.--Krause (Die Columella der VSgel, Berlin, R. Friedlinder and Sohn, 1901)
examined the form of the avian bony stapes in a number of groups, but no comprehensive
study has yet been achieved. As I reported earlier (Feduccia, Auk 91:427429, 1974),
most of the nonpasserine orders, and the entire "oscine" passerine assemblage of birds
possess a bony stapes resembling the primitive reptilian condition, with a flat footplate,
and a straight bony shaft. This type of stapes (Fig. 1) no doubt represents the retention
of the primitive condition. Both the New and Old World suboscines possess a charac-
teristic derived morphology of the bony stapes, and this I interpreted (op. cit.) as
evidence for common ancestry of the Old and New World groups. In the examination of
the coraciiform families (to be presented in detail later) I discovered that the Hoopoe
(Upupidae: Upupa epops) and the wood-hoopoes (Phoeniculidae: Phoeniculus purpu-
reus and Rhinopomastos cyanomelas) also possess a common derived morphology of the
bony stapes which is found in no other of the more than 1000 species I have examined.
This type of stapes (Fig. 1) is characterized by a flat bony footplate, but with a short
but wide shaft that bifurcates into 2 processes. There is a long, laterally directed thin
process, and a shorter, broad process, both of which connect to the tympanic membrane
via extracolumellar ligaments, and no doubt function in a complex lever system.
The possession of this bizarre type of stapes in both the Upupidae and Phoeniculidae
I interpret as a strong indication of monophyly of the 2 groups that points a fortJori to
September 1975 ß GENERAL NOTES 417
i
FIC. l. Alizarin-stained stapes of A, Upupa epops (Upupidae); B, Phoeniculus pur-
pureus (Phoeniculidae) ; C, Leptosomus discolor (Leptosomatidae) ; D, Coracias hoevia
(Coraciidae); and E, Berenicornis albocristatus (Bucerotidae). The stapes of Phoenicu-
lus is slightly damaged. C, D, and E represent the primitive condition for birds; Colius
has a similar stapes. All app;'oximatdy X 10-30.
their uniqueness. Because of this very unusual common character, and because there are
few "good" characters to ally them with the other members of the Coraciiformes, I sug-
gest that they are best lreated as 2 families within an order Upupiformes, as Stresemann
(Aves In Handbuch du Zoologic, vol. 7, No. 2, W. deGruyter, Berlin, 1927-34) sug-
gested, but that the hornbills (Bucerotidae) not be included within the order until
evidence for evolutionary affinity can be aptly demonstrated. Whether or not the wood-
hoopoes and Hoopoe are considered as separate famihes or subfamilies of a Upupidae
is a matter of arbitrary decision.
This research was supported by a grant from the University of North Carolina Research
Council. R. W. Storer (Univ. of Michigan), and R. L. Zusi (National Museum of Natural
History) kindly permitted me to use collections under their care. ALa FouccIa, Dept.
o! Zoology, Univ. o! North Carolina, Chapel Hill 27514. Accepted 19 Dec. 1974.