3Despite the overwhelming bacterial diversity present in the world's oceans, the majority of recognized marine bacteria fall into as few as nine major clades (36), many of which have yet to be cultivated in the laboratory. Molecular-based approaches targeting 16S rRNA genes demonstrate that the Roseobacter clade is one of these major marine groups, typically comprising upwards of 20% of coastal and 15% of mixed-layer ocean bacterioplankton communities (see, e.g., references 36, 37, 42, 98, and 109). Roseobacters are well represented across diverse marine habitats, from coastal to open oceans and from sea ice to sea floor (see, e.g., references 16, 28, 37, 42, 52, and 98). Members have been found to be free living, particle associated, or in commensal relationships with marine phytoplankton, invertebrates, and vertebrates (see, e.g., references 4, 6, 7, 44, 49, 115, and 119). Furthermore, representatives of the clade stand out as representing one of the most readily cultivated of the major marine lineages (36). These isolated representatives are serving as the foundation for an improved understanding of marine bacterial ecology and physiology.
DESCRIPTION OF THE GROUPThe Roseobacter clade falls within the ␣-3 subclass of the class Proteobacteria, with members sharing Ͼ89% identity of the 16S rRNA gene. The first strain descriptions appeared in 1991, about the time that 16S rRNA-based approaches for cataloging microbial diversity were revealing the immensity of prokaryotic diversity in the world's oceans. Interest in the clade has risen steadily since the initial discovery of these strains; at present the clade contains 36 described species, representing 17 genera, and literally hundreds of uncharacterized isolates and clone sequences. The first described members were Roseobacter litoralis and Roseobacter denitrificans, both pinkpigmented bacteriochlorophyll a-producing strains isolated from marine algae (99). Subsequent cultivation of clade members, however, revealed that many strains are neither pink nor bacteriochlorophyll a producers (see, e.g., references 20, 41, 43, and 61). With the exceptions of the described strains of the genus Ketogulonicigenium (113) and several clones from a South African gold mine (GenBank accession numbers AF546906, -13, -17, -22 to -24, and -26), the Roseobacter clade is exclusively marine or hypersaline, with characterized isolates demonstrating either a salt requirement or tolerance (see, e.g., references 60 and 62
ABUNDANCE AND DISTRIBUTION IN MARINE ENVIRONMENTSBased on culture collections, 16S rRNA clone libraries, and single-cell analyses, roseobacters have been identified in most marine environments sampled. The group is prevalent in 16S rRNA gene inventories of seawater (Table 1) and marine sediments (Table 2) and is noticeably absent from analogous inventories of freshwater and terrestrial soil environments. Fluorescent in situ hybridization (FISH) studies quantifying Roseobacter populations in coastal waters of the southeastern United States and the North Sea indicate ...