MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 
47 
been able to expand and diversify in neighboring regions. The recently described southern Angolan 
endemic Cordylus namakuiyus is a good example of these “exchanges.” Together with 
C. machadoi, its sister species, C. namakuiyus occurs in the “Southern Africa” region sensu 
Linder et al. (2012) (Stanley et al. 2016). However, C. machadoi is restricted to the highlands of 
southern Angola and the northern Nambia escarpment, in what could be considered as the 
“Angolan mountains and high plateaus” zone of the “Angolan-Rhodesian plateau district” sensu 
Crawford-Cabral (1991), whereas C. namakuiyus only occurs in the lowlands of the “Namibe 
desert and subdesert” zone. Another similar example, already explored above, is that of the Namib 
Day Geckos of the genus Rhoptropus, in which the majority of species occur in the “Namibe desert 
and subdesert,” but two taxa have their distributions in the “Angolan mountains and high plateaus” 
zone {R. montanus) and the “Escarpmenf ’ zone {R. benguellensis). 
There are several examples of the so-called paraethiopien amphbians and reptiles identified by 
Monard (1937b, 1938), e.g., Bitis arietans, Varanus niloticus, and Crocodylus niloticus. However, 
recently published and ongoing large-scale phylogeographic studies are starting to deconstruct 
some of these widely distributed taxa, revealing that they comprise numerous cryptic species, each 
with much more modest distributional ranges that can hardly be considered paraethiopien or even 
panethiopien. This is for example the case of Ptychadena mascareniensis, the same example used 
by Monard (1938) as a panethiopian amphibian species. This species complex was recently 
studied by Zimkus et al. (2017), who showed that the Angolan population is one of several differ¬ 
ent lineages contained in the complex. This is also true for reptile species such as Hemidactylus 
mabouia, which is currently under study. Preliminary results shows that it is a complex of several 
different lineages with still large, but considerably more retricted distribution ranges (Ishan 
Agarwal, pers. comm.). 
Recent modeling and statistical approaches to Angolan zoogeography have produced interest¬ 
ing, if limited, insights. Solely based on the data for ungulates compiled by Crawford-Cabral in a 
series of past publications, Rodrigues et al. (2015) were able to retrieve four main biogeographic 
regions for the country; 1) a northern Angola “Zaire-Lunda-Cuanza” region, encompassing the 
blend of humid evergreen and semi-deciduous forests, woodlands, shrublands and grasslands char¬ 
acterizing the Western Congolian forest-savanna mosaic, and including most of the Angolan scarp 
savannas and woodlands, as well as containing the northern parts of the deciduous broadleaf savan¬ 
nas and woodlands of the Angolan Miombo woodlands; 2) a “Central Plateau” area that intergrades 
with the previous region in a north-south gradient, corresponding roughly to the south and south 
eastern parts of the Angolan plateau, particularly dominated by the Angolan Miombo woodlands, 
and related to a more broad Zambezian region; 3) a southern “Cunene - Cuando-Cubango” region, 
comprising most of southern areas of the country with the exception of the Namibe area in the 
southwest, and corresponding to the northern limit of the Kalahari sub-region, and dominated by 
Zambezian Baikiaea woodlands and Angolan mopane woodlands; and finally 4) a “Namibe” region 
comprising the southwestern, low elevation areas of the country, deeply embedded in the Southern 
Africa Kaokoveld desert area, dominated by the Namibe Desert and the Namibian savanna wood¬ 
lands. These four regions can potentially be assigned to some of the divisions proposed by Craw¬ 
ford-Cabral (1991) and, in general, reflect a north-south turnover in the country, with a specific 
Angolan plateau group in the center/south of the country similar to what was proposed by Monard 
(1937b) for reptiles. However, Rodrigues et al. (2015) failed to illustrate the considerable within- 
region diversity proposed by Crawford-Cabral (1991), most likely because of the uneven 
geographic representation of the data, and the limitation of using only ungulates, a group that is not 
speciose and has a potentially weaker biogeographic signal because of high vagility and habitat 
generalism in some cases. 
