Appendix 2
Appendix Table A1: Comprehensive list of all hybrid taxa for which the hybridization-invasion (H-I) hypothesis has been suggested. We report basic
biological information, including nativity of parents relative to the region where hybrids are invasive/weedy (N = native, E = exotic); ploidy estimates
(and whether hybrids have increased ploidy relative to their parents); life form and life history. For each hybrid, we note whether any experimental tests
of the H-I hypothesis have been performed, whether observational data only have been reported or whether invasive/weedy hybrids have simply been
identified in the literature with an association between the two processes (exp, obs or ID, respectively). We also report whether hybrids have at least
one crop or domesticated parent, whether the system was previously noted in the latest review by Schierenbeck and Ellstrand 2009 (and if so, where),
and the relative timing of hybridization and introduction to the novel region (where relevant).
Parental Taxa
Parental
Nativity
Parents
polyploid?
(ploidy)
Hybrid
polyploid?
(ploidy)
Incr.
ploidy vs
parents?
ID/
exp/obs
Life form
Life
history
Crop
or
domest.
parent?
Plant Family,
Fungal/Animal
Class
Ref
from
S&E
2009
Timing of
hybridization
relative to
introduction
References
Achillea alpina
aggregate
A. acuminata ×
A. asiatica
NxN
N
Y (4X)
Y
ID
herb
perennial
N
Asteraceae
-----
pre-intro, but
after native RE
[1]
Amelanchier
erecta
A. humilis ×
A. "clade B"
NxN
Y
(2-4X x ?)
Y (4X)
?
ID
shrub
perennial
N
Rosaceae
Table 1
NA
[2]
Bothriochloa
bladhii
(=B. intermedia)
unknown
?x?
?
Y
(4X [to
8X])
?
ID
grass
perennial
N
Poaceae
-----
unknown
[3]
Bromus
hordeaceus
B. caroli-henrici ×
unknown
Ex?
Nx?
Y (4X)
?
ID
grass
annual
N
Poaceae
Table 1
unknown
[4]
Cardamine
insueta
C. rivularis × C. amara
NxN
N
Y (3X)
Y
obs
herb
various
N
Brassicaceae
Table 1
NA
[5]
Cardamine
schulzii
C. rivularis × C. amara
NxN
N
Y (6X)
Y
obs
herb
various
N
Brassicaceae
Table 1
NA
[5]
Cardamine
flexuosa "Asian"
C. fallax × C. scutata
ExE
Y
(6X x 4X)
Y (8X)
Y
ID
herb
various
N
Brassicaceae
Table 1
unknown
[6, 7]
Hybrid Taxon
Plant hybrids
Hybrid Taxon
Parental Taxa
Parental
Nativity
Parents
polyploid?
(ploidy)
Hybrid
polyploid?
(ploidy)
Incr.
ploidy vs
parents?
ID/
exp/obs
Life form
Life
history
Crop
or
domest.
parent?
Plant Family,
Fungal/Animal
Class
Ref
from
S&E
2009
Timing of
hybridization
relative to
introduction
References
Carpobrotus
hybrids
C. edulis × C.
chilensis
ExE
N
N
N
exp
shrub
perennial
N
Aizoaceae
Table 2
post-intro
[8-10]
Carpobrotus ×
cf.
acinaciformis
C. edulis ×
C. acinaciformis
ExE
N
N
N
exp*
shrub
perennial
N
Aizoaceae
-----
unknown
[11]
Carthamus
creticus
C. lanatus ×
C. leucocaulos
NxN
Y/N
(4X x 2X)
Y (6X)
Y
ID
herb
various
N
Asteraceae
-----
pre-intro, but
after native RE
[12]
Carthamus
turkestanicus
C. lanatus × C. glaucus
NxN
Y/N
(4X x 2X)
Y (6X)
Y
ID
herb
various
N
Asteraceae
-----
pre-intro, but
after native RE
[12]
Casuarina
hybrids
C. equisetifolia ×
C. glauca and/or C.
cunninghamiana
ExE
N
N
N
ID
tree
perennial
N
Casuarinaceae
-----
post-intro
[13]
Centaurea
hybrids
C. diffusa × C. stoebe
(=C. maculosa)
ExE
N/Y
(2X x 24X)
N
N
obs
herb
various
N
Asteraceae
-----
pre-intro
[14, 15]
Centaurea x
monchtonii
C. nigra × C. jacea
ExE
Y
(4X x 4X)
Y (4X)
N
obs
herb
perennial
N
Asteraceae
-----
pre-intro
[16]
Centaurea
stoebe s. l.
C. stoebe s. str. ×
unknown
Ex?
N/?
(4X x 2X)
Y (4X)
Y
exp?
herb
various
N
Asteraceae
-----
pre-intro, but
after native RE
[17]
Circaea x
intermedia
C. alpina ×
C. lutetiana
NxN
N
N
N
ID
herb
perennial
N
Onagraceae
Table 1
NA
[18, 19]
Glyceria x
pedicellata
G. fluitans × G. notata
NxN
Y/N?
(8X x ?)
N
N
ID
grass
perennial
N
Poaceae
Table 1
unknown
[19, 20]
Helianthus
annuus subsp.
texanus
H. annuus × H. debilis
NxN
N
N
N
exp
herb
annual
N
Asteraceae
Table 1
NA
[21]
Hieracium
pilosella-like
hybrids
H. pilosella ×
H. praealtum
ExE
Y
(4X x 4X)
Y (4-5X)
N
ID
herb
perennial
N
Asteraceae
-----
post-intro
[22]
Hieracium x
stoloniflorum
H. aurantiacum ×
H. pilosella
ExE
Y
(4X x 4X?)
Y (6X)
Y
ID
herb
perennial
N
Asteraceae
-----
pre-intro
[23]
Lactuca hybrids
L. serriola × L. sativa
NxE
N
N
N
exp
herb
annual
Y
Asteraceae
-----
post-intro
[24]
Hybrid Taxon
Parental Taxa
Parental
Nativity
Parents
polyploid?
(ploidy)
Hybrid
polyploid?
(ploidy)
Incr.
ploidy vs
parents?
ID/
exp/obs
Life form
Life
history
Crop
or
domest.
parent?
Plant Family,
Fungal/Animal
Class
Ref
from
S&E
2009
Timing of
hybridization
relative to
introduction
References
Mentha x
verticillata
M. aquatica ×
M. arvensis
NxN
Y
(8X x 6X)
Y (7X)
N
ID
herb
perennial
N
Lamiaceae
Table 1
NA
[20]
Myriophyllum
heterophyllum
hybrids
M. heterophyllum ×
M. laxum
ExE
N
N
N
obs
herb
perennial
N
Haloragaceae
Table 2
pre-intro
[25, 26]
Myriophyllum
spicatum hybrids
M. spicatum ×
M. sibiricum
E x E/N
Y
(6X x 6X)
unknown
unknown
exp
herb
perennial
N
Haloragaceae
-----
post-intro
[27]
Nasturtium x
sterile
(=Rorippa x
sterilis)
N. microphyllum ×
N. officinale
(=R. microphyllum × R.
nasturtium-aquaticum)
NxN
Y
(8X x 4X)
Y (6X)
N
obs
herb
perennial
Y?
Brassicaceae
Table 1
NA
[18, 28, 29]
Nuphar x
spenneriana
(=N. x
intermedia)
N. lutea × N. pumila
subsp. pumila
NxN
N
N
N
ID
herb
perennial
N
Nymphaeaceae
-----
NA
[30]
Oenothera
glazioviana
O. hookeri (=O. elata)
× O. biennis
ExE
N
N
N
ID
herb
biennial
N
Onagraceae
Table 1
post-intro, but
subsequent
introductions
[31, 32]
Onopordum
hybrids
O. acanthium ×
O. illyricum
ExE
N/N?
(2X x 2X)
N
N
ID
herb
biennial
N
Asteraceae
Table 2
pre-intro
[33]
Persicaria
maculosa
(=Polygonum
persicaria)
P. foliosa ×
P. lapathifolia
ExE
N
Y (4X)
Y
ID
herb
annual
N
Polygonaceae
-----
pre-intro, but
after native RE
[34]
Persicaria
punctata
P. hydropiper ×
P. hirsuta/setacea
ExE
N
Y (4X)
Y
ID
herb
various
N
Polygonaceae
-----
pre-intro
[34]
Persicaria
pensylvanica
(=Polygonum
pensylvanicum)
P. glabra/hispida ×
unknown
unknown
(Y/N)/?
(2X or 46X x ?)
Y (8X)
Y
ID
herb
various
N
Polygonaceae
-----
unknown
[34]
Parental
Nativity
Parents
polyploid?
(ploidy)
Hybrid
polyploid?
(ploidy)
Incr.
ploidy vs
parents?
ID/
exp/obs
Life form
Life
history
Crop
or
domest.
parent?
Plant Family,
Fungal/Animal
Class
Ref
from
S&E
2009
Timing of
hybridization
relative to
introduction
References
R.raphanistrum ×
R. sativus
ExE
N
N
N
exp
herb
annual
Y
Brassicaceae
Table 2
post-intro
[35, 36]
Reynoutria x
bohemia
(=Fallopia x
bohemica)
R. japonica ×
R. sachalinensis
(=F. japonica ×
F. sachalinensis)
ExE
Y (4/6/8X
x 4/8X)
Y
(6X
[4/8X])
N
exp
herb
perennial
N
Polygonaceae
Table 1
post-intro, but
subsequent
introductions
[37, 38]
Rhododendron
hybrids
R. ponticum ×
R. catawbiense
ExE
N
N
N
ID
shrub
perennial
N
Ericaceae
Table 2
post-intro
[39]
Rorippa x
armoracioides
R. sylvestris ×
R.austriaca
NxE
N/Y
(4/6X x
2X)
Y (3-5X)
N
obs
herb
perennial
N
Brassicaceae
Table 1
post-intro, but
subsequent
introductions
[40]
Salsola Type C
S. tragus × S. kali ssp.
Austroafricanus
ExE
Y/N
(4X x 2X)
Y
(6X [4X])
Y (N)
ID
herb
annual
N
Chenopodiaceae
Table 1
post-intro
[41]
Salsola lax
S. tragus × S. kali ssp.
Austroafricanus ×
S. paulsenii (?)
ExE
Y/N/Y
(6X x 2X x
6X)
Y (6X)
N
ID
herb
annual
N
Chenopodiaceae
-----
unknown
[41]
Sarcocornia
hybrids
S. perennis ×
S. fruticosa
NxN
Y/N
(2X x 8X)
Y
(6X [45X])
N
exp
shrub
perennial
N
Chenopodiaceae
Table 2
NA
[42, 43]
Senecio vulgaris
var. hibernicus
S. squalidus × S.
vulgaris var. vulgaris
ExN
N/Y
(2X x 4X)
Y (4X)
N
exp
herb
various
N
Asteraceae
Table 1
post-intro
[44]
Senecio
squalidus
S. aethnensis × S.
chrysanthemumifolius
ExE
N
N
N
exp
herb
various
N
Asteraceae
Table 1
pre-intro
[45]
Solanum x
edinense
S. tuberosum ssp.
andigena ×
S. demissum
NxN
Y
(4X x 6X)
Y (5X)
N
ID
herb
perennial
Y
Solanaceae
Table 1
NA
[46, 47]
Solanum x rechei
S. microdontum ×
S. kurtzianum
NxN
N
N
N
ID
herb
perennial
N
Solanaceae
-----
NA
[47, 48]
Solanum x
sambucinum
S. ehrenbergii ×
S. pinnatisectum
NxN
N
N
N
ID
herb
perennial
N
Solanaceae
-----
NA
[47]
Sorghum almum
S. halepense ×
S. bicolor
ExE
N/Y
(4X x 2X)
Y (4X)
N
exp
grass
perennial
Y
Poaceae
Table 1
post-intro
[49]
Hybrid Taxon
Parental Taxa
Raphanus
hybrids
Hybrid Taxon
Parental Taxa
Parental
Nativity
Parents
polyploid?
(ploidy)
Hybrid
polyploid?
(ploidy)
Incr.
ploidy vs
parents?
ID/
exp/obs
Life form
Life
history
Crop
or
domest.
parent?
Plant Family,
Fungal/Animal
Class
Ref
from
S&E
2009
Timing of
hybridization
relative to
introduction
References
Sorghum
halepense
S. bicolor ×
S. propiniquum
ExE
N
Y (4X)
Y
ID
grass
perennial
Y
Poaceae
Table 1
post-intro, but
subsequent
introductions
[50]
Spartina hybrids
S. alterniflora ×
S. foliosa
ExN
Y
(6X x 6X)
Y (6X)
N
exp
grass
perennial
N
Poaceae
Table 2
post-intro
[51, 52]
Spartina anglica
S. maritima ×
S. alterniflora
NxE
N
Y (12X)
Y
ID
grass
perennial
N
Poaceae
Table 1
post-intro, but
subsequent
introductions
[53]
Spartina
densiflora
S. arundinacea ×
S. alterniflora
NxN
Y
(4X x 6X)
Y (7X)
Y
ID
grass
perennial
N
Poaceae
-----
NA
[54]
Sphagneticola
hybrids
S. trilobata ×
S. calendulacea
ExN
Y
(4X x 4X)
Y (4X)
N
exp
herb
perennial
N
Asteraceae
-----
post-intro
[55]
Stachys x
ambigua
S. palustris ×
S. sylvatica
NxN
Y (12-13X
x 8X)
Y (10X)
N
ID
herb
perennial
N
Lamiaceae
Table 1
NA
[19]
Symphytum x
uplandicum
S. officinale ×
S. asperum
NxE
Y
(4X x 2X)
Y (3X+)
N
ID
herb
perennial
N
Boraginaceae
-----
pre-intro
[56, 57]
Tamarix hybrids
T. ramosissima ×
T. chinensis
ExE
N
N
N
obs
shrub
perennial
N
Tamaricaceae
Table 2
post-intro
[58, 59]
Tragopogon
mirus
T. dubius ×
T. porrifolius
ExE
N
Y (4X)
Y
ID
herb
biennial
N
Asteraceae
Table 1
post-intro
[60, 61]
Tragopogon
miscellus
T. dubius ×
T. pratensis
ExE
N
Y (4X)
Y
ID
herb
biennial
N
Asteraceae
Table 1
post-intro
[60, 61]
Typha x glauca
T. angustifolia ×
T. latifolia
ExN
N
N
N
exp
graminoid
perennial
N
Typhaceae
Table 1
post-intro
[62, 63]
Ulmus hybrids
U. rubra × U. pumila
NxE
N
N
N
ID
tree
perennial
Y
Ulmaceae
-----
post-intro
[64]
Viola hybrids
V. riviana ×
V. reichenbachiana
NxN
Y/N
(4X x 2X)
Y (4X)
N
ID
herb
perennial
N
Violaceae
Table 2
NA
[65]
Viola x tatrae
V. tricolor × V. lutea
ssp. sudetica
ExN
Y
(4X x 8X)
Y
(6X and
up)
N
exp
herb
perennial
N
Violaceae
-----
post-intro
[66]
Hybrid Taxon
Parental Taxa
Parental
Nativity
Parents
polyploid?
(ploidy)
Hybrid
polyploid?
(ploidy)
Incr.
ploidy vs
parents?
ID/
exp/obs
Life form
Life
history
Crop
or
domest.
parent?
Plant Family,
Fungal/Animal
Class
Ref
from
S&E
2009
Timing of
hybridization
relative to
introduction
References
Wisteria hybrids
W. floribunda ×
W. sinensis
ExE
N
N
N
obs
vine
perennial
Y
Fabaceae
-----
post-intro, but
subsequent
introductions
[67]
Melampsora x
columbiana
M. medusae ×
M. occidentalis
N/E x N
?
N
exp
fungus
NA
N
Urediniomycetes
-----
post-intro
[68, 69]
Ophiostoma
novo-ulmi
O. novo-ulmi ×
O. ulmi
ExE
?
N
ID
fungus
NA
N
Sordariomycetes
-----
post-intro, but
subsequent
introductions
[70, 71]
Phytophthora
alni alni
P. alni uniformis ×
P. alni multiformis
N? x E?
N
Y
ID
stramenopile
NA
N
Oomycetes
Table 4
post-intro
[72, 73]
Phytophthora
andina
P. infestans ×
unknown
E x ??
?
N
ID
stramenopile
NA
N
Oomycetes
-----
unknown
[74]
Phytophthora x
pelgrandis
P. cactorum ×
P. nicotianae
NxE
?
N
ID
stramenopile
NA
N
Oomycetes
-----
unknown
[75]
Phytophthora x
serendipita
P. cactorum ×
P. hedraiandra
NxE
?
N
ID
stramenopile
NA
N
Oomycetes
-----
unknown
[76]
Pythium hybrids
P. arrhenomanes (or
similar) × P. phragmitis
ExN
?
N
exp
stramenopile
NA
N
Oomycetes
-----
unknown
[77]
Verticillium
longisporum
V. dahliaea × unknown
?? X ??
N
Y (2X)
ID
fungus
NA
N
Ascomycota
-----
unknown
[78]
Ambystoma
hybrids
A. tigrinum mavortium
× A. californiense
ExN
?
N
exp
amphibian
NA
N
Amphibia
-----
post-intro
[79, 80]
Carcinus hybrids
C. maenas ×
C. aestuarii
ExE
N
N
N
ID
crab
NA
N
Malacostraca
-----
unknown
[81, 82]
Cottus hybrids
C. rhenanus ×
C. perifretum
NxN
?
N
N?
obs
fish
NA
N
Teleostei
-----
post-intro
[83]
Cyprinella
hybrids
C. lutrensis ×
C. venusta stigmatura
ExN
?
N
exp
fish
NA
N
Actinopterygii
-----
post-intro
[84]
Salvelinus
hybrids
S. fontinalis ×
S. alpinus
NxN
?
N
ID
fish
NA
N
Actinopterygii
-----
post-intro
[85]
Fungal hybrids
Y
Animal hybrids
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