The current link for the Wulff (1954) article.
In addition to dihaploidization, there is the matter of partial hybridization.
An unusual mechanism in crosses between cultivated sunflower and perennial Helianthus may lead to partial hybrid in first generation plants
N. Faure, H. Serieys, A. BervillÃ©, E. Cazaux and F. Kaan
"Hybridisation between the annual diploid sunflower (Helianthus annuus) and the perennial diploid species Helianthus mollis and Helianthus orgyalis was studied. Cultivated sunflower was either used as female or male parent. Progenies were obtained by means of a normal crossing or embryo rescue procedures, but hybridisation success was low. All plants examined cytologically appeared to be diploid. Plants showed a great morphological variation, but they resembled the female parent-type predominantly. In crosses with cultivated sunflower pollinated by one of the two Helianthus perennial species, 35% of the plants had a sunflower phenotype. We applied RAPD and RFLP markers to determine the genetic constitution of progenies. It appeared that the contribution of the wild male parent was reduced in comparison with the female parent: hybridisation occurred at different levels depending on the plants. On average, only 5% of the minimum number of expected RAPD and RFLP bands from the male parents were recovered in plants produced from mature seeds after pollination of sunflower by H. mollis. "
This phenomenon has also been reported in crosses between Solanum tuberosum and S. phureja. Some clones of the latter species produce “clean” dihaploid S. tuberosum, while other clones leave a small “genetic footprint”.
Also, C. G. van Tubergen, Junr. (1906) wrote:
“LILIUM.â€” Very numerous crosses among various species have been effected, and many seedlings are still under observation; a good and noteworthy race has sprung from the crossing of Lilium Martagon album with L. Hansoni. It is of particular interest to note that whereas L. Martagon album, if raised from seed, almost always comes perfectly true, scarcely ten among a thousand plants reverting to the typical purple Martagon Lily; out of the mingling of L. Martagon album with L. Hansoni not a single white martagon occurred. All plants (several hundreds) that showed no influence of the pollen-parent (L. Hansoni) reverted to the typical purple Martagon Lily. Those that showed the influence of L. Hansoni developed into stately, tall-growing lilies with broad, dark green foliage in whorls and pyramidal spikes, composed of very numerous flowers.”
I could go on.
As for the weirdness of ‘Baby Chateau’ producing tetraploid offspring with no external signs of Roxburghii (Wulff wrote, “Only anatomical studies revealed a certain similarity and relationship to the latter species.”) we may note that its grandparent ‘Eva’ (= Robin Hood x J.C. Thornton) also has more chromosomes than expected (4x rather than 3x).
I admit that I am surprised that no peonin was found in Tantau’s hybrids. Peonin + pelargonin usually gives a brighter orange than cyanin + pelargonin. Peonin (non-blueing) is apparently produced from cyanin, which often “blues”, and thus dulls the orange.
The pelargonin presumably comes from ‘Baby Chateau’. Stoddard (1980) crossed ‘Orangeade’ with a broad assortment of roses. “Crosses by R. laxa, R. suffulta, and Victor Hugo unexpectedly produced strong and pure oranges among their very sparse populations.” Suffulta, at least, has peonin-colored flowers.
Has anyone seen ‘Baby Chateau’ in the U.S.? I saw Tantau’s three hybrids, and R. roxburghii, at the Heritage Rose Garden, San Jose, CA, but might have learned more if I could have compared them with ‘Baby Chateau’. I wonder if it is as vigorous as its offspring. They reportedly have more resistance to fungal infections than ‘BC’.