Genes are located on chromosomes
The location of genes on chromosomes was concluded by Morgan from his research on fruit fly, Drosophila melanogaster.
Chromosomal theory of inheritance
Walter Sutton and Theodor Boveri studied the behavior of chromosomes and genes during the process of meiosis and proposed the chromosomal theory of inheritance.
Both of them observed that the movement of the chromosome and the gene were similar. Based on their observation, they proposed that genes are located on chromosomes.
Chromosomal theory of inheritance states that the "Mendelian factors or genes are located on chromosomes and it is the chromosomes that segregate and assort independently during meiosis."