Seven species were included in this complex before 《Fl. Reip. Pop.
Sin.》, which recognized 3 species. Carried out in this work were population
sampling ( 46 populations in total), cultivation, experimental observation,
chromosome observation (published), crossing, isozymogramatic comparison, character analysis (including numerical taxonomical analysis), stem anatomy and
palynological observation. The results are shown as follows.
1. This complex is found almost always out-pollinated as it is always
protandrous. No apomixis is found. Seedlings are of only basal leaves in the first
year, and it is only in the second year that they develop stems and come into
2. Among 32 chromosome-counted populations, all in eastern Liaoning, most
in western Liaoning and two in Haoxian County of Shanxi Province are found to
be 2x (2n=34), and all the others 4x (2n=68).
3. A rather wide range of zymogramatic variation of leaf esterase was detected
in the complex. But seed esterase was quite constant.
4. Based on the character analysis (including numerical taxonomical analysis)
and stem anatomy, the populations'are found to be different from one another in
size and form of corollas, size of fruits, size and indumentum of calyces, size and
form of leaves (ratios of length and width), teeth of leaves, relative length of
styles and corollas, seed form as well as presence or absence of medullary fibre.
As a result, 8 races are recognized. A diploid distributed in western Liaoning,
which was recognized as a variety, A. polantha var. contracta, is so distinct in
gross morphology that it is treated here as a species, A. contracta (Kitag.) J.Z.
Qiu et Hong. With the other 7 races, 3 species (A. gmelinii, A. polyantha and A.
elata) and 5 subspecies are recognized.
5. As a result of crossing among 9 populations, some F1 seeds were produced
between the tetraploids, but none between the tetraploids and the diploids from
6. Inferred from hybrid index method, the tetraploid populations of A.
gmelinii and A. polyantha may have hybridized in eastern Shanxi Province and
western Hebei Province, which may explain the intergradation of these two
The evolution of this complex is finally deduced. Its ancestor might occupy in
the past a rather wide area where it differentiated into three diploid races. From
the one now distributed in eastern Liaoning, the tetraploid of A. polyantha might
have evolved and migrated southwards; from the one in Haoxian (Shanxi), a
tetraploid of A. gelinii might have been derived and expanded its area northwards, and finally, its subspecies might have differentiated. No diploid has been
found in A. elata. Two diploids might be involved in its origin. One might be
the diploid of A. gmelinii, and the other a diploid with calyx teeth (not includedin the complex).
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