Do microbes have macroecology? Large-scale patterns in the diversity and distribution of marine benthic ciliates


Andrey Azovsky, Dept. of Hydrobiology, Moscow State University, Moscow 119899, Russia. E-mail:


Aim  The biogeography and global distribution of protists has long been disputed, with two primary, opposing views. To test these two sets of views in greater detail, we have compiled the available data for marine benthic ciliates and assessed the general patterns of their diversity and distribution compared with Metazoa.

Location  World-wide.

Methods  A comprehensive database (1342 species, over 350 sources) was used to analyse the diversity, distribution, species occurrences and range size distribution of free-living ciliates that inhabit marine sediments in 17 geographical regions.

Results  Twenty-five per cent of the species have been found in a single region only, whereas 18% are widespread (they occur in more than half the regions covering both hemispheres). Only 5–7% of regional faunas are endemic, which is much lower than for macroorganisms. Regional diversity depends neither on total area nor on coastline length and does not show any obvious latitudinal trends, but correlates highly with the investigation effort expended in a region and (negatively) with the average salinity. A comparison of species composition reveals distinctions between the Arctic Area (the White, Barents and Kara seas), Laurasian Area (north Atlantic, north Pacific and European seas), Gondwanian Area (Southern Ocean) and the Antarctic. No clear geographical correlations are found for faunistic composition at the genus or family levels. There is the tendency to narrow the latitudinal ranges for species found at high latitudes (reversal of Rapoport's rule).

Main conclusions  Undersampling and data insufficiency are the key factors affecting the observed diversity and distribution of microorganisms. Nevertheless, marine benthic ciliates demonstrate certain patterns that generally agree with the ‘moderate endemicity’ model (Foissner, 2004, 2008), but consistently contradict the regularities commonly observed for multicellular taxa. Thus, ciliates do have a biogeography, but their macroecological patterns may be different in some respects from that of macroorganisms.