Microbial mats of coexisting bacteria and archaea date back to the early Ar
chaean: many of the major steps in early evolution probably took place with
in them. The earliest mats may have formed as biofilms of cooperative chemo
lithotrophs in hyperthermophile setting-a, with microbial exploitation of d
iversifying niches. Anoxygenic photosynthesis using bacteriochlorophyll cou
ld have allowed mats, including green gliding bacteria, to colonize anaerob
ic shallow-water mesothermophile habitats. Exploitation of the Calvin-Benso
n cycle by purple bacteria allowed diversification of microbial mats, with
some organisms in more aerobic habitats, while green sulphur bacteria speci
alized in anaerobic niches. Cyanobacterial evolution led to more complex ma
ts and plankton, allowing widespread colonization of the globe and the crea
tion of further aerobic habitat. Microbial mat structure may reflect this e
volutionary development in broad terms, with anaerobic lower levels occupie
d by archaeal and bacterial respirers, fermenters and green bacteria, while
the higher levels contain aerobic purple bacteria and are dominated by cya
nobacteria. A possible origin of eukaryotes is from a fusion of symbiotic p
artners living across a redox boundary in a mat. The geological record of A
rchaean mats may be present as isotopic fingerprints: with the presence of
cyanobacteria, mats may have had a nearly modern structure as early as 3.5
Ga ago (1 Ga = 10(9) years).