To get insight into the early evolution of the primate Alu elements, w
e characterized sequences of these repeats from the Malagasy prosimian
s, lemurs (Lemuridae) and sifakas (Indriidae), as well as from galagos
(Lorisidae). These sequences were compared with the oldest Alu specie
s known from the human genome: dimeric Alu J and S and free Alu monome
rs. Our analysis indicates that about 60 Myr ago, before the prosimian
divergence, free left and right monomers formed an Alu heterodimer co
nnected by a 19-nucleotide-long A-rich linker. The resulting elements
successfully propagated in diverging primate lineages until about simi
lar to 20 Myr ago, conserving similar sequence features and essentiall
y the same Alu RNA secondary structure. We suggest that until that tim
e the same ''retropositional niche'', molecular machinery making possi
ble the proliferation by retroposition, constrained the evolution of A
lu elements in extant primate species. These constraints became subseq
uently relaxed. In the Malagasy prosimians the dimeric Alu continued t
o amplify after acquiring a 34- to 36-nucleotide extension of their li
nker segment, whereas in the galago genome the ''retropositional niche
'' was occupied by novel short elements.