / O's are
integrated with other facets of a plant's
breeding system. Second, I show that
there is a strong correlation between the
breeding system, hence P/ O's, and suc-
cessional stage. In addition, I briefly
circumscribe what constitutes a sufficient
number of pollen grains to assure maxi-
mum seed set and discuss evidence that
supports the notion that some plants tend
to minimize pollen production. Finally, I
discuss deviations from the general pattern
to show that P / O's are a better predictor
of a plant's breeding system than other
morphological characteristics.
MATERIALS AND METHODS
The pollen-ovule ratios were determined
by dividing the estimate of the number of
pollen grains per flower by the number of
ovules per flower. Several methods were
used to estimate the number of pollen
grains. If the number of pollen grains per
anther was 2000 or less, all of the pollen
grains in an anther were counted and then
multiplied by the number of anthers. In
some instances half an anther was used
and the number of pollen grains was multi-
plied by two to obtain the number of pollen
grains per anther. If the number of pollen
grains per anther was greater than 2000
then the pollen grains were suspended in a
solution of aniline-blue in lactophenol and
the number of pollen grains in a known
volume of the suspension was counted.
This number was multiplied by the dilution
32
 BREEDING SYSTEMS IN PLANTS
TABLE 1. Breeding systems and Mean pollen-ovule ratios of species in Appendix I.
OeI Breeding System N=
0 Cleistogamy 6 4.7
1 Obligate autogamy 7 27.7
2 Facultative autogamy 20
3 Facultative xenogamy 38
4 Xenogamy 25
factor and then by the number of anthers.
The number of ovules was determined by
counting the number of ovules in the ovary
of the flower used for the pollen count.
With the exception of Geranium maculatum
it was assumed that all ovules would
develop into a seed if fertilization occurred.
For each population an "outcrossing in-
dex" (OCl) was determined from either
preserved or living material. Each OCI
is the sum of assigned values for three
characteristics of the flower and floral
behavior: 1) diameter of the flower, or
head in Compositae, i.e., the target area.
Flowers were assigned to one of four
classes; corolla up to one mm wide = 0,
1-2 mm wide =1; 2-6 mm wide =2;
more than 6 mm wide = 3. 2) temporal
separation of anther dehiscence and stigma
receptivity. Homogamy and protogyny re-
ceived a value of O. Protogyny, although
promoting out-crossing, does not preclude
self-pollination. Protandry was given a
value of 1. 3) spatial relationship of
stigma and anthers. If the stigmas and
anthers were at the same level and contact
between anther and stigma seemed possible
the value was O. If the stigma and anthers
were spatially separated and contact
seemed unlikely the value was 1. Out-
crossing indices were computed prior to
the pollen-ovule ratios.
Reproductive characteristics (see Table
2) were determined by the examination
and manipulation of plants in the field and
greenhouses. Cages were used in the field
to exclude insect visitors from facultatively
xenogamous and xenogamous species.
Plants that set no seed with pollinators
excluded but set seed when selfed were
classified as self-compatible, pollinator re-
33
Pia log Pia
x of x ± S.E. x of x ± S.E.
± .7 .65 ± .07
± 3.1 1.43 ± .05
168.5 ± 22.1 2.15 ± .06
796.6 ± 87.7 2.81 ± .05
5859.2 ± 936.5 3.65 ± .06
quired. Presence of nectar was determined
by observing nectar in the flowers, or in-
ferred from the repeated and systematic
probing of flowers by nectivorous insects.
Habitats were classified as 1) highly
disturbed, i.e., those that were subject to
repeated disturbance, such as mowed
lawns; 2) early successional, e.g., unmowed
roadsides, old fields, etc.; 3) late succes-
sional stages in which pollinator activity
was unreliable; and 4) late successional
stages in which pollinator activity was re-
liable. The late successional classes include
communities in which successional change
is slow or not likely to occur but in which
seasonal changes including canopy closure
and weather conditions exclude or restrict
pollinator activity. For example, a bee
pollinated plant growing in a habitat which
received rain every day was put in class
three because rain decreases pollinator
activity which in turn decreases fecundity
(Cruden, 1972). A hummingbird flowered
plant growing in the same habitat was put
in habitat four, because hummingbird
activity is unaffected by rain.
RESULTS
Pollen-Ovule Ratios and Breeding Sys-
tem.-Using the outcrossing index as a
relative measure of outcrossing, it is clear
that P / O's increase with the likelihood of
cross-pollination (Table 1; Appendices 1.
and 2.). The difference in P/ O's between
groups (Table 1) is significant (F s =
204.4; df = 4,91; P < .001) and the
increase in P /0 from group to group ap-
pears to be logarithmic (r =
.947; P <
.01) rather than linear (r =
.538; P <
.01) .
34 ROBERT WILLIAM CRUDEN

TABLE 2. Comparison of reproductive characteristics of species in Appendices I and II (for which

data is available).

Outcrossing Index 3 4

Number species in class 5 20 36 25

Self-compatible 5/5 11/11 18/18 4/8
Bud autogamy 4/5 5/11 0/18 0/8
Stigma exposed after selfing 1/4 2/5
Stigma exposed before selfing 1/5 3/6 12/12 4/4
Selfing but mechanism unknown 0 3 6 0
Pollinator required 0/5 0/10 4/18 7'/9
Produce nectar 15/16 21/25'
Protogynous 6/7 73/22
Protandrous 1/7 12/22
'Does not include Pectls pringlei which is partially autogamous, pollinator required for maximum seed set. (Keil,
pers, comm.)
'Those that did not produce nectar were bee flowers that are strictly pollen flowers.
3 Includes Anemone patens, Ranunculus [ascicularis, Anemonella thalictroides which are early spring species; Crypto-
taenia canadensis, a woods species.

Plants with the same OCI share sets of
morphological and behavioral characteris-
tics that delimit particular breeding sys-
tems (Table 2). Flowers with an OCI
o are cleistogamous. Plants whose flowers
have an OCI = 1 are self-compatible,
obligately autogamous weeds. An OCI 2
characterizes plants that are self-compati-
ble and autogamous, but which probably
outcross to a limited degree. This group
includes native colonizers as well as many
weeds. Plants with an OCI =
3 are self-
compatible, and if dichogamous, tend to be
protogynous. Most of the species studied
= produce nectar and some require a pollina-
tor. An OCI =
4 characterizes species that
are adapted for outcrossing and a few of
= those studied are self-incompatible. Dichog-
amy is common, particularly protandry, and
most of the species for which there is data
require a pollinator.
PIO's of Plants With "Sweepstakes"

TABLE 3. Pollen-ovule ratios of plants with "Sweepstakes" reproduction

# PolIen grains in a
Pollen-Ovule pollinium or polyad
ratio
Taxon Sonrce x ± S.E. Ovules per carpel
Asclepias'
verticillata L. Dickinson Co., Iowa 10 10.1 ± 0.4 1.99
oenotheroides Cham. & Schlecht. Durango, Mexico 6 7.5 ± 0.3 1.48
subverticillata (Gray) Vail Durango, Mexico 10 10.8 ± 0.6 2.13
Sarcostema'
clausum (Jacq.) Roem. & Schult. J aIisco,Mexico 10 7.4 ± 0.4 1.46
Matellea'
chrysantha (Greenm.) Woods. Morelos, Mexico 10 4.9 ± 0.4 0.97
pilosa (Benth.) Woods. Michoacan, Mexico 8 5.7 ± 0.2 1.14
pavonia (Decaisne) Woods. J aIisco, Mexico 10 3.8 ± 0.4 0.73
Calliandra'
palmeri S. Wats. J aIisco,Mexico 11 863.3 ± 42.3 0.71
Nayarit, Mexico 11 828.3 ± 37.2 0.79
callistemon (Schlect.) Benth. J aIisco,Mexico 14 526.5 ± 59.0 1.18
kunthii Benth. Michoacan, Mexico 15 328.7 ± 36.6 0.95
anomala (Kunth.) Macbr. Mexico, Mexico 12 246.1 ± 24.5 0.92
, Asclepiadaceae
2 Mimosaceae
 BREEDING SYSTEMS IN PLANTS 35

TABLE 4. Pollen-ovule ratios of some grosses from Johnson County, Iowa.

Species Habitat N= x ± S.E.

Digitaria sanguinalis (L.) Scop. roadside 13 1234.0 ± 19.5

Echinochloa crusgalli (L.) Beauv. roadside 15 1267.1 ± 38.0
Eleusine indica (L.) Gaertn. greenhouse 15 1111.1 ± 58.1
Eragrostis cilianensis (All.) Link. roadside 11 498.0 ± 37.1
Eragrostis pectinacea (Michx.) Hitch. roadside 5 302.4 ± 3.9
Glyceria striata (Lam.) Hitch. 1 woods 15 389.0 ± 7.9
Setaria faberii Herrm. parking lot 10 1212.0 ± 41.3
Setaria glauca (L.) Beauv. roadside 14 754.7 ± 42.1
Trisetum spicatum (L.) Richt.' tundra 8 1839.3 ± 123.4
1 Collected in Emmet Co., Iowa.
• Collected in Clear Creek Co., Colo. ele. ca. 3650 m.

Reproduction.-Plants in Asclepiadaceae latter species, in a small number of cases

and Mimosaceae have low P/ a's compared
to other xenogamous species (Table 3). The
pollen grains are incorporated in pollinia
or polyads, thus the important comparison
is not the P/ a but the ratio of pollen
grains in a pollinium or polyad to the num-
ber of ovules per carpel. This ratio in the
Asclepiadaceae studied varied from 0.73 to
2.13. In Calliandra the number of ovules
was usually greater than the eight pollen
grains in a polyad.
PIa's of Grasses.-The PIa's of nine
grasses (Table 4) are much smaller than
those reported for various wind pollinated
species (Pohl, 1937). Self-pollination oc-
curs in the buds of Echinochloa crusgalli,
Eragrostis cilianensis, E. pectinacea, and
Trisetum spicatum, although the anthers
and stigmas may subsequently become ex-
serted. The anthers and stigmas of Setaria
glauca and Digitaria sanguinalis are ex-
serted simultaneously from the tip of the
floret and dehiscence occurs just prior to
or concomitantly with exsertion. In the
the anthers and stigmas remained within
the flowers.
Pollen-Ovule Ratios and H abitat.-
There is a significant difference between
the P / a's of plants that grow in different
habitats (Table 5; F s = 65.11; df =
3,82; P < .001). Plants that were col-
lected in highly disturbed habitats are
obligately or facultatively autogamous with
OC/'s = 1 or 2. Plants in early succes-
sional stages have OC/'s = 2 or 3. All are
self-compatible and most are autogamous.
Plants that occur in advanced successional
stages but flower when pollinators are not
reliable are self-compatible, usually produce
nectar, and may require a pollinator. If
dichogamous, the species are usually
protogynous, including those xenogamous
species that flower in early spring (Table
2) . In contrast, all the species from
habitats with reliable pollinators produce
nectar, require a pollinator, and are fre-
quently protandrous.
Pollen Grains Required for Pollination.-

TABLE 5. Correlations between pollen-ovule ratios and successional stage.

Number PIO log PlO

Successional stage species x of x ± S.E. x of x ±S.E.

Highly disturbed 23 135.6 ± 23.5 1.93 ± .09

Early successional 24 588.7 ± 100.3 2.65 ± .07
Late successional 23 1877.4 ± 423.6 3.09 ± .09
Pollinators unpredictable
La te successional 15 7251.5 ± 1396.1 3.72 ± .10
Pollinators predictable
36 ROBERT WILLIAM CRUDEN
TABLE 6. Seed set as a function of pollen grains
per stigma in Mirabilis jalapa.
Pollen grains/Stigma' 2 3 4 5 6 >6
Flowers pollinated 39 55 49 52 45 21 15
% Seed set 49 67 73 87 76 86 100
1 Only one ovule per flower.
Two sets of data indicate that several
pollen grains per ovule must reach a stigma
to assure maximum seed set. In Mirabilis
jalapa four pollen grains per ovule are
required to produce the level of seed set
that results from self-pollination, i.e., ap-
proximately 90% (Table 6). A comparison
of seed set in cleistogamous flowers from
two populations of Viola nephrophylla sug-
gests that approximately six pollen grains
are necessary per seed. The PIO's of the
two populations (6.1 and 5.4) are essen-
.tially equal (P > .4). However' seed set
III the two populations (x = 97% and
84%, respectively) is significantly differ-
ent (t = 5.3; P < .001) as is the number
of viable pollen grains per ovule (6.1 ±
S.E. 0.4 and 4.8 ± S.E. 0.4 respectively;
t = 2.18; P < .05). The smaller number
of viable pollen grains is due 1) to flowers
with fewer pollen grains and 2) flowers
with aborted pollen grains.
Intraspecific Variation.-Pollen-ovule ra-
tios for two or more populations of several
species were determined and in most cases
the differences do not appear to be bio-
logically significant, e.g., Viola nephrop-
hylla (Appendix 1A), Veronica arvensis
(Appendix 1B), Verbena bracteata (Ap-
pendix 1C), Pectis prostrate, Circaea
canadensis, Polygonum virginianum, Ver-
bena stricta and Tribulus terrestris (Ap-
pendix 1D), Pectis multiflosculosa (Appen-
TABLE 7. Breeding system characteristics and fecundity in Hedeoma hispida.
Source GCI PIG
Dickinson Co., Kan. 1 39
Dickinson Co., Iowa 2 64.2
Bernalillo Co., N.M. 2 109.9
McLennon Co., Texas 3 244.7
dix lE), Calliandra palmeri (Table 5) and
four species of M irabilis (Appendix 2).
Dramatic intraspecific variation occurred
in Glecoma hederacea, Hedeoma hispida,
Stellaria media, and Draba reptans. In
Hedeoma (Table 7) the differences in
length and width of the corolla were cor-
related with the relative position of the
anthers and stigmas. In the Kansas plants
the receptive stigma lay between the
anthers, whereas in the Iowa and New
Mexico plants the stigmas extended be-
yond but curled back into the anthers. In
the Texas population the stigmas extend
beyond the anthers, which they barely
reached when curled back. The Texas
population was reproductively unsuccessful
in a disturbed habitat compared to the
other populations (Table 7) and un-
doubtedly required a pollinator.
Similarly, the PI O'S of the two popula-
tions of Stellaria media reflect flower size.
The flowers of the Kansas population (PI 0
= 22.6) had very small petals that were
included within the sepals. In contrast,
the petals of the Iowa flowers (PI 0 =
76.1) extended beyond the sepals and pro-
vided a target for a potential pollinator.
Casual observations in other populations
suggest that the two populations studied
represent parts of a continuum with re-
spect to petal size and corolla diameter.
Individuals of Draba reptans had
cleistogamous flowers with no petals or
flowers with noticeable petals, although
petal size was variable. Flowers with
petals had PIO's from 4 to 85 (x = 30.4
± S.E. 8.8, N = 11). Mature flowers
could not be used to determine PI O'S
hence it was impossible to correlate PI 0
with petal length. Petal size and Pl O's
Length of Width of
flower/calyx coroUa mm Fecundity
1:1 1-1.5 98
1.2: 1 2-3 97
1.5:1 2.5-3.5 91
1.8: 1 3-4 31
 BREEDING SYSTEMS IN PLANTS
varied within individuals as well as be-
tween individuals. In one individual, a
flower with petals had a Pia of 33.2 and
a flower with no detectable petals, a Pia
of 5.2.
DISCUSSION
The inescapable conclusion from this
survey is that pollen-ovule ratios are cor-
related with the breeding systems of plants.
Using an outcrossing index, populations
were assigned to one of five classes (Table
1). The breeding systems of the taxa in
each class are more similar to each other,
than to a typical taxon in another class.
Although the number of species studied in
each class is relatively small (Table 2)
the trends are clear. Xenogamous species
are primarily outcrossers, protandrous or
self-incompatible, and most require a pol-
linator. Exceptions to this rule are pri-
marily self-compatible, protogynous species
that flower when and/or where pollinators
are unreliable. In contrast, facultatively
xenogamous species are regularly self-com-
patible and protogynous or homogamous.
Although some species require a pollinator,
most self-pollinate when the flowers close.
Even though selfing may occur it is clear
that the adaptations favor xenogamy and
that selfing occurs in the absence of, or in
addition to outcrossing. The relative im-
portance of autogamy and xenogamy is
reversed in facultatively autogamous
plants. Flowers of facultatively autoga-
mous plants tend to self-pollinate prior to
or during flower opening and the exposure
of receptive stigmas to potential pollinators
occurs after self-pollination has occurred.
The opportunity for outcrossing in obli-
gately autogamous species is very low.
From class to class there is a distinct even
if subtle change in the likelihood of out-
crossing, accompanied by a significant
decrease in Pia's. The evolutionary shift
in breeding systems from xenogamy to
cleistogamy is thus accompanied at each
step by a large decrease in Pla's.
The results of other studies are consistent
with mine, including studies on Eupatori-
37
um (Baker, 1967), Leavenworthia (Lloyd,
1965), Limnanthes (Arroyo, 1973), Mirab-
ilis (Cruden, 1973; Pilz, 1974), and
Senecio (Gibbs et al., 1975). In particular
the studies on Eupatorium and Senecio
show that decreases in P/ a's go hand in
hand with the switch from xenogamy to
autogamy and that intermediate Pia's are
correlated with intermediate breeding sys-
tems. In the above taxa the derived
breeding systems exhibit shifts toward
autogamy. In Cotula (Lloyd, 1972) the
derived breeding systems exhibit shifts
toward greater xenogamy. All the species
tested were self-compatible, including
gynomonoecious and monoecious species.
The gynomonoecious species produced
fewer pollen grains per floret than
monecious species, which in turn produced
fewer pollen grains per floret than related
dioecious and subdioecious taxa.
I examined a series of grasses to test
the patterns described above. The results
show clearly that the Pia's of anemophi-
lous plants and their autogamous relatives
also are correlated with their breeding
systems. Rye (Secale cereale), an obligate
outcrosser, has a Pia of approximately
57,310 (Pohl, 1937) . Wheat (Triticum
aestivum) is primarily autogamous or
geitonogamous and has Pia's that range
from 1200 to 12,000 (Vries, 1971, 1974).
The varieties with the lowest PIa's are
autogamous, as are a number of weedy
species with similar PI a's (Table 4), and
the number of pollen grains shed outside
the flowers increases with the number
produced (Beri and Anand, 1971), i.e., as
the Pia increases so does the likelihood of
outcrossing. Kannenberg and Allard
( 1967) report that cleistogamous flowers
of Festuca microstachys complex produce
but one tiny anther which contains "half
a dozen or so pollen grains" and is en-
tangled in the stigmatic hairs. Only the
examination of many more species will per-
mit the discrimination of major groups as
can be done with zoophilous species and
their relatives.
The Pia's of plants in Asclepiadaceae
38 ROBERT WILLIAM CRUDEN
and Mimosaceae are exceptions in the
xenogamous group. In these groups the
pollen grains are grouped in pollinia or
polyads and the PI O'S (Table 3) are
quite low for xenogamous plants. How-
ever, if pollination is successful, sufficient
pollen grains are deposited to fertilize
most, though rarely all of the ovules in a
carpel. The parallel between the re-
productive biology of these groups and the
gambling lottery, i.e., minimal energetic
investment in each pollen packet, low
reproductive return on each pollinia or
polyad, but large reproductive return when
pollination is successful, suggests the name
"sweepstakes" reproduction.
Pollen-Ovule Ratios, Breeding Systems,
and Successional Stage.-A comparison of
PlO's with successional stage (Table 5)
shows that PI O'S increase significantly
from early to late successional seres. In ad-
dition to corroborating the notion that
autogamy is adaptive in disturbed habitats
and xenogamy in advanced successional
stages (Baker, 1959, 1974; Stebbins,
1958), the data show there is a spec-
trum of breeding systems and that
obligate autogamy and obligate xenogamy
represent the extremes of that spectrum.
Between the extremes the successful breed-
ing systems display a balance between
autogamy and xenogamy.
The balance between autogamy and
xenogamy in intermediate habitats is
easily seen in a comparison of breeding
systems in early successional and late suc-
cessional stages with unreliable pollinators.
Plants of early successional stages (Table
5) are primarily facultatively autogamous
or facultatively xenogamous. All of the
species for which I have data (N = 12)
are self-compatible and are regularly
autogamous although some have adapta-
tions that favor xenogamy, for example,
protandry in Malva neglecta and Hous-
tonia minima, and nectar production in
Verbena bracteata. Both M. neglecta
(Mulligan, 1972) and V. bracteata
(Cruden, unpubl.) are occasionally visited
by small bees, which may mediate some
outcrossing. Facultatively autogamous
species may also occur in more advanced
successional stages, particularly in arid
habitats (Arroyo, 1973; Moore and Lewis,
1965) and/or habitats in which pollinators
are not reliable (Hagerup, 1950, 1951;
Kevan, 1972 ; Lloyd, 1965; Mosquin,
1966) . Two grasses, Trisetum spicatum
and Glyceria striata, grow in habitats in
which wind is an unreliable pollen vector
and T. spicatum at least is facultatively
autogamous.
Xenogamous and facultatively xenoga-
mous species from habitats with ephemeral
(Beattie, 1971) or unreliable pollinators,
although self-compatible, regularly require
pollinators. The emphasis is on outcross-
ing although many species can and do
self-pollinate. Estimates of outcrossing in
facultatively xenogamous plants vary
greatly, from 5-10% in homostylous
Primula vulgaris (Crosby, 1959) to 70%
in Cheiranthus cheiri L. (Bateman, 1956).
The extremes of variation in this group
are well illustrated by Anthericum nanum
(PIO = 271) in which all the ovules
develop into seeds even if pollinators are
excluded from the flowers and Adenocaulon
bicolor (PI 0 = 884) which requires a
pollinator. The potential for both xenog-
amy and autogamy in facultatively
xenogamous species is exemplified by
Mirabilis nyctaginea. Flowers are visited
in late afternoon and early morning by
bees and at night by moths. Those flowers
that remain unpollinated may self-pollinate
as the flowers close (Cruden, 1973).
In many self-compatible and/or autog-
amous plants, successful outcrossing
must result from the ability of "foreign"
pollen tubes to grow faster than "self"
pollen tubes, e.g., in Cheiranthus cheiri
(Bateman, 1956) . This is particularly
true if selfing precedes exposure of the
stigmas to foreign pollen, e.g., in Bidens
lemmonii (Cruden, unpubl.) . In some
species the differential in growth rates be-
tween foreign and self pollen tubes is
considerable. For example, in Iris tenax
self pollen tubes reach the ovary in 50 hrs,
 BREEDING SYSTEMS IN PLANTS
whereas foreign pollen tubes reach the
ovary in 40 hrs (Smith and Clarkson,
1956).
Plants with cleistogamous flowers are
not associated with a particular habitat,
and many species produce both cleistog-
amous and chasmogamous flowers. The
cleistogamous flowers of some species are
produced when pollinators of the chasmog-
amous flowers are presumably absent
from the habitat. For example, in Viola,
cleistogamous flowers are initiated sub-
sequent to canopy closure, which effectively
excludes many bees from the habitat. The
cleistogamous flowers of Triodanis per-
foliata are produced in late spring prior to
regular bee activity.
How Many Pollen Grains are Suffi-
cient? . -The answer to this question
establishes the minimum PIa, that PIa
below which fecundity is reduced. With
the exception of species with cleistogamous
flowers and those with "sweepstakes"
reproduction, effective PIa's should be in
excess of the minimum. The available
data suggest that two to seven pollen
grains per ovule on a stigma are sufficient
to maximize seed set. In Mirabilis jalapa
four pollen grains produce 87 percent 'Seed
set and in Viola nephrophylla six viable
pollen grains result in 97 percent seed set.
Cleistogamous flowers of Triodanis lepto-
carpa set 98 percent seed (Cruden, un-
publ.) with 3.3 pollen grains per ovule.
These figures are similar to the pollen
grain to seed ratios of 2.0: 1 and 4.8: 1 for
the pin and thrum flowers of J epsonia
parryi (Ornduff, 1970) and 1.3: 1 and
5.2: 1 for the pin and thrum flowers of J.
heterandra (Ornduff, 1971). Seed set in
J. heterandra was less than 85% which
suggests 1.3 pollen grains is not sufficient
to achieve maximum seed set. Silander
and Primack (pers. comm.) found that
seed set did not increase if more than
three pollen grains per seed were placed
on the stigmas of Oenothera jruticosa.
Finally, the pollen per pollinia/ovules per
ovary ratios of Asclepiadaceae and
Calliandra are not sufficient to give per-
39
fect seed set but give high seed set
(Cruden, 1976; unpubl.). These observa-
tions suggest that 2-6 pollen grains per
ovule must reach each stigma to assure
relatively high seed set, but not necessarily
perfect seed set.
The necessity of more than a single
pollen grain per ovule reaching a stigma is
explained, at least in part, by Brewbaker
and Majumder (1961). These workers
demonstrated that in hanging drop cultures
the percentage of pollen grains germinating
is directly proportional to the number of
pollen grains. Jennings and Topham
(1971) experimentally reduced the amount
of viable pollen by diluting it with talc or
dead pollen. As the percent of viable pol-
len applied to stigmas was reduced through
dilution, the percent of viable pollen grains
that germinated also decreased and fecun-
dity decreased. Brewbaker and Majumder
(1961) attribute the differences in pollen
grain germination to compounds produced
by the pollen grains that diffuse into the
stigmatic or stylar tissue and promote
pollen grain germination and pollen tube
growth. A second reason for the necessity
of several pollen grains per ovule, is that
some pollen tubes never reach the ovary,
e.g., in Phaseolus vulgaris (Weinstein,
1926).
Are PI a's Minimal? .-If slight differ-
ences in fecundity affect fitness then Pia's
should reflect a trade-off between being
energetically conservative by minimizing
pollen production and providing sufficient
excess pollen to assure maximum fecundity.
The comparison of PIa's and seed set in
the cleistogamous flowers from two popula-
tions of Viola nephrophylla suggests that
in cleistogamous flowers PIa's are mini-
mal. A reduction of viable pollen from 6.1
to 4.8 pollen grains was correlated with a
significant decrease in seed set from 97 to
84 percent. In wheat, seed set was con-
siderably less in plants whose flowers were
cross-pollinated compared to self-pollinat-
ing controls. Pollen grain to seed ratios
of 4.4:1 produced only 51.8% seed set in
the wind pollinated plants compared to
40 ROBERT WILLIAM CRUDEN
87.7% in controls (Sorokina and Laptev,
in Vries, 1971). It is clear that in the
variety tested the PI 0 was not sufficient
for wind pollination although large enough
to achieve considerable outcrossing. The
low fecundity of the chasmogamous flowers
of Mimulus guttatus (Vickery, 1959;
Lindsay and Vickery, 1967) may well have
been a consequence of extremely low
PIO's (range 2.5 to 25) rather than in-
efficient pollinators. Although these re-
sults do not permit a conclusion as to
whether or not PI O'S are minimal, they
do suggest that less than minimal PI O'S
result in decreased fecundity.
Pollen-Ovule Ratios as Indicators of
Breeding Systems.-Careful study of floral
behavior shows that the PI O'S of several
species are better indicators of their breed-
ing system than their OCI. The 6-7 mm
wide flowers and protandry of Houstonia
minima indicate xenogamy. The PIO of
54 suggests regular autogamy. Examina-
tion of preserved material showed that
style elongation carries the stigma to the
level of the dehisced anthers, which still
contain pollen and lie within the narrow
floral tube. Autogamy is inevitable.
In contrast, the flowers of Amorpho
canescens and Abronia maritima are
smaller than those usually associated with
xenogamous plants. However, the flowers
of Amorpha canescens are protogynous and
set no seed if pollinators are excluded. The
large PIO (16,807) suggests that the
plants are obligate outcrossers. Abronia
maritima (PIO = 2520) is self-incompat-
ible (Tillet, 1967), hence xenogamous.
Although the flowers of both species are
small, they are aggregated in showy in-
florescences which attract potential pol-
linators.
The Pl O's of the various Mirabilis
species indicate facultative xenogamy
whereas the OCl's indicate xenogamy. The
morphology and behavior of the flowers of
M. jalapa (PIO = 382) and M. multiflora
(PIO = 595-910) are virtually the same
except for flower closure. In the former
both the style and stamens curl within the
floral tube, thus self-pollination may occur
during flower closure (Cruden, 1973),
whereas in M. multiflora the style remains
outside the floral tube during closure, thus
precluding autogamy (Cruden, 1970). The
former is facultatively xenogamous where-
as the latter, being self-incompatible
(Baker, 1964), is obligately xenogamous.
The PI O's of these species reflect basic
differences in their breeding systems.
SUMMARY
Pollen-ovule ratios (PI O's) of flowering
plants, including grasses, reflect their
breeding system and these can be divided
into five classes: xenogamy, facultative
xenogamy, facultative autogamy, obligate
autogamy and cleistogamy. The evolu-
tionary shift from class to class is accom-
panied by a significant decrease in the
mean p) O. This pattern is found in
anemophilous plants and their relatives as
well as zoophilous plants and their rela-
tives. Plants in Asclepiadaceae and
Mimosaceae are exceptions in the xenog-
amous group. These plants invest mini-
mal energy in pollen production and their
fecundity is low, but when pollination is
successful the reproductive return is
relatively high.
A comparison of PI O's with successional
stage shows that PI O'S increase signifi-
cantly from disturbed habitats to late
successional seres. In addition to suggest-
ing that autogamy is adaptive in disturbed
habitats and xenogamy in advanced suc-
cessional stages, the data show that in
intermediate habitats PIO's, hence breed-
ing systems, also are intermediate, and
display a balance between autogamy and
xenogamy.
PIO's may be minimal and if PI O's
fall below a certain minimum, fecundity
decreases. This is probably a consequence
of insufficient pollen grains reaching the
stigmas. Several examples are discussed
showing that PI O's are a better indicator
of a plant's breeding system than floral
size and morphology.
 BREEDING SYSTEMS IN PLANTS
ACKNOWLEDGMENTS
The technical assistance of Harold
Brotzman, Nancy Crocker, Mary Ann
Reihman, Vicky Kinnamon, Sharon Kins-
man and Sharon Hermann-Parker is
greatly appreciated. Bud collections of
various species were contributed by Robert
Ballard, Charles Hart, Thomas Melchert,
George Pilz and William Platt. David
Keil identified and provided information
on the reproductive biology of Pectis. Jane
Bock, Suzanne Clark, Thomas Duncan
and Elizabeth Ramsey provided informa-
tion on the reproductive biology of various
other species. Rogers McVaugh and Lorin
I. Nevling, Jr., provided identifications of
various species. The constructive criticism
of Herbert Baker, Nels Lersten, William
Platt and an anonymous reviewer of an
earlier draft of the manuscript are greatly
appreciated as are the comments of S.
Hermann-Parker on the present draft.
LITERATURE CITED
ARROYO, M. T. K. 1973. Chiasma frequency
evidence on the evolution of autogamy in
Limnanihes floccosa (Limnanthaceae). Evo-
lution 27:679-688.
BAKER, H. G. 1955. Self-compatibility and es-
tablishment after "long-distance" dispersal.
Evolution 9:347-348.
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speciation in flowering plants. Cold Spr.
Harb. Symp. Quant. BioI. 24:177-190.
- - . 1964. Variation in style length in rela-
tion to out-breeding in Mirabilis (Nyctagina-
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- - . 1967. The evolution of weedy taxa in
the Eupatorium microstemon species aggregate.
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- - . 1974. The evolution of weeds. Ann. Rev.
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BATEMAN, A. J. 1956. Cryptic self-incompati-
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Heredity 10:257-261.
BAWA, K. S. 1974. Breeding systems of tree
species of a lowland tropical community. Evo-
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BAWA, K. S., AND P. A. OPLER. 1975. Dioecism
in tropical forest trees. Evolution 29: 167-
179.
BEATTIE, A. J. 1971. Itinerant pollinators in a
forest. Madrofio 21: 12<J-.124.
BERI, S. M., AND S. C. ANAND. 1971. Factors
41
affecting pollen shedding capacity of wheat.
Euphytica 20:327-332.
BOCK, J. H., AND S. J. PETERSON. 1975. Re-
productive biology of Pulsatilla patens (Ra-
nunculaceae). Am. MidI. Natur. 94:476-478.
BREWBAKER, J. L., AND S. K. MAJUMDER. 1961.
Cultural studies of the pollen population ef-
fect and the self-incompatibility inhibition.
Amer. J. Bot. 48:457-464.
CROSBY, J. L. 1959. Outcrossing in homostylous
primroses. Heredity 13:127-131.
CRUDEN, R. W. 1970. Hawkmoth pollination of
Mirabilis (Nyctaginaceae). Bull. Torr. Bot.
Club. 97:89-91.
- - . 1972. Pollinators in high elevation eco-
systems: The relative effectiveness of birds
and bees. Science 176: 1439-1440.
- - . 1973. Reproductive biology of weedy and
cultivated Mirabilis (Nyctaginaceae). Amer.
J. Bot. 60:802-809.
- - . 1976. Fecundity as a function of nectar
production and pollen-ovule ratios. pp. 171-
178. In Variation, Breeding and Conservation
of Tropical Forest Trees. J. Burley and B. T.
Styles (eds.). Academic Press.
GIBBS, P. E., C. MILNE, AND M. CARRILLO.
1975. Correlation between the breeding sys-
tem and recombination index in five species
of Senecio. New Phytol. 75:619-626.
HAGERUP, O. 1950. Rain pollination. Dan. Vi-
densk. Sels. BioI. Medd. 18, nr5, 19 p.
- - . 1951. Pollination in the Faroes-in spite
of rain and poverty in insects. Dan. Vidensk.
Sels. BioI. Medd. 18, nr15, 48 p.
JENNINGS, D. L., AND P. B. TOPHAM. 1971.
Some consequences of raspberry pollen dilu-
tion for its germination and for fruit develop-
ment. New Phytol. 70:371-380.
KANNENBERG, L. W., AND R. W. ALLARD. 1967.
Population studies in predominantly self-pol-
linated species. VIII. Genetic variability in
the Festuca microstachys complex. Evolution
21: 227-240.
KEVAN, P. G. 1972. Insect pollination of high
Arctic flowers. J. Ecol. 60:831-848.
KNUTH, P. 1908. Handbook of Flower Polli-
nation. Vols. II and III. Trans. J. R. Ains-
worth Davis. Clarendon Press, Oxford.
LINDSAY, D. W., AND R. K. VICKERY, JR. 1967.
Comparative evolution in Mimulus guttatus
(Scrophulariaceae). Evolution 21:439-456.
LLOYD, D. G. 1965. Evolution of self-compati-
bility and racial differentiation in Leaven-
worthia (Cruciferae). Contr. Gray Herb. Har-
vard. 195:3-133.
- - . 1972. Breeding systems in Cotula L.
(Compositae, Anthemideae). I. The array of
monoclinous and diclinous systems. New Phy-
tol. 71: 1181-1194. "
MOORE, D. M., AND H. LEWIS. 1965. The evo-
42 ROBERT WILLIAM CRUDEN

lution of self-pollination in Clarkia xanthiana.
Evolution 19: 104-114.
MOSQUIN, T. 1966. Reproductive specialization
as a factor in the evolution of the Canadian
Flora. pp. 43-65. In The Evolution of Can-
ada's Flora. R. L. Taylor and R. A. Ludwig
(eds.) , Univ. Toronto Press.
MULLIGAN, G. A. 1972. Autogamy, allogamy
and pollination in some Canadian weeds.
Can. J. Bot. 50:1767-1771.
ORNDUFF, R. 1969. Reproductive biology in re-
lation to systematics. Taxon 18: 121-133.
- - . 1970. Incompatibility and the pollen econ-
omy of Jepsonia parryi. Amer. J. Bot. 57:
1036-1041.
1971. The reproductive system of Jep-
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PIJL, L. VAN DER. 1937. Disharmony between
Asiatic flower-birds and American bird-flowers.
Annl. Jard. Bot. Buitenzorg 48:17-26.
PILZ, G. 1974. Systematics of Mirabilis sub-
genus Quamoclidion. Ph.D. Thesis-Botany,
University California, Berkeley.
POHL, F. 1937. Die Pollenerzeugung der Wind-
bluter. Beih. Bot. Centralbl. 56:365-470.
SMITH, F. H., AND Q. D. CLARKSON. 1956. Cyto-
logical studies of interspecific hybridization
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43: 582-588.
STEBBINS, G. L. 1950. Variation and Evolution
in Plants. Columbia Univ. Press. New York
643 p.
- - . 1958. Longevity, habitat, and release of
genetic variability in the higher plants. Cold.
Spr, Harb. Symp, Quant. BioI. 23 :365-378.
TILLET, S. S. 1967. The maritime species of
Abronia (Nyctaginaceae). Brittonia 19:299-
327.
VICKERY, R. K., JR. 1959. Barriers to gene ex-
change within Mimulus guttatus (Scrophu-
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VRIES, A. PH. DE. 1971. Flowering biology of
wheat, particularly in view of hybrid seed
production-a review. Euphytica 20: 152-170.
1974. Some aspects of cross-pollination
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Euphytica 23:11-19.
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Phaseolus vulgaris. Amer. J. Bot. 13:248-263.

ApPENDIX 1. Pollen-ovule ratios of some angiosperms.

Habitat Breeding
Species and family Location and habitat class' system> PlO (x + S.E.)
A. Class 1 (OCI =
0) Cleistogamous
Triodanis perfoliata (L.) Nieuwl. Dickinson Co., Kansas 2.7 ±.3
Campanulaceae Roadside park
Triodanis leptocarpa
(Nutt.) Nieuwl. Dickinson Co., Kansas 3.3 ±.2
Campanulaceae Roadside park
Draba reptans (Lam.) Fern. Cedar Co., Iowa 3.7 ±.2
Cruciferae Sand prairie
Veronica peregrina L. Johnson Co., Iowa 6.7 ±.4
Scrophulariaceae Mowed lawn
Viola nephrophylla Greene Johnson Co., Iowa 6.1 ±.4
Violaceae Flower gardens
5.4 ±.3

B. Class 2 (OCI =
1) Autogamous-obligate
M ollugo verticillata L. Johnson Co., Iowa SC,A 38.9 ± 3.1
Aizoaceae Garden
Stellaria media (L.) Cyrille Douglas Co., Kansas 1 SC,A 22.6 ± 1.3
Caryophyllaceae Mowed lawn
Cardamine hirsuta L. Johnson Co., Iowa SC,A 22.2 ± 1.0
Cruciferae Greenhouse weed
Hedeoma hispida Pursh Dickinson Co., Kansas SC,A 39.0 ± 3.0
Labiatae Mowed lawn, roadside park
Veronica arvensis L. Dickinson Co., Kansas 18.1 ± 0.5
Scrophulariaceae Mowed lawn
Johnson Co., Iowa SC,A 25.0 ± 1.5
Mowed lawn
Douglas Co., Kansas 1 28.3 ± 1.5
Mowed lawn
 BREEDING SYSTEMS IN PLANTS 43

ApPENDIX 1. Continued.

Habitat Breeding
Species and family Location and habitat class' system b PlO (x + S.E.)
C. Class 3 (OCI =
2) Autogamous-facultative
\'
Microsteris humilis
(Doug!.) Greene Pueblo Co., Colo. 99.1 ± 3.4
Boraginaceae Gravel along highway
Heliotropium sp. Sinaloa, Mexico 134.5 ±3.5
Boraginaceae Camping area, trailer park
Stellaria media (L.) CyriIIe Dickinson Co., Iowa SC,Ae 76.1 ± 4.8
Caryophyllaceae Disturbed area next to
dormitory
Bidens lemmonii Gray Chihuahua, Mexico (4) SC,Ak 113.7
Compositae Plants reared in greenhouse
Conyza ramosissimum Cron. Johnson Co., Iowa SC,A 190.4 ± 9.4
Compositae Open ground by parking lot
Matricaria matricariodes
(Less.) Porter Johnson Co., Iowa SC,A 154.0 ± 4.6
Compositae Gravel area along road
Camelina microcarpa Andrz. Boulder Co., Colo. 287.2 ± 3.9
Cruciferae Mowed weedy area by field
Descurainia richardsonii Boulder Co., Colo. 275.4 ± 11.6
(Sweet) O. E. Schulz Mowed roadside
Cruciferae
Glecoma hederacea L. Douglas Co., Kansas 143.6 ± 16.1
Labiatae Mowed lawn
Hedeoma hispida Pursh Dickinson Co., Iowa SC,A 64.2 ± 3.6
Labiatae Mowed lawn
Torrance Co., New Mexico 2 109.8 ± 4.9
Dry, grazed pine woods
Malva neglecta Wallr." Torrance Co., New Mexico 2 SC,Aef 226.4 ± 4.7
Malvaceae Weedy area by parking lot
Sida ciliaris L. Sinaloa, Mexico 133.7
Malvaceae Camping area, trailer park
Androsace occidentalis Pursh Johnson Co., Iowa 111.1 ± 3.8
Primulaceae Mowed lawns
Houstonia minima Beck" Cedar Co., Iowa 3 Pr,SC,A 54.1 ± 11.4
Rubiaceae Opening on sand prairie
Chaenorrhinum minus (L.) Lange Johnson Co., Iowa 2 SC,A 344.4 ± 17.6
Scrophulariaceae Occasionally mowed lawn
Collinsia parviflora Doug!. Pueblo Co., Colo. 2 396.9 ± 13.7
Scrophulariaceae Along trail in pine woods
Veronica serpyllifolia L. Pueblo Co., Colo. (7) 1 SC,Aer 31.9 ± 1.4
Scrophulariaceae Along a stream
Verbena bracteata Lag. & Rodr. Dickinson Co., Iowa 2 SC,A 201.9 ± 11.0
Verbenaceae Abandoned driveway
Johnson Co., Iowa 221.0 ± 9.3
Mowed lawn .

D. Class 4 (OCI =
3) Xenogamous-facultative
Lap pula echinata Bilib. Dickinson Co., Iowa 2 1767.7 ± 112.4
Boraginaceae Abandoned farm yard
Lappula redowskii
(Hornem.) Greene Lincoln Co., Colo. (5) 2 1894.0
Boraginaceae Unmowed roadside
Adenocaulon bicolor Hook. Humboldt Co., Calif. 3 SC,Pg,P 884.7 ± 92.6
Compositae Redwood forest
Adenocaulon himalaicum Edgew. Japan 3 SC,Pg,P 925.6 ± 45.9
Compositae Forest or woods
44 ROBERT WILLIAM CRUDEN

ApPENDIX 1. Continued.

Habi tat Breeding

Species and family Location and habitat class» system> PlO (x + S.E.)
Bidens leptocephala Sherff. Arizona SC,Ak 988.6 ± 42.6
Compositae
Pectis prostrate Cav. Sinaloa, Mexico SC,Ag 285.7 ± 9.1
Compositae Parking area, trailer park
Durango, Mexico 2 SC,Ag 257.8 ± 6.4
Roadside ditch near road
Tagetes /ili/olia Lag. Nayarit, Mexico 2 355.9 ± 29.4
Compositae Edge of path
Quamoclit ct. pinnata J aIisco, Mexico 2 SC",P 160.7 ± 4.9
Convolvulaceae Uncut roadside
Capsella bursa-past oris (L.) Medic. Johnson Co., Iowa 2 SC,A f 344.2 ± 21.4
Cruciferae Weedy road between fields
Thlaspi arvense L. Johnson Co., Iowa 2 SC,Af 755.6 ± 34.3
Cruciferae Weedy road between fields
Ellisia nyctelea L. Pueblo Co., Colo. 2 1470.5 ± 82.9
Hydrophyllaceae Unmowed roadside
Hedeoma hispida Pursh. McLennon Co., Texas 2 244.7 ± 8.8
Labiatae Roadside park
Lamium amplexicaule L. Douglas Co., Kansas 2 SC,Ae 517.3 ± 27.9
Labiatae Mowed lawn
Lepechinia spicaia WiIId. Mexico, Mexico 3 710.4 ± 28.1
Labiatae Pine forest
M oldavica parvi/lora (Nutt.) Britt. Dickinson Co., Iowa 2 965.0 ± 45.9
Labiatae Abandoned farm yard
Monarda /~tulosa L. Dickinson Co., Iowa 4 Pr 825.1±21.2
Labiatae Virgin prairie
Scutellaria parvula Michx. Dickinson Co., Iowa 3 1011.6 ± 47.4
Labiatae Virgin prairie
A nthericum nanum Baker Mexico SC,A 271.0 ± 26.0
Liliaceae Grazed areas
Kosteletzkya paniculata Benth. J alisco, Mexico 2 583.5 ± 32.0
Malvaceae Unmowed roadside
Circaea canadensis HilI. Dickinson Co., Iowa (5) 3 880.5
Onagraceae Oakwoods
Linn Co., Iowa (7) 3 835.3 ± 22.4
Oak-hickory forest
Phryma leptostachya L. Dickinson Co., Iowa 3 SC,A 2558.6 ± 22.9
Phrymaceae Oakwoods
Polygonum virginianum L. Johnson Co., Iowa 3 439.4 ± 16.1
Polygonaceae Small woods
Cedar Co., Iowa 3 382.2 ± 12.9
Deciduous woods
Primula incana Jones Park Co., Colo. (1) 2 SC,AJ 245.0
Primulaceae Wet meadow
Ranunculus abortivus L. Johnson ce., Iowa 3 SC,Pg 980.0 ± 94.2
Ranunculaceae Woods on flood plain
Ranunculus longirostrus Godr. Dickinson Co., Iowa 3 SC,Pg,P 1071.6 ± 28.2
Ranunculaceae J emerson Slough
Galium boreale L. Dickinson Co., Iowa 3 1902.4 ± 45.0
Rubiaceae Sand prairie
Mitella diphylla L. Dubuque Co., Iowa 3 1280.0 ± 212.5
Saxifragaceae Deciduous forest
Phyla nodi/lora (L.) Greene Muscatine Co., Iowa 3 733.3 ± 35.6
Verbenaceae Marsh
Verbena simplex Lehm, Muscatine Co., Iowa 2 327.0 ± 12.8
Verbenaceae Sandy areas near Little Lake
 BREEDING SYSTEMS IN PLANTS 45

ApPENDIX 1. Continued.

Species and family
Verbena strict a Vent.
Verbenaceae
Verbena wrightii Gray
Verbenaceae
Tribulus terrestris L.
Zygophyllaceae
Habitat Breeding
Location and habitat class- system b PlO (x + S.E.)
Cedar Co., Iowa 2 535.4 ± 19.0
Disturbed area along road
Dickinson Co., Iowa 2 SC,A 796.6 ± 31.8
10 yr old grassland
McLennon Co., Texas 2 782.0 ± 25.4
Edge of roadside park
Torrance Co., New Mexico 2 574.7 ± 20.7
Weedy area near parking lot
Sinaloa, Mexico (1) Pg 377.1
Camping area, trailer park
Sedgwick Co., Kansas (4) 340.3
Parking lot

E. Class 5 (DCI = 4) Xenogamous

Peciis multiflosculosa
(DC) Sch-Bip. Guerrero, Mexico (1) Pr 3973
Compositae Sand dunes
Sinaloa, Mexico 3581 ± 118
Sand dunes
Pectis pringlei Fernald Durango, Mexico (6) 3 SC,±A",Pg 2651 ± 180
Compositae Open ground in desert
Geranium maculatum L. Johnson Co., Iowa 4 Pr 1062 ± 59
Geraniaceae Sand prairie
Glecoma hederacea L. Johnson Co., Iowa 3 Pr 2108 ± 67
Labiatae Unmowed lawn
Monarda citriodora Cerv. Chihuahua, Mexico 3 Pr 1684 ± 58
Labiatae Juniper woodland
Salvia cardinalis Kunth Mexico, Mexico (2) 4 Pr 19525
Labiatae Pine woods
Salvia chiapensis Fernald Chiapas, Mexico (5) 4 Pr 11687 ± 399
Labiatae Pine woods
Satureja mexicana (Benth.) Brig. Oaxaca, Mexico (5) 4 Pr 1685 ± 93
Labiatae Chaparral
Stachys coccinea Jacq. Durango, Mexico (3) 4 Pr 5870 ± 148
Labiatae Pine woods
Stachys eriantha Benth. Mexico, Mexico (3) 4 Pr 3538 ± 96
Labiatae Zacatone, above tree line
Stachys palustris L. Dickinson Co., Iowa 4 Pr 6182 ± 211
Labiatae Slough on upland prairie
Amorpha canescens Pursh'' Dickinson Co., Iowa 4 Pg,P 16807 ± 750
Leguminoseae Upland prairie
Abronia maritima Nutt. ex Wats." Sinaloa, Mexico SI,P 2520 ± 144
Nyctaginaceae Sand dunes
Abronia umbellata Lam. Monterey Co., Calif. (5) 4 SI,P 7912 ± 462
Nyctaginaceae Coastal bluffs
Sanguinaria canadensis L. Cedar Co., Iowa 3 3242 ± 313
Papaveraceae Oak forest
Aneomone canadensis L. Dickinson Co., Iowa 4 Pg,P 9895± 440
Ranunculaceae Upland prairie
Anemone patens L. Dickinson Co., Iowa (2) 3 SC',Pg 3492
Ranunculaceae Upland prairie
Anemonella thalictroides (L.) Spach Cedar Co., Iowa 3 Pg 10053 ± 426
Ranunculaceae Sand prairie
Ranunculus fascicularis Muhl. Cedar Co., Iowa 3 SC,Pg 3049 ± 207
Ranunculaceae Sand prairie
Ranunculus repens L. Johnson Co., Iowa 4 SI,Pr,P 5791 ± 183
Ranunculaceae Church yard
46 ROBERT WILLIAM CRUDEN

ApPENDIX 1. Continued.

Habitat Breeding
Species and family Location and habitat class' system. P /0 (x + S.E.)
Ceanothus thrysijlorus Esch. Monterey Co., Calif. 4 Pr 7994 ± 485
Rhamnaceae Hills above ocean
Cryptotaenia canadensis (L.) DC. Dickinson Co., Iowa 3 SC,Pg,A 2180 ± 83
Umbelliferae Oak woods
Zizia ourea (L.) Koch Dickinson Co., Iowa 4 Pg 4957 ± 338
Umbelliferae Upland prairie
Tribulus cistoides L. Tabasco, Mexico (1) 4 Pr 5043
Zygophyllaceae Open area near beach
• I-highly disturbed; 2-early successional; 3-late successional (pollinators unreliable); 4-late successional (pol-
Iinators reliable).
• SC-Self-compatible; SI-Self-incompatible; A-autogamous; P-Pollinator necessary; pg-Protogynous; Pr-e-Pro-
tandrous.
CFigures indicate number of individuals examined. Where no figure is shown the mean is based on 9 or more indi-
viduals. 40 of tbe 9S means are based on 14 or more individuals.
d Outcrossing Index = 3. The reasons for inclusion in Class 3 or S are discussed in the text.
• Knuth, 1905.
f Mulligan, 1972.
• Keil, pers, comm,
• Pijl, 1937.
J Bock, pers, comm.
k Hart, pers. comm,
1 Bock & Peterson, 1975.

ApPENDIX 2. Pollen-ovule ratios of Mirabilis.

Breeding
Species Source Pollinator System N= PlO (x ± S.E.)

M. multiflora Texas hawkmoths and XC 15 910'

(including hummingbirds
M. [orebelli)
New Mexico 15 900'
Utah 15 595'
California 20 685'
Calif. & Ariz. 8 800·
M. jalapa Coahuila, Mex. hawkmoths FX 15 382.4 ± 11.4
J alisco, Mex. hawkmoths FX 15 383.0 ± 14.7
M. grandiflora Durango, Mex. hawkmoths FX 15 298.0 ± 7.5
M. triilora Baja Calif. Sur. hummingbirds 15 646.0 ± 18.6
J alisco, Mex. 15 618.0'
M. hirsuta Dickinson Co., moths FX 17 368.8 ± 19.0
Iowa bud autogamy FA 13 118.5 ± 6.6
M. nyctaginea Dickinson Co., moths and bees FX 15 559.0 ± 17.3
Iowa
Johnson Co., Ia. bud autogamy FA 20 186.6 ± 13.9
Cedar Co" Ia. bud autogamy FA 22 156.5 ± 5.2
• Pilz, 1974.
• Cruden, 1973.
CX = Xenogamy; FX = Facultative Xenogamy; FA = Facultative Autogamy.
NOTE ADDED IN PROOF: Since the revision of the manuscript two papers have come to my
attention: Murray, B. G., 1974, Heredity 33:285-292 and Lloyd, R. M., 1974, Bot. J. Linn. Soc.
69:23-35. Murray provides evidence that grasses with high PIO's are outcrossers and those with
low P10's are seIfers. Lloyd presents evidence from ferns that seIfers occur in pioneer habitats
and outcrossers in climax communities.