606 PROCEEDINGS OF THE NATIONAL MUSEUM vol.87 



from ranids, or vice versa, in early Cretaceous times. The classifica- 

 tion of Cardioglossa is doubtful and its opalinids are unknown. 



The Ranidae and their Cepedeas and Opalinas may have devel- 

 oped in either Ethiopia or Lemuria. Apparently the Protoopalinas 

 of subgeneric group VIII (ancestors of Cepedea), found now in eastern 

 Asia and Malaysia, passed from Asia-Malaysia to Lemuria. Wliile 

 Cepedea was evolving in ranid hosts and later giving rise to Opalina 

 in the same hosts, the southward spread to Etliiopia may weU have 

 been in progress. 



BUFO (Fig. 157) 



The absence of Bufo from Madagascar and the Seychelles and from 

 Australasia including Papuasia indicates (1) origin in Asia in Creta- 

 ceous times (no longer in communication with Australasia and 

 Lemuria); or (2) origin in Ethiopia later than the early Cretaceous, 

 after Ethiopia and Madagascar had separated; or (3) origin in South 

 America, probably the northwestern highland region. Origin in Asia 

 in the Cretaceous would leave the way open to South America by way 

 of the circum-Pacific land-strip, but if Bufo carried Cepedea with it 

 there would be certain difficulties, to be discussed later. If it arose 

 in northwestern South America, the same circum-Pacific land-strip 

 could have carried it to eastern Asia. If it arose in Ethiopia this 

 must have been after the isolation of Madagascar in the middle 

 Cretaceous. Bufo did not enter South America from North and 

 Central America after Rana and its Opalinas were in these northern 

 lands, or it would have carried Opalina, if indeed Rana itself would 

 not have accompanied it, and neither Opalina nor Rana is in South 

 America, except one species of Rana (palmipes) , which seems to be a 

 late Pliocene or Pleistocene immigrant. 



A little discussion of the origin and distribution of Cepedea is neces- 

 sary at this point. It is found today throughout the earth where 

 climate is suitable, except in (1) Australasia, including Papua and 

 New Zealand, (2) northeastern United States, and (3) northern South 

 America (probably merely not yet reported here). Cepedea evolved 

 from Protoopalina through species resembling those now found in 

 Asia-Malay asia (P. formosae, P. quadrinucleata, P. axonucleata). 

 This would seem to indicate Asia-Malaysia as its place of origin. 

 If so, how did it reach Lemuria? No Protoopalina is today known 

 from lands reputed to have been once a part of Lemuria (Madagascar, 

 the Seychelles, Ceylon, southern India), except one species in a 

 gastrophrynid from just about the boundary line between southern 

 India and northern India. Similarly Bufo is not known from south- 

 ern Lemurian lands (Madagascar, the Seychelles). 



The data known seem to fit the following hypothesis: (1) Origin 

 of Cepedea in Asia-Malaysia or Lemuria, before Bujo was present 

 there, perhaps in the latest Jurassic or earliest Cretaceous, when 



