The conservation of the core structure and diversification of the external features among the turreted reoviruses appear to be relevant to structural evolution in facilitating the infection of diverse host species. The structure of Rice ragged stunt virus (RRSV), in the genus Oryzavirus of the family Reoviridae, is determined to show a core composed of capsid shell, clamps, and long turrets. The RRSV core structure is equivalent to the core structure of Orthoreovirus and the virion structure of Cytoplasmic polyhedrosis virus (CPV). In RRSV, five peripheral trimers surround each long turret and sit at the Q trimer position in the T31؍l icosahedral symmetry, a structural feature unique to turreted reoviruses. That is, the core of RRSV is partially covered by 60 copies of the peripheral trimer. In contrast, the core of Orthoreovirus is covered by 200 copies of the trimer that sit at the Q, R, S, and T trimer positions. Our results suggest that among the three viruses, RRSV has a structure intermediate between that of Orthoreovirus and the CPV virion. This conclusion coincides with the results of the phylogenetic analysis of amino acid sequences of RNA-dependent RNA polymerases.Reoviridae is the largest and most diverse family of doublestranded RNA (dsRNA) viruses. It includes 12 established genera, namely Aquareovirus, Coltivirus, Cypovirus, Fijivirus, Idnoreovirus, Mycoreovirus, Orbivirus, Orthoreovirus, Oryzavirus, Phytoreovirus, Rotavirus, and Seadornavirus. The hosts of these viruses include plants, vertebrates, insects, and fungi (22). All known viruses in this family are 600 to 800 Å in diameter and consist of an inner core that is surrounded by a few layers of protein, with the exception of the single-layered Cytoplasmic polyhedrosis virus (CPV), which encapsidates 9 to 12 segments of dsRNA and the enzymes involved in transcription. Whereas the precise morphology varies among genera, the morphologies of the innermost capsid shells are similar in spite of the absence of significant sequence homology among component proteins. The conserved innermost capsid of reoviruses is composed of 120 copies of thin crescent-shaped proteins, and the respective subunits exhibit similar overall folding, with substantial modifications that appear to have developed during viral evolution (11,24,28,37,42). Except in CPV, the innermost capsid shell is covered by additional outer layers, the organization and structure of which vary among the genera in the family. The outer capsid shell appears to play important roles in maintaining the stability of the thin innermost capsid shell and sequestering the dsRNA genome, as well as in conferring host specificity and mediating entry into host cells.Reoviruses can be divided structurally into two subgroups, the turreted and nonturreted reoviruses, on the basis of a critical structural feature. Members of the seven genera Aquareovirus, Cypovirus, Fijivirus, Idnoreovirus, Mycoreovirus, Orthoreovirus, and Oryzavirus are classified as turreted reoviruses (22), with distinctive pentameric tu...