Research Article

Seed dormancy and germination vary within and among

species of milkweeds

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Thomas N. Kaye*, Isaac J. Sandlin and Matt A. Bahm
Institute for Applied Ecology, 563 SW Jefferson Avenue, Corvallis, OR 97333, USA
Received: 8 November 2017  Editorial decision: 12 February 2018  Accepted: 27 February 2018  Published: 2 March 2018
Associate Editor: Gabriela Auge
Citation: Kaye TN, Sandlin IJ, Bahm MA. 2018. Seed dormancy and germination vary within and among species of milkweeds. AoB
PLANTS 10: ply018; doi: 10.1093/aobpla/ply018

Abstract.  Pollinators in general and monarch butterflies in particular are in decline due to habitat loss. Efforts to
restore habitats for insects that rely on specific plant groups as larvae or adults depend on the ability of practition-
ers to grow and produce these plants. Monarch larvae feed exclusively on milkweed species, primarily in the genus
Asclepias, making propagation and restoration of these plants crucial for habitat restoration. Seed germination pro-
tocols for milkweeds are not well established, in part due to the large number of milkweed species and conflicting
reports of seed dormancy in the genus. We tested for seed dormancy and the optimum period of cold stratification
in 15 populations of A. speciosa and 1–2 populations of five additional species, including A. asperula, A. fascicularis,
A. subulata, A. subverticillata and A. syriaca. We exposed seeds to cold (5 °C) moist conditions for 0, 2, 4, 6 and 8
weeks and then moved them to 15 °C/25 °C alternating temperatures. In A. speciosa, dormancy was detected in
eight populations, and this dormancy was broken by 2–4 weeks of cold stratification. The remaining seven popula-
tions showed no dormancy. Seed dormancy was also detected in two populations of A. fascicularis (broken by 4–6
weeks of cold stratification) and a single population of A. syriaca (broken by 2 weeks of cold stratification). No dor-
mancy was detected in A. asperula, A. subulata or A. subverticillata. Seed dormancy appears to be widespread in the
genus (confirmed in 15 species) but can vary between populations even within the same species. Variation in seed
dormancy and cold stratification requirements within and among Asclepias species suggests local adaptation and
maternal environments may drive seedling ecology, and that growers should watch for low germination and use
cold stratification as needed to maximize seed germination and retain genetic variability in restored populations.

Keywords: Cold stratification; conservation biology; habitat restoration; monarch butterfly; plant propagation; pol-
linator conservation.

Introduction may have serious impacts on ecologic function and eco-

nomic stability (Potts et al. 2010) because pollinators are
Declines in insect pollinator abundance and diversity
important or essential for 65 % of wild flowering plants
have been documented over the past decade on regional
(Ollerton et al. 2011) and 75 % of domestic food crops
and global scales (Ghazoul 2005; Steffan-Dewenter
(Klein et  al. 2007). Monarch butterflies (Danaus plexip-
et al. 2005; Biesmeijer et al. 2006; Williams and Osborne
pus) in particular have declined up to 90 % over the past
2009; Cameron et al. 2011). The loss of these pollinators

*Corresponding author’s e-mail address: [email protected]

© The Author(s) 2018. Published by Oxford University Press on behalf of the Annals of Botany Company.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/
licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

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 Kaye et al. – Seed dormancy and germination in milkweeds

15 years (Jepson et al. 2015), due in part to loss of milk- suggested germination protocols for Asclepias species
weed in their summer breeding habitat from increased are inconsistent or even contradictory (Borland 1987).
use of herbicides in intensive farming practices (Brower For example, cold stratification has been recommended
et al. 2012). to release seeds of A. speciosa from dormancy but some
Plants in the milkweed family (Asclepiadaceae) are practitioners report that no cold stratification is neces-
the exclusive food source for monarch butterfly larvae. sary for large-scale propagation (Stevens 2000).
Monarchs in North America feed on at least 27 species Here we present the results of research on seed dor-
in the genus Asclepias and a few closely related genera mancy and germination in six Asclepias species. In par-
(Malcolm and Brower 1986). Milkweed populations have ticular, we examine variation in dormancy and cold
suffered considerable declines in habitat in the central stratification requirements among populations of A. spe-

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USA due mainly to the use of genetically modified, gly- ciosa as well as between five other species. We test the
phosate resistant crops, and the placement of an add- hypothesis that dormancy varies among populations of
itional 10 million hectares of herbicide-tolerant corn A.  speciosa from across the species’ geographic range,
into production since 2007 (Pleasants and Oberhauser and populations of the five additional species. We pre-
2012). Restoring milkweed habitat is crucial to the dict that dormancy varies substantially, thus explaining
recovery of the monarch butterfly and will require plant- conflicting reports of the importance of cold stratifica-
ing large numbers of milkweeds throughout its geo- tion in this genus. We also compile and review the pri-
graphic range (Fallon et  al. 2015; Jepsen et  al. 2015; mary literature and unpublished reports on dormancy
Luna and Dumroese 2013). Milkweeds also benefit many and germination of several additional milkweed species.
other insects and pollinators, such as milkweed beetles
in the Cerambycidae (Evans 2014), bumblebees (Bombus
Methods
spp.), honey bees (Apis mellifera), and other bees and
lepidoptera of various body sizes (e.g. Fishbein and
Venable 1996), so conservation of these plants has cas- Germination and viability tests
cading positive effects on ecosystem function and mul- We obtained seed from 15 populations of A.  speciosa
tiple groups of organisms (Buckley and Nabhan 2016; (Fig.  1) and one or two populations of five additional
Dumroese et al. 2016). North American species, including A.  asperula, A.  fas-
To produce milkweeds for habitat conservation at a cicularis, A.  subulata, A.  subverticillata and A.  syriaca
large scale, reliable propagation protocols must be avail- (Table  1) from multiple locations in the USA (Fig.  2).
able. One method of milkweed propagation is by vegeta- These seeds had either been collected during the sum-
tive rhizome cutting, which can be an effective method mer immediately prior to germination testing, or had
for growing and establishing several species of milk- been in dry, cold (<0 °C) storage at the US Forest Service
weeds in gardens and habitat restoration sites (Ecker and Bend Seed Extractory or the US Agricultural Research
Barzilay 1993; Landis 2014; Landis and Dumroese 2014). Service, National Plant Germplasm System.
Although cuttings of this type can produce large plants To determine if seeds from each species and source
relatively quickly, the process generally results in clones population required a period of cold stratification to
of fewer genotypes, and thus lower genetic diversity and release them from dormancy, we exposed seeds to a
fitness in restored populations, than propagation from range of cold, moist periods. Dormancy was defined
seed (Williams 2001). Germination is the first step in here as the inability of a seed to germinate in a specified
producing plants from seed and requires an understand- period of time under combinations of normal physical
ing of dormancy and germination requirements for effi- factors (e.g. temperature, light, etc.) that are otherwise
cient and successful plant propagation at a large scale. favourable for germination, following Baskin and Baskin
But seed germination protocols for Asclepias species are (2004). Seeds were stratified for 0, 2, 4, 6 or 8 weeks at
relatively poorly developed, despite the large number of 5 °C at the Oregon State University Seed Lab. We used
species in the genus. Over 100 species of Asclepias are four or five replicates (depending on seed availabil-
known to occur in North and Central America (Mabberley ity, Table 1) of 50 seeds from each seed source. For all
1997) with ~76 species in the USA and Canada (USDA, replicates, seeds were placed on moistened germina-
NRCS 2017). Related species, and populations within tion paper in 15 cm × 15 cm × 3 cm transparent plastic
species, are often assumed to have the same germin- boxes with fitted lids. The paper was moistened weekly
ation requirements because of phylogenetic constraints as needed with distilled water. After stratification treat-
(Seglias et  al. 2018). It is generally unknown if seed ments were applied, seeds were placed in a growth
dormancy and cold stratification requirements vary chamber with 15  °C/25  °C alternating temperatures,
across Asclepias species or populations. In some cases, with 8 h of darkness at 15 °C and 16 h of fluorescent light

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Figure 1.  Asclepias speciosa (A) in flower and (B) dehiscing fruit with seeds. Plant photographed in Willamette Valley, OR.

at 25 °C. We defined germination as emergence of the trials (not shown) we found that 95  % or more seeds
radicle at least 3  mm, and counted germinated seeds germinated within this period of time, and any addi-
after 10  days (Baskin and Baskin 2001). In preliminary tional germination took many weeks to complete. Seed

Table 1.  Asclepias species and populations included in dormancy and germination tests, with seed sources, samples (50 seeds each) per
treatment and years in storage. †BSE indicates the US Forest Service, Bend Seed Extractory; GRIN indicates the US National Germplasm
Resources Information Network.

Species Population Seed source† (accession number) Samples per treatment Years in storage

Asclepias speciosa Denver, CO BSE (CO932-306-Jefferson-12) 4 5

Montrose, CO Colorado Plateau Native Plant Program 4 2
Minidoka, ID US Fish and Wildlife Service 5 0
Treasure Valley, ID Idaho State University 5 0
Navajo Dam, NM BSE (NM930N-86-SanJuan-12) 4 5
Vernal, NM BSE (SOS-NM930N-12-10) 4 7
Galice, OR BSE (SOS-OR110-904-Josephine-15) 4 2
Malheur, OR US Fish and Wildlife Service 5 0
Ontario, OR Institute for Applied Ecology 5 0
Tub Springs, OR BSE (OR110-637-Jackson-13) 4 4
Willamette Valley, OR Heritage Seedlings, Inc. 5 0
Escalante, UT BSE (UT030-256-Garfield-15) 4 2
Maeser, UT BSE (SOS-UT080-140-UINTAH-13) 4 4
Little Pend Oreille, WA US Fish and Wildlife Service 5 0
Kane, WY BSE (SOS-WY020-11-11) 4 10
Asclepias asperula Santa Cruz, AZ GRIN (W6-48232) 4 4
Asclepias Agate Reservoir, OR (1) BSE (SOS-OR110-608-Jackson-13) 4 4
fascicularis Gold Hill, OR (2) BSE (SOS-OR110-968-Jackson-15) 4 2
Asclepias subulata Lake Havasu City, CA GRIN (W6-46777) 4 5
Asclepias Anvil Points, CO BSE (SOS-CO932-160-08) 4 9
subverticillata UT GRIN (W6-36792) 4 10
Asclepias syriaca NJ GRIN (W6-48817) 4 4

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Figure 2.  Location in the USA of each Asclepias source population included in this study. The arrow indicates a population of A. syriaca from
New Jersey.

viability was tested with tetrazolium (TZ) by the Oregon A.  speciosa, with substantial improvement in germin-
State University Seed Lab using standard procedures ation after stratification for eight out of the 15 popu-
(Elias et  al. 2012). The TZ test was used to estimate lations tested (grouped for ease of visual inspection in
the percentage of live and dead seeds in each seed Fig. 3A), but only slight or no dormancy in the remain-
source, regardless of dormancy level (Baskin and Baskin ing seven populations (Fig.  3B). Seed viability was over
2001). Seeds were cut longitudinally to expose interior 90 % in most A. speciosa populations as tested with TZ,
tissues and facilitate the entrance of TZ solution, and with the exception of two populations with lower viabil-
incubated in a 1 % TZ solution for 24 h at 35 °C. Seeds ity, Willamette Valley, OR (73 ± 6.0 %) and Malheur, OR
were inspected for TZ staining, specifically the pattern (28 ± 6.1 %). In populations with substantial dormancy,
and intensity of red colour in live tissues in seeds. Seed <60 % of seeds germinated without cold stratification,
with uniform stain colour were recorded as viable seeds. but with 2 or more weeks of cold stratification germin-
Tetrazolium tests were performed with samples of 157 ation increased to the levels of seed viability, or nearly
to 210 seeds per source population. so. For example, without cold stratification 32  ±  5  %
of seeds from Little Pend Oreille, WA, germinated, but
Analysis after 2 weeks of cold stratification germination rose to
We tested for effects of cold stratification and popula- 91  ±  5  %, close to the viability estimate of 96  ±  2.6  %
tion with a general linear model of analysis of variance, (Fig.  3A). Seed from Malheur, OR, required 4 weeks of
with cold stratification treatment as a fixed effect and cold stratification to fully break dormancy, while ger-
seed source as a random effect using NCSS statistical mination of seeds from Willamette Valley, OR, did not
software (Hintze 2008). Separate analyses were per- germinated to the level of their estimated viability.
formed for Asclepias speciosa populations, and the five Populations with largely non-dormant seeds (Fig.  3B)
additional Asclepias species and their populations. Data germinated to very high rates at or near their viability
are reported with the mean ± 95 % confidence interval. estimate without any cold stratification. Seed germin-
Confidence intervals for the viability estimates were cal- ation was not higher after 8 weeks of cold stratification
culated from the normal approximation of the binomial than 6 weeks in any population tested, so this treatment
distribution using the proportion of seeds that were con- is not shown (Fig. 3).
sidered viable, the proportion of seeds that were consid-
ered non-viable and the total number of seeds tested. Interspecific differences in dormancy and
germination
Dormancy and germination varied among species of
Results
Asclepias examined here. Again there was a signifi-
cant interaction between cold stratification treatment
Intraspecific population differences in dormancy and population for the five taxa examined (F  =  40.29,
and germination of A. speciosa df  =  24;105, P  <  0.00001), with two species showing
There was a significant stratification treatment by popu- positive effects on germination of cold stratification of
lation interaction (F = 17.62, df = 56;255, P < 0.0001) for 2 or more weeks, but the remaining taxa possessing

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Figure 3.  Seed germination in response to different periods of cold stratification for 15 populations of Asclepias speciosa (A) dormant seeds
and (B) non-dormant seeds. Seed viability as estimated with TZ is shown with horizontal hatched lines. Error bars represent 95 % confidence
intervals.

little or no dormancy. Without cold stratification, both Discussion

populations of Asclepias fascicularis that were sampled
had germination of only 17 ± 5.1 % and 35 ± 5.1 %, des-
Seed dormancy and viability
pite viability estimates of 93  ±  3.5  % and 90  ±  4.2  %,
respectively, indicating substantial dormancy. Both of We found that seed dormancy varied widely within and
these populations required 6 weeks of cold stratifica- among Asclepias species, and that the period of time in
tion to achieve maximum germination of 80 ± 5.1 % and cold stratification needed to break dormancy varied as
82.5  ±  5.1  %, respectively (Fig.  4). Eight weeks of cold well. In A. speciosa in particular, seeds from seven source
stratification (not shown) did not improve seed ger- populations had essentially no dormancy, while seeds
mination any further in A. fascicularis. Asclepias syriaca from eight other populations generally needed 2 weeks
also had partial dormancy, with only 38.5  % of seeds (but up to 6 weeks) of cold stratification to reach germi-
germinating without cold stratification, despite viability nation levels similar to viability. Of the remaining species
estimated at 95 ± 3.0 %. After 2 weeks of cold stratifica- tested here, the seeds of three (A. asperula, A. subulata
tion of this sample, germination increased to 92 ± 5.1 % and A.  subverticillata) were non-dormant, while two
(Fig. 4). The remaining species, A. asperula, A. subulata (A. fascicularis and A. syriaca) required cold stratification
and A. subverticiallata, germinated to levels close to (or to break dormancy. Seed germination tests in A. subu-
within 95  % confidence intervals) of their seed viabil- lata have also found no dormancy in populations from
ity as estimated by TZ even without cold stratification. California (Everett 2012). Previous reports have sug-
Again, cold stratification for 8 weeks (not shown) did gested little or no dormancy in A. speciosa (Bartow 2006;
not increase seed germination over the 6-week period Skinner 2008), but our results indicate that the benefits
(Fig. 4). of cold stratification differ substantially among source

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Figure 4.  Seed germination in response to difference periods of cold stratification for five Asclepias species from the USA. Seed viability as
estimated with TZ is shown with horizontal hatched lines. Error bars represent 95 % confidence intervals.

populations. Even in populations where seed germina- Cold stratification

tion was enhanced by cold stratification, many seeds Baskin and Baskin (2001) suggest that Asclepias seeds
germinated without a cold treatment, suggesting that may have physiological dormancy, which is typically bro-
when dormancy was present, it typically affected only ken by cold stratification, seed coat removal or chem-
a portion of the seeds from a given location. This was ical inducements, all of which improve germination in
true when dormancy was present in A. speciosa as well A.  syriaca (Oegema and Fletcher 1972). Our review of
as A. fascicularis and A. syriaca, which had 3 % to 54 % published and unpublished accounts of seed germin-
germination even without cold stratification. ation and dormancy in 17 species of Asclepias suggests
Seed viability was typically very high (over 85 %) in our that variation in the period of cold stratification needed
samples, with two exceptions out of the 22 populations to break dormancy appears to be common within and
we tested. High seed viability has also been reported among milkweed species (Table 3). For example, Green
in A.  cordifolia (Barner 2009) and A.  syriaca (Bhowmik and Curtis (1950) examined five species of milkweed
1978). Unpublished records of seed viability and dor- from Wisconsin and found a range of periods of cold
mancy in Asclepias species tested by the US Department stratification needed to completely release seeds from
of Agriculture National Laboratory for Genetic Resources dormancy, from none at all to 5 months. Most research-
Preservation are compiled in Table 2. Their records from ers use cold stratification temperatures of 4  °C to 5  °C
germination tests of small seed samples (typically two (Luna and Dumroese 2013), although some placed
replicates of 50 seeds, but in one case only 44 seeds) seeds outdoors in winter to expose seeds to ambient
with specific periods of cold stratification ranging from cold conditions (e.g. Green and Curtis 1950).
0 to 21 days suggest wide variation in viability and dor- Species with non-dormant seeds (i.e. in which no
mancy within and among populations of nine Asclepias benefit of cold stratification has been reported) include
species. For example, among eight populations of A. fas- A.  erosa (Everett 2012), A.  meadii (Bowles et  al. 1998),
cicularis, seed viability ranged from 68 to 98 %, and seven A.  perennis (Edwards et  al. 1994), A.  speciosa (Bartow
of the source populations possessed no seed dormancy 2006; Skinner 2008), A. syriaca (Radivojevic et al. 2016)
(defined here as <80 % germination of viable seeds) when and A.  tuberosa (Green and Curtis 1950; Phillips 1985).
no cold stratification was provided. This is in contrast to However, in some of these same species cold stratifi-
our findings of substantial dormancy in both populations cation has been found to improve germination in stud-
of A. fascicularis that we examined. In A. speciosa, seed ies from other populations, and the optimal period of
viability ranged from 65 to 98 % among 13 populations, cold stratification varies as well. For example, A. syriaca
and six out of nine populations were dormant without and A.  tuberosa are important for monarch butterfly
cold stratification (Table 2), a pattern that agrees with our populations (Seiber et al. 1986; Borders and Lee-Mader
observation of wide variation in dormancy in that species. 2014) and have received the most attention for their

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Table 2.  Summary of seed dormancy in Asclepias species tested at the USDA/ARS National Laboratory for Genetic Resources Preservation,
Seed Quality Lab, with duration of cold stratification (if any), number of populations sampled and number of populations with and without
dormancy. Germination is relative to viability, and is shown as the amount or range determined for each period of stratification. Populations
were classified as possessing seed dormancy if <80 % of viable seeds germinated. Each population represents a separate seed collection.

Species Stratification No. of populations Viability Populations without dormancy Populations with dormancy
sampled
No. of populations Germination No. of populations Germination

A. asperula 7 days 1 100 % 1 100 % 0 –

A. fascicularis 0 day 8 68–98 % 7 92–100 % 1 37 %
A. hirtella 0 day 1 100 % 0 – 1 33 %

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A. latifolia 0 day 1 100 % 1 100 % 0 –
A. speciosa 0 day 9 73–98 % 3 89–100 % 6 8–78 %
7 days 4 65–95 % 4 93–100 % 0 –
14 days 1 98 % 1 100 % 0 –
A. subulata 0 day 1 100 % 1 100 % 0 –
A. subverticillata 0 day 2 92–96 % 2 100 % 0 –
14 days 1 96 % 1 100 % 0 –
A. syriaca 0 day 8 92–100 % 0 – 8 10–67 %
A. tuberosa 0 day 3 96–100 % 1 88 % 2 50–72 %
7 days 1 55 % 1 93 % 0 –
14 days 1 98 % 0 – 1 50 %
21 days 3 50–76 % 3 100 % 0 –

germination requirements, and both vary widely in individual mother plants (Andersson and Milberg 1998).
cold stratification needs. Asclepias syriaca populations Seed dormancy can affect interactions within and
with seed dormancy may require 56  days or more of among species by determining the seasonal timing of
cold stratification (Green and Curtis 1950; Oegema and germination, seedbank dynamics, and exposure of seeds
Fletcher 1972), or as little as 7 days (Evetts and Burnside and seedlings to hazards and competition for resources
1972; Farmer et al. 1986). We found 2 weeks of cold strati- (Harper 1977; Baskin and Baskin 2001). Dormancy has
fication was sufficient to release dormancy in this spe- been shown to be under genetic control in some spe-
cies, but we did not try a shorter period. In A. tuberosa, cies, often in response to natural selection, such as in
as much as 90–120 days of cold stratification (Salac and Digitaria (Hacker 1984), Arabidopsis (Alonso-Blanco et al.
Hesse 1975; Cullina 2000; Blessman and Flood 2001) or 2003; Bentsink et  al. 2010) and Oryza (Gu et  al. 2004),
as little as 21 days (AOSA 2016) may be needed for opti- or it can result from conditions during seed maturation
mal germination of dormant seed lots. Similarly, 30 days in the environment of the maternal plant and zygote
(Vandevender and Lester 2014a) down to 1 week or less (Bodrone et al. 2017; Penfield and MacGregor 2017), or
(Lincoln 1983; Schultz et al. 2001a) of cold stratification both (Postma and Agren 2015). To our knowledge, nei-
may be needed to improve germination in A. incarnata. ther genetic nor environmental controls on dormancy
Among species tested by the National Laboratory for have been documented in Asclepias. Despite the wide-
Genetic Resources Preservation, most populations that spread presence of seed dormancy in Asclepias species,
received cold stratification of 7–21  days were released persistent seed banks have not been detected even
from dormancy (Table 2). when milkweeds are present in the above-ground vege-
tation (Smith and Kadlec 1983; Johnson and Anderson
Factors that affect dormancy and germination in 1986). Seedling emergence of A.  syriaca exceeds 80  %
Asclepias for seeds buried 0.5–4  cm (Yenish et  al. 1996) leaving
Seed dormancy can vary among species, popula- only a small proportion of seeds in the soil unaccounted
tions (Keith and Myerscough 2016; Siles et  al. 2017), for and which could contribute to a seedbank or suc-
collections from the same population but different cumb to mortality. Germination of Asclepias seeds
years (Green and Curtis 1950; Kaye 1999) and among can also be affected by light, scarification, substrate,

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 Kaye et al. – Seed dormancy and germination in milkweeds

Table  3.  Duration of cold stratification needed to break dormancy in Asclepias species from published sources, with post stratification
temperatures used or recommended for germination, where reported.

Species Period of cold stratification needed Germination temperature References

to break dormancy

Asclepias amplexicaulis 60 days Heon and Larsen (1999)

Asclepias erosa 0 day Everett (2012)
Asclepias exaltata 0–60 days Heon and Larsen (1999)
Asclepias floridiana 120 days 18–21 °C Green and Curtis (1950)
Asclepias incarnata 0–60 days Heon and Larsen (1999)

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5 days 27 °C/16 °C Lincoln (1983)
7 days 18 °C Schultz et al. (2001a)
30 days Vandevender and Lester (2014a)
90 days Cullina (2000)
Asclepias hirtella 0–60 days Heon and Larsen (1999)
Asclepias meadii 0 day Bowles et al. (1998)
70 days Betz (1989)
Asclepias ovalifolia 120 days Green and Curtis (1950)
Asclepias perennis 0 day 10–30 °C Edwards et al. (1994)
Asclepias purpurascens 0–60 days Heon and Larsen (1999)
Asclepias speciosa 0 day 21 °C/10 °C Bartow (2006)
0 day Skinner (2008)
Asclepias subulata 0 day Everett (2012)
Asclepias sullivantii 0–60 days Heon and Larsen (1999)
90–120 days 21–27 °C/18–24 °C Blessman et al. (2002)
150 days Green and Curtis (1950)
Asclepias syriaca 0 day 26 °C/21 °C Radivojevic et al. (2016)
0–60 days Heon and Larsen (1999)
7 days 20 °C/30 °C Evetts and Burnside (1972)
7 days 20 °C/30 °C Farmer et al. (1986)
14–21 days 30 °C/15 °C Baskin and Baskin (1977)
21 days 25 °C Jeffery and Robison (1971)
30 days 18 °C Schultz et al. (2001b)
30 days 20 °C/30 °C Lincoln (1976)
56 days 26 °C Oegema and Fletcher (1972)
60 days Green and Curtis (1950)
Asclepias tuberosa 0 day Phillips (1985)
0 day Green and Curtis (1950)
0–60 days Heon and Larsen (1999)
21 days 30 °C/10 °C AOSA (2016)
30 days Vandevender and Lester (2014b)
60 days Bir (1986)
60–90 days Grabowski (1996)
71 days Nichols (1934)

(Continued)

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 Kaye et al. – Seed dormancy and germination in milkweeds

Table 3.  Continued

Species Period of cold stratification needed Germination temperature References

to break dormancy

90–120 days 21–27 °C/18– Blessman and Flood (2001)

24 °C
105 days 33 °C/19 °C or 26 °C Salac and Hesse (1975)
Asclepias verticillata 0–60 days Heon and Larsen (1999)
30 days Green and Curtis (1950)

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Asclepias viridiflora 0–60 days Heon and Larsen (1999)

temperatures, after-ripening and other factors. Light Baskin and Baskin (1977) suggest that with after-ripen-
may be required for germination in some Asclepias spe- ing A.  syriaca seeds can germinate at lower tempera-
cies, such as A. incarnata (Lincoln 1976; Deno 1993), or tures and after shorter periods of cold stratification, and
have no effect on others, including A. tuberosa (Mitchell Bhowmik (1978) found that germination increased grad-
1926; Deno 1993). Light has little or no effect on ger- ually with time in storage from 1 to 11  months. Seed
mination of A.  syriaca once adequate (≥2 weeks) cold dormancy in A.  speciosa populations included in our
stratification has been provided (Baskin and Baskin experiments may have been affected by the amount of
1977; Lincoln 1983). Mechanical scarification of the time seeds were in storage as well as local environmen-
seed coat improves germination in A. syriaca (Oegema tal conditions. We used fresh seeds that had been col-
and Fletcher 1972; Evetts and Burnside 1972). Smoke lected in the same year of the experiment and had been
treatments that mimic wildfire smoke can improve stored for ~3 months, as well as seeds that had been in
germination of A.  syriaca as well (Mojzes and Kalapos dry, cold (<0  °C) storage for up to 6  years. Fresh seeds
2015), but burning and soil disturbance reduced seed- from six populations had >40 % dormancy, while seeds
ling emergence in A.  meadii (Roels 2013). Experiments stored for 2, 5 and 6  years had <15  % dormancy. But
with temperature and substrate have found that A. syri- seeds stored for 4 years from two populations had dor-
aca germinates best under alternating temperatures of mancy levels similar to fresh seeds, suggesting that dor-
20 °C/30 °C on clay or clay mixed with peat (Farmer et al. mancy can persist even in stored seeds and may involve
1986), or 21 °C/26 °C on sand or loam (Radivojevic et al. other factors beyond storage. Further confounding the
2016). Temperatures that promote successful germin- effects of storage time was that most of our fresh seeds
ation after cold stratification of Asclepias also vary con- came from latitudes north of 40° and all of the stored
siderably, and most reports recommend an alternating seeds came from farther south, and no seed dormancy
temperature regime. For example, a temperature cycle was found in the southern populations; in other words,
of 10 °C/30 °C is effective for A. perennis (Edwards et al. most non-dormant seeds had been stored and were
1994) and the standard for seed testing in A. tuberosa from southern latitudes. Therefore, it is not possible for
(AOSA 2016). Alternating 16 °C/27 °C (Lincoln 1983) has us to separate the effects of storage time from local
been successfully used for A.  incarnata, and cycles of environments, but we speculate that both seed storage
15  °C/30  °C (Baskin and Baskin 1977) and 20  °C/30  °C and local conditions (Seglias et  al. 2018) could influ-
(Evetts and Burnside 1972; Lincoln 1976; Farmer et  al. ence seed dormancy in A.  speciosa and possibly other
1986) have been recommended for A. syriaca. Bhowmik Asclepias species.
found germination in A. syriaca increased with increas-
ing temperature from 10 °C to 27 °C. We used alternat-
ing 15 °C/25 °C in our trials and this cycle was generally Conclusions
effective for germination of the species and populations Growers of Asclepias species should consider cold strati-
we examined. In the case of the A. speciosa population fication to break seed dormancy when it is encountered.
from Malheur, OR, we also used a cycle of 20 °C/30 °C but We found widespread evidence of seed dormancy in
found germination to be reduced by about 30  % com- the genus through germination tests reported here as
pared to the 15 °C/25 °C cycle (data not shown). well as published and unpublished reports. Seed dor-
After-ripening, a period of time after seed dispersal in mancy has been detected in at least some populations
which changes in the seeds affect their ability to germin- of 15 species in the genus, including A. amplexicaulis,
ate, can affect some species of Asclepias. For example, A.  exaltata, A.  fascicularis, A.  floridiana, A.  incarnata,

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 Kaye et al. – Seed dormancy and germination in milkweeds

A.  hirtella, A.  meadii, A.  ovalifolia, A.  purpurascens, Management Seeds of Success Program; A.  Pilmanis
A. speciosa, A. sullivantii, A. syriaca, A. tuberosa, A. ver- and R.  Hosna from the Bureau of Land Management;
ticillata and A.  viridiflora (Table  3). Cold stratification D. Perkins from College of Idaho; L. Boyer from Heritage
is a relatively simple method to break this dormancy Seedlings, Inc.; K.  See from the Uncompahgre Native
and increase seed germination substantially, and can Plant Program; and K.  Herriman from the US Forest
shorten germination time (Salac and Hesse 1975) and Service Bend Seed Extractory. M.  Cashman of the US
even out differences in germination across populations Department of Agriculture, Agricultural Research Service
(Milberg and Andersson 1998). For native plant special- Western Regional Plant Introduction Station in Pullman,
ists, commercial growers and managers who want to WA, assisted with making seeds available through the
grow milkweed to support habitat restoration for mon- US National Germplasm Resources Information Network

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arch butterflies (Borders and Lee-Mader 2015) and polli- and National Plant Germplasm System. A.  Miller from
nators, the use of cold stratification treatments may be the US Department of Agriculture National Laboratory
integral to successful large-scale production. For most for Genetic Resources Preservation in Fort Collins, CO,
milkweed species and populations, some germination generously provided access to unpublished results of
of seeds is likely to occur even in the absence of cold seed germination trials. We thank D. Brown, S. Elias and
stratification, but incomplete germination could result the Oregon State University Seed Lab for access to ger-
in loss of genetic variability in plants or harvested seeds mination chambers, viability testing and consultation on
and reduced adaptive potential of restored populations the project.
in the long term (Basey et al. 2015). Variation in seed
dormancy and cold stratification requirements among
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