Williams - p. 2 
The South American beta anole groups are all invaders from the north. 
There have been multiple invasions and hence a layered succession includes 
older groups with strong or even relict endemism and younger groups in 
which there may even be forms conspecific with those in Central America. 
The southernmost of South American beta anoles - A. meridionalis - 
appears, on Etheridge's osteological characters, to be the most primitive 
of South American betas, a relict of the oldest beta invasion of the 
continent. It is clearly an isolated species deserving a group of its own. 
Etheridge has described as the petersi species group the most primitive 
Central American anoles. Two species of the group occur in South America: 
one, A. biporcatus, extends from Mexico to Ecuador and Venezuela, with a 
distinctive race in southwestern Colombia and western Ecuador. A close 
relative of biporcatus - apollinaris - occurs in the inter-Andean valleys 
of Colombia, perhaps the result of an earlier invasion than that which has 
resulted in only racial difference in the trans-Andean region. The other 
species, A, sulcifrons in Colombia, both west of the Andes and in inter- 
Andean valleys, is the geographic representative of pentaprion, a species 
ranging from Mexico to Panama. A. biporcatus, A. apollinaris and A. sulcifrons 
are probably best compared to West Indian crown anoles. 
A. lemurinus is the Central American-Mexican member of another series 
of three. A. vittigerus replaces it in southeastern Panama and trans-Andean 
Colombia. Trans-Andean Ecuador has a third undescribed member of the series. 
This is another independent invasion of South America. 
The humilis group is distinguished by its peculiar axillary pockets. 
Its South American representative is A. notopholis in trans-Andean Colombia. 
A. notopholis appears to be a grass-bush anole. 
* The four groups - petersi, lionotus, humilis, lemurinus - may be the 
most recent invaders of South America. They appear not to have got very far 
and, though endemic South American species when present are amply distinct, 
their northern relationships are very clear. 
A northern relationship is equally clear for the fuscoauratus group 
(in Williams' usage a more restricted and surely more natural group than 
envisioned by Etheridge). A. fuscoauratus of Amazonia, the Guianas and 
the Atlantic Forest is not easy to separate from the limifrons complex in 
Central America. The trans-Andean populations - probably to be called 
maculiventris - in northwestern Colombia are in contact with undescribed 
species of the limifrons complex and in central Ecuador abut on or inter- 
grade with A. fuscoauratus and are more different from the members of the 
limifrons complex than these are from Amazonian A. fuscoauratus. A boldly 
patterned undescribed fuscoauratoid occurs on Gorgona Island, Another 
quite distinct and also patterned undescribed species occurs locally in 
northern Colombia. A. antonii represents fuscoauratus in the inter-Andean 
valleys of Colombia. 
