162 EDWARD L. RICE 
the great majority make the entire reptilian columella (with the 
possible exception of part of the insertion plate) a derivative of 
the hyoid arch—thus Bender (’11, 712), Cords (09), Gaupp (’98 
and later papers, contrary to his earlier view of a double origin), 
Gregory (13), Howes and Swinnerton (’01), Kingsley (’00), 
Kunkel (711, 712 b), Peyer (12), Schauinsland (’00), Shiino (14), 
Versluys (98). The view is an old one, having been held by 
Rathke (’39, cited by Moller, ’05). The opposite extreme view, 
that the columella is of exclusive otic origin, is held by Moller 
~ (05) and Noack (07). The intermediate view of Parker (’79) . 
and Hoffmann (’89) finds a vigorous supporter to-day in Fuchs 
(06 and later papers). According to this view, the columella 
consists of two genetically distinct divisions—an otostapes, 
derived from the otic capsule, and a hyostapes, of hyoid origin. 
As first suggested by Hoffmann and as used by Fuchs, these 
terms are synonymous with stapes and extracolumella; Versluys 
(03), however, who accepts the terms as representing somewhat 
distinct centers of chondrification, but rejects their genetic 
significance, uses them in a slightly different sense. According 
to his view, the division between the permanently cartilaginous 
extracolumella and the bony stapes of the adult is not coincident 
with the junction of the cartilage nuclei characterizing hyostapes 
and otostapes, but is located slightly nearer to the footplate. 
According to Hoffmann and Fuchs, the processus internus and 
processus dorsalis belong to the hyostapes; according to Versluys, 
they belong to the otostapes; all agree as to their extracolu- 
mellar nature. Conditions in Eumeces seem to accord with the 
usage of Versluys, which I shall follow. 
Three questions, then, demand consideration: 1. Is the hyo- 
stapes genetically related to the hyoid arch? 2. Is the otostapes 
genetically related to the otic capsule? 3. Are the hyostapes 
and otostapes genetically related to one another? 
Is the hyostapes genetically related to the hyoid arch? As already 
noted, a structural connection of the hyostapes with the hyoid 
arch has been observed, at some stage of development, in repre- 
sentatives of all the main groups of living reptiles—Rhyncho- 
cephalia, Chelonia, Squamata (both lizards and snakes), and 
