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In the genus Agrobacterium, terrestrial and plant pathogenic species and marine star-shaped-aggregate-forming species were reported (Conn, 1942; Jordan, 1984; Ophel and Kerr, 1990; Ruger and Hofle, 1992; Starr and Weiss, 1943). The phylogenetic positions of the terrestrial and plant pathogenic species have already been made clear, but those of the marine species have not been published yet. Since Stapp and Knosel (1954) grouped a marine species as Agrobacterium stellulatum based on their formation of star-shaped aggregation, many marine isolates resemble to A, stellulatum as a species of the genus Agrobacterium (Ahrens, 1968; Ahrens and Rheinheimer, 1967). However, intrageneric relationships between marine and terrestrial Agrobacterium species remain uncertain. Ruger and Hofle (1992) performed a phenotypic analysis on these Agrobacterium species in addition to their marine isolates, and they insisted that among these marine strains, five species could be recognized as a distinct taxon. They believe these five species form a group independently from the terrestrial Agrobacterium species, but there were no distinctive phenotypic characteristics to separate the marine species from the genus Agrobacterium. Therefore, they concluded that the genus Agrobacterium must be divided into two subdivisions, accommodating the terrestrial and plant pathogenic species in subdivision 1 and the marine star-shaped-aggregate-forming five species Agrobacterium atlanticum, Agrobacterium ferrugineum, Agrobacterium gelatinovorum, Agrobacterium meteori and A. stellulatum in subdivision 2, until further taxonomic data are available (Ruger and Hofle,1992). In this communication, we determined 165 ribosomal RNA gene sequences of these marine subdivision species, including the non-validated species "Agrobacterium agile" and "Agrobacterium kieliense ," and investigated their phylogenetic position to clarify the classification of marine Agrobacterium species. The bacterial strains used in this study were A. atlanticum IAM 14463T, A, ferrugineum IAM 12616T, A. gelatinovorum IAM 12617T, A. meteori IAM 14464T, A. stellulatum IAM 12614, A. stellulatum IAM 12621T , "A. agile" IAM 12615 and "A. kieliense" IAM 12618. All the strains were grown aerobically on the plate of Difco marine agar 2216 for 2 days at 25°C. Cells grown in the late exponential phase were harvested, washed with sterile water and stored at -20°C until use. The 165 rRNA gene was amplified by PCR using Takara 7 aq (Takara Shuzo, Kyoto, Japan), the three forward primers 8F (5'-AGAGTTTGATCCTGGCTCAG-3'), 520F (5'-CAGCAGCCGCGGTAATAC-3') and 926F (5'-AAACTCAAAGGAATTGACGG-3'), and three reverse primers 704R (5'-TCTACGCATTTCACC-3'), 1110R (5'GGGTTGCGCTCGTTG-3') and 1510R (5'-GGCTACCTTGTTACGA-3') [Escherichia coli numbering system], by the combination of 8F-704R, 520F1110R and 926F-1510R. Amplified fragments were cloned using an Invitrogen TA CloningTM Kit (Invitrogen, CA, U.S.A.). Extraction and purification of the plasmids were carried out using a QlAprep Spin Plasmid Kit (Qiagen, Hilden, Germany). The cloned plasmids were used for sequencing. Sequencing was carried out using an ABI PRISMTM Dye Primer Cycle Sequencing Ready Reaction Kit (Perkin-Elmer Co., CA, U.S.A.) and a model ABI 3735 automatic DNA sequencer (Perkin-Elmer Co.). Previously published 165 rRNA gene sequences, which were already aligned, * Address reprint requests to: Dr . Akira Yokota, ular and cellular Biosciences, The University Yayoi; Bunkyo-ku, Tokyo 113, Japan. Institute of Molec-

Phylogenetic position of the marine subdivision of Agrobacterium species based on 16S rRNA sequence analysis.