146 
R. Ruggles Gates. 
segregation is apparently the same type of behaviour as was found 1 
in the F 2 and F 3 of crosses between the large-flowered CE. rubricalyx 
and the small-flowered (E. biennis , in which large and small flowers 
frequently appeared on the same plant, and even long and short 
petals, of varying lengths, on the same flower. It is possible that 
this race of CE. lamavckiana var. cruciata , which emanated from 
the Bremen Botanic Garden, may have originated by a cross 
between CE. biennis leptomeres and CE. lamavckiana. This hypo¬ 
thesis is supported by the short style in the flowers of this race, so 
short that, like CE. biennis , nearly all the flowers were self-pollinated. 
But in crosses between CE. biennis and its cruciate mutation there are 
no reports of this vegetative segregation, with both types of flowers 
on the same plant. 
As a result especially of the studies of MacDougal (1905) and 
Bartlett (1914a), a number of cruciate species and varieties are 
now accurately known. As Bartlett points out, it is clear that 
these have not had a common origin, but they have arisen 
independently from different stocks, some more remotely and some 
more recently, just as var. leptomeres apparently originates even 
now from CE biennis L. The first cruciate form to be discovered 
was CE. cruciata Nutt. Its taxonomic history is given by Bartlett. 
Nuttall found it in Massachusetts and sent seeds of it to several 
botanists. Don grew it in 1824 and described it under Nuttall’s 
name. The species which is still cultivated in botanical gardens 
under this name is quite likely descended from this original source. 
Its variable petals indicate that it has been crossed with a broad- 
petalled species, and it is now called CE. cruciata , var. varia, de 
Vries. 2 In the sixth edition of Gray’s Manual (1889) it was classed 
as probably a rare garden sport, since no specimens had been 
collected except in gardens, subsequent to the original collection 
over 60 years earlier. In later years cruciate specimens were 
collected in Vermont. 
The interest awakened in wild North American CEnotheras 
by the work of de Vries, soon led to the discovery of other forms. 
Seeds of a cruciate form which had been collected at Sandy Hill, 
New York, near Lake George, in 1902, were divided, part of them 
being grown by MacDougal (1905) at New York, and a larger 
number by de Vries at Amsterdam. This material contained two 
1 See Gates, 1917a. 
3 De Vries found that this variety segregated roughly into three groups, 
(1) cruciate, (2) intermediate, (3) broad-petalled. The latter is constant except 
for rare cruciate bud mutations. 
