Appendix: Supplementary Information
Table A1 Theoretical studies simultaneously examining dispersal through space and time
Table A2 Empirical studies investigating combinations of dispersal through space and time in wild populations
TABLE A1
Type of dispersal
Authors
Approach
Type of
regulation
Temporal
Den
Boer
(1968)
Review /
Idea
Reddingt
us &
Den
Boer
(1970)
Numerical
simulations
Southwo
od
(1977)
Review /
Idea
Venable
&
Lawlor
(1980)
Bakshtan
sky
(1980)
Analytical
results
Review /
Idea
Densitydependent /
Densityindependent
Densityindependent
Spatial
Covariat
ion of
dispersal
strategie
s
Level of
biological
organizatio
n
Spatiotemporal
correlation
of the
environmen
t
Conditi
ondepende
nt
dispersa
l
Approach
for
studying
consequen
ces
Metric of
dispersal
(Spatial/Tem
poral)
Stages of the
dispersal
process
(Spatial/Tempo
ral)
Dormancy / Diapause /
Age-structure
Unspecifie
d
Individual,
Population,
and Species
No
Unspecified/
Unspecified
Unspecified /
Unspecified
Age-structure
Unspecifie
d
Population
No
Dispersal rate
/ Age
structure
variability
Emigration /
Emigration
Dormancy
Unspecifie
d
Individual
No
Dispersal
distance /
Dormancy
duration
Emigration /
Emigration
Dormancy
Unspecifie
d
Individual
(plants
producing
two morphs)
Yes
(predicta
bility of
germinat
ion)
Age-structure /
reproductive strategy
(iteroparity vs semelparity)
Straying of
adults
outside of
their natal
river
Negative
Negative
Species
No
germination
fraction
Dispersal rate
/ Dormancy
rate
Emigration /
Emigration
Straying rate
/ Age
structure
variability
Unspecified /
Unspecified
Main results
Spatial and
temporal
dispersal allow
spreading of risk
and contribute to
population
stability.
Heterogeneity of
the environment
and within a
population
contribute to
population
stability.
Organisms adopt
strategies that
vary in space
and time to
maximize
reproductive
success.
Negative
covariation
between spatial
and temporal
dispersal
strategies
evolves in an
unpredictable
environment.
The age
complexity
among salmonid
fishes is a form
of “protection in
time” while
spatial dispersal
between
populations is a
Age-structure /
reproductive strategy
(iteroparity vs semelparity)
Straying of
adults
outside of
their natal
river
Negative
Species
Straying rate
/ Age
structure
variability,
reproductive
strategy
Quinn
(1984)
Review /
Idea
Levin et
al.
(1984)
Analytical
results and
numerical
simulations
/
Population
model
without
structure
Densitydependent/F
requencydependent
Dormancy
Unspecifie
d
Negative
Individual
No
No
Cohen &
Levin
(1987)
Analytical
results and
numerical
simulations
/
Population
model
without
structure
Densityindependent
Dormancy
Unspecifie
d
Negative
Individual
No
No
Evolutiona
rily Stable
Strategy
Emigration /
Emigration
Dispersal rate
/ Dormancy
rate
Emigration /
Emigration
Dispersal rate
/ Dormancy
rate
Emigration /
Emigration
form of
“protection in
space”
Temporal
dispersal may be
balanced by
spatial dispersal
within the
Pacific
salmonids
Negative
covariation
between
dormancy and
spatial dispersal
evolve because
both are
mechanisms for
escaping
environmental
unpredictability.
Both spatial and
temporal
dispersal are
expected to
increase if the
variability of the
environment
increases. The
covariation
between spatial
and temporal
dispersal is also
affected by
temporal
autocorrelation
of environmental
conditions.
Negative
covariation
between spatial
and temporal
dispersal evolve
in an
unpredictable
environment but
the covariation is
affected by costs
Seger &
Brockma
nn
(1987)
Review /
Idea
Klinkha
mer et al.
(1987)
Analytical
results and
numerical
simulations
Venable
& Brown
(1988)
Numerical
simulations
Densitydependent /
Densityindependent
Densityindependent
/Frequencyindependent
Dormancy
Dormancy
Unspecifie
d
Unspecifie
d
Negative
Negative
Dispersal rate
/ Dormancy
rate
Emigration /
Emigration
Individual
(one
population
with many
patches)
No
No
Dispersal rate
/ Germination
rate
Emigration /
Emigration and
Transfer
Individual
Yes (with
/without
temporal/sp
atial
correlation,
respectively
)
No
Dispersal rate
/ Dormancy
rate
Emigration /
Emigration and
Transfer (via
survival)
of dispersal.
Spatial and
temporal
dispersal can be
interpreted as
bet-hedging
strategies and
both can
promote
coexistence of
species.
Observed
negative
correlation
between spatial
dispersal and
delayed
germination in
annuals plants,
including from
models with
both densityindependent and
the densitydependent
regulation.
Spatial and
temporal
dispersal allow
an escape from
sib-competition.
Dormancy,
dispersal, and
seed size interact
to reduce risk in
temporally and
spatially variable
environments,
and are partly
substitutable.
Increasing
(positive) spatial
autocorrelation
of environmental
conditions
favors decreased
spatial dispersal
and increased
dormancy.
Philippi
& Seger
(1989)
Review /
Idea
Cohen &
Levin
(1991)
Analytical
results
Densityindependent
Dormancy / Diapause
Unspecifie
d
Dormancy
Unspecifie
d
Negative
Individual
No
No
Dispersal rate
/ Dormancy
rate
Emigration /
Emigration
Yes
(temporal
autocorrelati
on)
No
Dispersal rate
/ Dormancy
rate
Emigration /
Emigration
Increasing
(positive)
temporal
autocorrelation
favors decreased
spatial dispersal
and dormancy.
Temporal and
spatial dispersal
are
complementary
and partially
substitutable.
Increasing
number of
patches within a
meta-population
favors spatial
dispersal as a
risk-spreading
strategy while
temporal
dispersal and
spatial dispersal
both decrease
with increasing
(positive)
temporal
autocorrelation
in the
environment.
Temporal
dispersal reduces
spatial dispersal
at all levels of
temporal
autocorrelation
in the
environment.
The effect of
temporal
dispersal is
strongest when
there is a
negative
temporal
environmental
correlation, is
moderate at zero
Wiener
&
Tuljapur
kar
(1994)
Analytical
results and
numerical
simulations
/ Matrix
model with
structure
Eriksson
(1996)
Eriksson
&
Kiviniem
i (1999)
Dormancy / Age-structure
Unspecifie
d
Negative
Individual
(one
population
with two
subpopulations)
Review /
Idea
Dormancy
Unspecifie
d
Negative
Species
Review /
Idea
Dormancy
Unspecifie
d
Densityindependent
Yes
No
No
No
Long-term
growth rate
Dispersal rate
/ Dormancy
rate
Emigration /
Emigration and
Transfer (via
costs of
migration)
Unspecified/
Unspecified
Unspecified /
Unspecified
Dispersal
distance /
Unspecified
Emigration /
Unspecified
correlation, and
is weak when
the environment
is positively
autocorrelated.
Spatial dispersal
is favored in
variable
environments,
by negative
spatial
autocorrelation,
and by a large
population
structure but the
benefits are
reduced when
temporal
dispersal also
occurs.
Temporal
dispersal
contributes to
population
buffering and
increases the
time to
extinction even
in declining
populations.
Proposed the
concept of
remnant
populations
(populations
which disperse
in space and
time) as an
alternative to
metapopulation
and source-sink
approaches.
Review of
spatial and
temporal
dispersal
strategies in
plants that
revealed the
Eriksson
(2000)
Review /
Idea
McPeek
& Kalisz
(1998)
Analytical
results and
numerical
simulations
/ Stochastic
matrix
model
Laterra
&
Solbrig
(2001)
Numerical
simulations
Densityindependent
Densitydependent
Dormancy
Unspecifie
d
Dormancy
Unspecifie
d
Dormancy
Unspecifie
d
Negative
Negative
Species
No
Individual
No
Population,
Species
Yes
Dispersal
distance /
Unspecified
No
No
Geometric
mean
fitness
Emigration,
Transfer, and
Settlement /
Unspecified
Dispersal rate
/ Dormancy
rate
Emigration and
Transfer /
Emigration and
Transfer
Dispersal rate
/ Dormancy
rate
Emigration,
Transfer, and
Settlement /
Emigration,
Transfer, and
Settlement
existence of
trade-offs
between
dispersal
strategies. The
authors
summarized
multiple
hypotheses to
explain this
pattern and the
joint evolution
of alternative
dispersal
strategies in
plants.
Highlighted the
conservation
implications of
dispersal
through space
and time for
plants.
Spatial dispersal
is favored over
temporal
dispersal with
increasing
number of
patches and until
the fitness cost
of spatial
dispersal greatly
exceeds the
fitness cost of
temporal
dispersal.
A short fire-free
interval favors
plants species
with low spatial
and high
temporal
dispersal
strategies, while
high spatial and
low temporal
dispersal
strategies are
Olivieri
(2001)
Numerical
simulations
/
determinist
ic
markovian
model
Mathias
& Kisdi
(2002)
Numerical
simulations
/ Adaptive
dynamic
Bohonak
&
Jenkins
(2003)
Review /
Idea
Snyder
(2006)
Analytical
results
Densitydependent /
Densityindependent
Densitydependent
Densitydependent
Dormancy
Unspecifie
d
Negative
Individual,
Population
No
No
Evolutiona
ry Stable
Strategy
Dispersal rate
/ Dormancy
rate
Emigration and
Transfer /
Emigration and
Transfer
No
No
Long-term
growth rate
Dispersal rate
/ Dormancy
rate
Emigration /
Emigration
No
Unspecified/
Unspecified
Unspecified /
Unspecified
Dispersal
distance and
rate /
Dormancy
rate
Emigration and
Transfer /
Emigration and
Transfer
Dormancy
Unspecifie
d
Negative
Individual
(one
population
in two
habitats)
Diapause
Active
dispersal/P
assive
dispersal
Substitut
able
Individual,
Population,
Species
No
Dormancy
Distance
dispersal
Negative/
Positive
Individual
Yes
(positively
correlated in
space and
time)
No
Evolutiona
ry Stable
Strategy
favored when
the fire-free
interval
increases.
Evolutionary
stable dormancy
rates increase
with local
extinction rates,
decrease with
increasing cost
of dormancy,
and increase
with the cost of
dispersal.
Selective tradeoff between
dormancy and
dispersal. The
long-term
growth rate of a
strategy is
determined by
the joint
distribution of
the environment
and population
density.
Authors
highlight that
diapause and
migration can be
thought of as
alternative
strategies for
spreading risk
through time and
space.
Author reports a
positive
correlation
between spatial
and temporal
dispersal in the
presence of
positive
temporal
autocorrelation
Rajon et
al.
(2009)
Vitalis et
al.
(2013)
Analytical
results and
numerical
simulations
/ Adaptive
dynamic
Analytical
results and
numerical
simulations
/
Individualbased
model
Densitydependent
Densitydependent
Dormancy
Dormancy
Unspecifie
d
Unspecifie
d
Negative
Negative/
Positive
Individual
(one
population
in two
habitats)
Individual
No
Uncorrelate
d (constant
vs.
stochastic
environment
)
No
Yes
(condtio
nal vs.
uncondti
onal
dormanc
y)
Adaptive
dynamics
with
Evolutiona
ry Stable
Strategy
Evolutiona
rily Stable
rate of
dormancy
Dispersal rate
/ Dormancy
rate
Dispersal rate
/ Dormancy
rate
Emigration and
Transfer /
Emigration and
Transfer
Emigration,
Transfer, and
Settlement /
Emigration,
Transfer, and
Settlement
in environmental
conditions. She
also found that
dormancy
reduces the
optimal fraction
of dispersers
when the
environment is
correlated only
over short time
intervals and
increases the
optimal dispersal
fraction when
the environment
is correlated
over longer time
intervals.
When the spatial
dispersal rate is
high, an
intermediate
dormancy
strategy (i.e., a
generalist
strategy) can be
selected because
it confers higher
mean fitness at
the
metapopulation
level than either
local specialist.
Increasing the
cost of
dormancy
selected against
dormancy and
favored spatial
dispersal, while
increasing the
cost of spatial
dispersal
selected against
spatial dispersal
and favored
dormancy.
TABLE A2
Type of dispersal
Authors
Approach
Type of organisms
Temporal
Covariation
of dispersal
strategies
Level of
biological
organization
Geographical
location /
Type of
environments
Metric of dispersal
(Spatial/Temporal)
Stages of the
dispersal process
(Spatial/Temporal)
Spatial
Emigration,
Transfer, and
Settlement /
Emigration,
Transfer, and
Settlement
Observed higher
dispersal for a
semelparous
species (coho)
than for an
iteroparous
species
(steelhead) in two
streams.
Reported a
negative
relationship
between
dormancy and
spatial dispersal
among individuals
for most species.
Found an apparent
trade-off between
dormancy and
spatial dispersal
(i.e., reported
combinations of
reduced spatial
dispersal/delayed
germination and
distance
dispersal/quick
germination.
Reported seed
dimorphism with
different
properties of
temporal and
spatial dispersal.
Emigration /
Emigration
Found that the
occurrence of a
Shapovalov &
Taft (1954)
Field
observations /
Comparison of 2
sites
Coho salmon
(Oncorhynchus
kisutch) and steelhead
(O. mykiss)
Semelparity
vs
Iteroparity
Dispersal to a
non-natal
river
Negative
Species
USA /
Mediterranean
climate
Straying rate /
parity
Emigration/
Emigration
Venable &
Lawlor (1980)
Meta-analysis /
Comparative
study
List of desert plants in
the families
Asteraceae and
Brassicaceae
Dormancy
Qualitative
measure of
dispersal
capability
Negative
Individual
(with species
comparisons)
Unspecified /
Arid
environment
Unspecified/
Unspecified
Emigration/
Emigration
McEvoy (1984)
Laboratory /
Comparative
study of seeds
germination
Dormancy
Qualitative
measure of
dispersal
capability
Unspecified/
Germination rate
Emigration /
Emigration and
Transfer
germination times
Senecio jacobaea L.
(Asteraceae)
Venable & Levin
(1985)
Laboratory /
Comparative
study of seeds
germination
Heterotheca latifolia
(Asteraceae)
Eriksson (1992)
Meta-analysis /
Comparative
61 angiosperm species
Main results
Negative
Individual
USA /
Temperate
environment
Dormancy
Qualitative
measure of
dispersal
capability
Negative
Individual
USA / arid
and semi-arid
environment
Seed dimorphism /
Seed dimorphism
Dormancy
Qualitative
measure of
No
Species
Sweden /
Temperate,
Long distance
dispersal /
study
Rees (1993)
Meta-analysis /
Comparative
study
dispersal
capability
British plants (171
species, representing
34 families and 114
genera)
Dormancy
Qualitative
measure of
dispersal
capability
Qualitative
measure of
dispersal
capability
artic and
alpine
environments
Negative
Unspecified
seed bank was not
related to seed
spatial dispersal.
Species
UnitedKingdom /
Temperate but
with wide
range of
habitats
Unspecified /
Germination
success
Unspecified/
Emigration and
Transfer
Species
Unspecified /
Mostly
temperate –
but wide
range of site
conditions
(e.g. prairie,
crop fields,
sand dune)
Dispersal distance/
Germination time
Emigration /
Emigration and
transfer
Meta-analysis /
Comparative
study
89 species of
herbaceous plants
Eriksson (1996)
Meta-analysis /
Comparative
study
Herbs (N=66) and
grasses (N= 86)
inhabiting Swedish
semi-natural
grasslands and
deciduous forests
Dormancy
Qualitative
measure of
dispersal
capability
Negative
Species
Sweden /
Temperate
and boreal
landscapes
Unspecified /
Dormancy rate
Emigration /
Emigration
Imbert (1999)
Experimental /
Comparison of
the viability of
dimorphic seeds
stored for up to 5
years
Crepis sancta
(Asteraceae)
Dormancy
Qualitative
measure of
dispersal
capability
Negative
Species
France /
Mediterranean
environment
Dispersal distance /
Germination rate
and seedling
survival
Emigration /
Emigration and
Transfer
Diapause
Qualitative
measure of
dispersal
capability
Species
Germany /
Temperate
environment
(Temporary
flooded
grassland)
Unspecified /
Emergence and
survival to diapause
Unspecified /
Emigration and
Transfer
Willson et al.
(1993)
Frisch (2002)
Field
observations and
laboratory study
Cyclopoid copepods
(Cyclopoida,
Copepoda)
Dormancy
Negative but
weak
Unspecified
Reported a
reduction of seed
dormancy in
species that had
an efficient means
of dispersing
through space.
Few species with
poor mechanisms
for dispersal
through space had
the capacity for
better dispersal
through time.
The majority of
species evaluated
displayed some
form of temporal
dispersal (e.g.,
seed banks) but
lacked traits that
favored spatial
dispersal.
Presence of
dimorphic seeds
that can disperse
in space and time
may explain the
persistence of this
colonizing species
in communities
dominated by
perennial species.
Both spatial
dispersal and
dormancy in
temporary water
bodies are
important for the
survival of
cyclopoid
copepods in
floodplains.
Bégin &
Roff (2002)
Field
observations and
laboratory
experiment
Spring field cricket
(Gryllus veletis)
Gravuer (2003)
Field
observations /
Comparison of
14 sites
Northern blazing star
(Liatris scariosa var.
novae-angliae)
Dostál (2005)
Meta-analysis /
Comparative
study
5 annual plant species
(Arenaria
serpyllifolia,
Androsace elongata,
Myosotis ramosissima,
Saxifraga tridactylites
and Veronica
arvensis)
Robinet et al.
(2008)
Release–
recapture
experiments to
parameterize a
stochastic
individual based
model
Gypsy moth
(Lymantria dispar L.)
Diapause
Wing
dimorphism
(microptery
vs.
macroptery)
Negative
(positive
between
direct devel
opment and
macroptery).
Individual
North
America /
Temperate
environment
Dispersal rate /
Diapause rate
Emigration /
Emigration
Germination
success
Measured
drop time
and flight
distance
Negative
Population
USA /
Temperate
environment
Dispersal distance /
Germination rate
Emigration /
Emigration and
Transfer
Dormancy
Quantitative
measure of
dispersal
capability
(distance
from mother
plant)
Negative but
weak
Species
East Europe /
Temperate
perennial
grassland
Dispersal distance /
Germination rate
Temporal
sexual
asynchrony
Qualitative
measure of
dispersal
capability
Unspecified
Individual
USA /
Temperate
environment
Dispersal distance /
Unspecified
Emigration /
Emigration and
Transfer
Emigration /
Emigration
Observed a
negative
covariation
between wing
morphology and
diapause
occurrence, as
well as positive
phenotypic and
genetic
correlations
between diapause
occurrenceand
wing morphology.
Observed negative
covariation
between estimated
drop time and
germination
success, and
estimated flight
distance and
germination
success among
populations.
Cautioned that
germination
success could be
influenced by
variation in seed
dormancy or
intrinsic viability
in different
populations.
Seed banks were
extensive, but had
only weak effect
on dynamics.
When either
temporal or spatial
dispersal was
high, males were
more easily lost
from the
population and
Siewert &
Tielbörger
(2010)
Semiexperimental/
Controlled for
spatial dispersal
distance
Stipa capensis
(Poaceae), Filago spp
(Asteraceae),
Hymenocarpos
circinnatus
(Fabaceae), Anagallis
arvensis (Primulaceae)
& Plantago cretica
(Plantaginaceae)
Dormancy
Stevens et al.
(2013)
Meta-analysis /
Comparative
study
Pélisson et al.
(2013)
Field
observations and
experiments
4 weevil species
(Curculio spp.)
Diapause
Westley et al.
(2013)
Field
observations and
hatchery releases
Chinook salmon (O.
tshawytscha) and
steelhead (O. mykiss)
Semelparity
vs
Iteroparity
Butterflies (>15
species)
Diapause
Quantitative
measure of
dispersal
capability
(distance
from mother
plant)
Mean
dispersal
distance,
Frequency of
long-distance
dispersal,
Dispersal
propensity,
Gene flow
Negative but
weak
No
Population
and Species
Israel /
Subhumid
over the
semiarid to
the arid zone
Species
Europe /
Temperate
environment
France /
Temperate
environment
Dispersal distance
USA /
Temperate
climate
Dispersal rate /
parity
Flight
performance
and distance
Negative but
weak
Species
Dispersal to a
non-natal
river
Negative
Species
Dispersal distance
and rate /
Dormancy rate
Dispersal distance,
rate and gene flow /
Diapause rate
/ Diapause rate
Emigration /
Emigration
mating success
decreased.
Concluded that
pre- mating
dispersal may be
inversely related
to probability of
establishment.
Found that the
importance of
spatial dispersal
for population and
community
dynamics was
extremely low
relative to local
reproduction and
dormancy.
Emigration and
Settlement /
Emigration
Emigration /
Emigration
Emigration/
Emigration
Half of the four
species studied
showed evidence
of a trade-off
(negative
correlation).
Observed higher
dispersal for a
semelparous
species (Chinook)
than for an
iteroparous
species
(steelhead).