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Ectosymbioses between invertebrates and sulfur-oxidizing bacteria are widespread in sulfidic marine environments and have evolved independently in several invertebrate phyla. The first example from a freshwater habitat, involving  Niphargus ictus  amphipods and filamentous  Thiothrix  ectosymbionts, was recently reported from the sulfide-rich Frasassi caves in Italy. Subsequently, two new  Niphargus  species,  N. frasassianus  and  N. montanarius , were discovered within Frasassi and found to co-occur with  N. ictus . Using a variety of microscopic and molecular techniques, we found that all three Frasassi-dwelling  Niphargus  species harbor  Thiothrix  ectosymbionts, which belong to three distinct phylogenetic clades (named T1, T2, and T3). T1 and T3  Thiothrix  dominate the  N. frasassianus  ectosymbiont community, whereas T2 and T3 are prevalent on  N. ictus  and  N. montanarius . Relative distribution patterns of the three ectosymbionts are host species-specific and consistent over different sampling locations and collection years. Free-living counterparts of T1–T3 are rare or absent in Frasassi cave microbial mats, suggesting that ectosymbiont transmission among  Niphargus  occurs primarily through inter- or intraspecific inoculations. Phylogenetic analyses indicate that the  Niphargus-Thiothrix  association has evolved independently at least two times. While ectosymbioses with T1 and T2 may have been established within Frasassi, T3 ectosymbionts seem to have been introduced to the cave system by  Niphargus .

Repeatedly Evolved Host-Specific Ectosymbioses between Sulfur-Oxidizing Bacteria and Amphipods Living in a Cave Ecosystem