Angiosperm plastid genomes are generally conserved in gene content and order with rates of nucleotide substitutions for protein-coding genes lower than for nuclear protein-coding genes. A few groups have experienced genomic change, and extreme changes in gene content and order are found within the flowering plant family Geraniaceae. The complete plastid genome sequence of Pelargonium X hortorum (Geraniaceae) reveals the largest and most rearranged plastid genome identified to date. Highly elevated rates of sequence evolution in Geraniaceae mitochondrial genomes have been reported, but rates in Geraniaceae plastid genomes have not been characterized. Analysis of nucleotide substitution rates for 72 plastid genes for 47 angiosperm taxa, including nine Geraniaceae, show that values of dN are highly accelerated in ribosomal protein and RNA polymerase genes throughout the family. Furthermore, dN/dS is significantly elevated in the same two classes of plastid genes as well as in ATPase genes. A relatively high dN/dS ratio could be interpreted as evidence of two phenomena, namely positive or relaxed selection, neither of which is consistent with our current understanding of plastid genome evolution in photosynthetic plants. These analyses are the first to use protein-coding sequences from complete plastid genomes to characterize rates and patterns of sequence evolution for a broad sampling of photosynthetic angiosperms, and they reveal unprecedented accumulation of nucleotide substitutions in Geraniaceae. To explain these remarkable substitution patterns in the highly rearranged Geraniaceae plastid genomes, we propose a model of aberrant DNA repair coupled with altered gene expression.comparative genomics ͉ genome evolution ͉ plastid genome A ngiosperm plastid genomes are generally highly conserved in gene order, gene content, and organization (1). Whereas the rates of nucleotide substitutions are highly variable in protein-coding genes of angiosperm nuclear genomes, rates in plastid genes are generally lower (2). Rates of nonsynonomous substitutions (dN), those that cause an amino acid change, are substantially lower than rates of synonymous substitutions (dS), those that do not cause an amino acid change. Aside from a recent report describing elevated dN for a single gene in Oenothera and lineages within Caryophyllaceae (3), plastid genes of photosynthetic plants are under strong purifying selection and the rapid accumulation of either dN or dS has not been described.The plastid genomes of nonphotosynthetic plants reveal accelerated rates of nucleotide substitutions in many proteincoding genes; furthermore, these genomes exhibit extensive gene loss and genome rearrangement (4-6). However, analyses involving either few genes or few taxa for photosynthetic angiosperm plastid genomes generally reveal that modest rate variation is locus-and lineage-specific. A few groups of angiosperms have experienced lineage-specific rate variation, including the lineages leading to the grasses (7), pea (2), Gnetum (8), and Welwitschia...