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most cases, introgression was the wild type form and as a weed Helianthus annuus is believed to
viewed to have led either to an throughout much of North America. have been introduced to Texas by
increase in genetic diversity within Wild type H. annuus - the progeni- native North Americans; however,
a species, broken down or rein- tor of the domesticated sunflower28 historical details are lacking in
forced reproductive barriers, or - is believed to be indigenous to regard to the date of origin of H.
resulted in the transfer or origin of the Great Plains of the Mississippi annuus ssp. texanus and its sub-
adaptations. However, in six cases River, but has since been intro- sequent spread throughout the
there was good evidence for the ori- duced elsewhere*‘. Following its north-eastern part of the State. In
gin of a new intraspecific taxon or introduction to other parts of the contrast, such information is avail-
ecotype, and in a further eight USA, it has hybridized with other able in regard to the recent origin
examples, the origin of a new sunflower species that are either and spread in Britain of an intro-
species at the homoploid level. The native to the region or introduced gressant radiate form of the com-
list produced by Rieseberg and along with H. annuuS?O. As a conse- mon groundsel, Senecio vulgaris
Wendel was not exhaustive, and quence, H. annuus has acquired var. hibernicus. The radiate variant
further examples of the origin of genes that may have enabled it to of groundsel differs from the more
new taxa via introgression adapt rapidly to the new con- common nonradiate form, S. vul-
have appeared in the recent litera- ditions it has encountered, and in garis var. vulgaris, in that it pro-
ture’9,27.Some of the best evidence at least one case, the process has duces flower heads (capitula) with
for the introgressive origin of new led to the origin of a stabilized ray florets in addition to the disc
plant taxa comes from studies of the introgressant form of H. annuus. florets produced by var. vulgaris.
evolutionary consequences of inter- Morphological and cytological evi- The difference in capitulum type
specific hybridization involving a dence3’ suggests that the sub- results from allelic variation for a
parent or parents of alien origin species H. annuus ssp. texanus single gene which controls pres-
(Table 1). Frequently the history of originated in Texas following intro- ence/absence of ray florets”.
these events is known, allowing gression of genes from native H. Radiate groundsel was first col-
examination of the factors which debilis ssp. cucumerifolius into H. lected from the wild in 1832 from
may have favoured the spread of a annuus. Recent molecular evi- wasteground in Oxford following
new taxon following its origin. dence18 supports this hypothesis, the reported escape in 1794 of the
Recent studies on North showing that 13 of 14 populations alien radiate species, Senecio
American sunflowers (Helianrhhus of H. annuus ssp. texanus sur- squalidus (Oxford ragwort), from
spp~)18,21.25
and British ragworts and veyed possessed chloroplast DNA the nearby Oxford Botanic Garden”
groundsels (Senecio sp~.)‘~,*~are of (cpDNA) and/or nuclear ribosomal (Fig. I ). The escape of S. squalidus
particular note in this respect. DNA (rDNA) markers diagnostic of occurred after its introduction to
Helianthus annuus occurs both in H. debilis ssp. cucumerifolius. the Garden from Sicily towards the
Table 1. Plant taxa produced following a plant invasion and subsequent interspecific hybridization”
Stabilized introgressants
Intraspecific taxa Helianthus annuus H. annuus* x H. debilis Texas, USA m,c,cpDNA,rDNA 18,31
ssp. texanus ssp. cucumerifolius
Allopolyploids
Species Senecio cambrensis S. vulgaris x S. squalidus* North Wales and m,c,as,i,rDNA 24,45
Edinburgh, UK
Spartina anglica S. alterniflora* x S. maritima UK m,c,i 42
402
TREE vol. 7, no. 12, December 7992
(a) (b)
alleles in the two or more parents British allopolyploids are products times lead to the rapid evolution of
become combined and sub- of hybridization between a native new plant taxa - i.e. either stabil-
sequently maintained via linkage and an introduced species. ized introgressants or allopoly-
and/or selection in the new taxon. Each new allopolyploid species ploids. If recognized soon after
Once created, the stability of the was identified soon after its origin their origin, further study of these
gene complexes of a new and thus it has been possible to new taxa can provide an under-
introgressant taxon will depend document the establishment and standing of the factors which influ-
greatly on mechanisms preventing spread of each species up to the ence their establishment and
recombination and segregation. In present. Isozyme evidence sug spread, and thus in certain cases
the Helianthus hybrid species, and gests there was a single origin of S. the processes which lead to suc-
possibly Iris nelsonii, this appears anglica in Britain and that genetic cessful speciation. (For a discussion
to have been accomplished by diversity has remained virtually of the factors likely to affect the ori-
‘recombinant speciation”7, i.e. the absent in the species despite its gin, establishment and persistence
establishment of a new chromo- current wide distribution42. In con- of polyploid plants, see the recent
some arrangement following re- trast to S. anglica, the geographical TREE review by Thompson and
combination of two or more distinct distributions of the other new Lumaret46.) Such material also pro-
arrangements that distinguish the allopolyploids are much more vides model systems for the
parental species. The new recom- restricted; however, there is evi- assessment of risks associated with
binant types are fertile inter se, but dence in each case that the species hybridization and the release of
at least partially sterile with both are spreading4’,44, and that this has genetically engineered organisms
parents. been aided by multiple origins into the environment47. In view of
Strong ‘internal’ reproductive of the allopolyploids following the obvious advantages that this
barriers between an intraspecific hybridization between parental material holds for the study of
introgressant taxon and its parental species at different locations. evolutionary processes, it can be
taxon are unlikely to occur to the Restriction analysis of chloroplast expected that its further analysis
same degree, but should not be DNA and rDNA has revealed at will lead to significant improve-
ruled out. In Senecio vulgaris, the least two independent origins of T. ments to our current understanding
introgressant radiate taxon var. mirus and two origins of i7 miscel- of how plants evolve in the wild.
hibernicus is only found in the Ius in the Palouse region of eastern
Acknowledgements
wild, as a component of stands with Washington, USA”. lsozyme and
I thank Clive State for providing me with a
nonradiate var. vulgaris. Both vari- cpDNA data have identified at breakdown of the number of native and
ants reproduce by predominant least two independent origins for alien species included in his New Flora of
self-fertilization’7; however, the S. cambrensis, one in North Wales the British Isles, Serena Marner for slides of
and one in Edinburgh24r45.Whereas specimens of Senecfo squalidus from the
level of intervariant crossing is such
historic collections of the Fielding-Druce
that the two variants might be ex- T. miscellus, which is more abun-
Herbarium, Chris Preston for a map of the
pected to have a common genetic dant than T. mirus, has spread to current distribution of S. squalidus in the
background and be differentiated locations beyond the range of at British Isles, and Christophe Thebaud for
only by capitulum type. In fact, the least one of its diploid progeni- comments on an earlier draft of the manu-
script.
two variants also differ for certain tor.+, both T. mirus and S. cam-
allozyme traits’9,36 (controlled by brensis are normally found only at
genes unlinked to the locus con- sites where both parent species References
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