Clos (1849) followed Bentham in keeping Sa/pig/ossis in the 
Scrophulariaceae; at the same time Gay (1849) in describing the 
new but obviously related genus Revesia, assigned it to the Bigno- 
niaceae. Sucha disparity of opinion concerning the proper alloca- 
tion of these three genera in the system can be easily understood if 
one takes into account both the floral zygomorphy and the tetra- 
dynamous androecium. 
Even as late as 1888, Baillon accepted the system of Bentham, 
keeping the tribe Salpiglossideae in the Scrophulariaceae. The 
first author to disagree with this classification was John Miers. In 
a brilliant research published in 1849, he transferred the whole 
tribe Salpiglossideae to the Solanaceae (including four genera: 
Salpiglossis, Leptoglossis, Browallia and Pteroglossis; this last 
name, as it is now understood, is a synonym of Revesia). When 
Bentham elaborated the Gamopetalae for the Genera Plantarum 
(2: 882, 913, 1244. 1876), he willingly accepted the ideas of Miers 
and placed. the Salpiglossideae (comprising Salpiglossis, Lepto- 
glossis, Revesia and twelve additional genera) in the Solanaceae. 
By then, the German school was making important contribu- 
tions for the proper delimitation between Solanaceae and Scro- 
phulariaceae. As early as 1866 (pag. 518), Wydler published an 
epoch-making article for the study of floral morphology: by 
checking carefully the symmetry of each of the four verticils in 
Schizanthus, he arrived at the conclusion that the So/anaceae 
display a unique type of oblique zygomorphy, different from the 
median one characteristic of the Scrophulariaceae, Labiatae, etc.; 
in his own words: “. . .Der Unterschied in der Bliitenbildung 
zwischen den Solaneen mit symmetrischer Bliithe und den Antir- 
hineen (Labiaten, etc.) ist ein sehr wesentlich.”; Wydler was later 
fully supported by Eichler (1869: 105; 1875, 1: 203). 
A further step forward towards an understanding of differences 
between these two families, was taken by Vesque (1885), Schlepe- 
grell (1892) and Fedde (1896), using an anatomical approach; 
these authors were able to show convincingly that the presence of 
internal phloem is a characteristic feature shared by all genera of 
Solanaceae, while the absence of it is a significant characteristic of 
the Scrophulariaceae. These findings supported the inclusion of 
Salpiglossideae in the Solanaceae, as Wettstein (1891:34) had 
already proposed on the basis of external morphology. 
