Phylogenetic Implications of Grooming Behavior in Cockroaches (Insecta: Blattaria)

The use of behavioral data for the investigation and delineation of evolutionary relationships has been increasingly heuristic (Evans, 1952; Speith, 1952; Alexander, 1962; etc.). In insects, grooming behavior seems particularly useful because it is a widespread and prominent part of their behavioral repertory. Comparative studies of several orders have been based on grooming behavior (Szymanski, 1918; Heinz, 1949; Gangwere, 1958; Jander, 1966; Farish, 1972; Valentine, 1973; Valentine and Glorioso, 1979). Lipps (1973) provides an excellent review of grooming literature. The cockroaches are a diverse group with an extensive nontaxonomic literature, There are many descriptions of the general biology of the order (Gould and Deay, 1938; Roth and Willis, 1954, etc.) and of particular species (Qadri, 1938; Rau, 1940; Dow, 1955; Heslop and Ray, 1959, etc.). Roth and Willis (1954) and McKittrick (1964) summarize, many such papers, especially on biology. Our taxonomy follows McKittrick (1964), and is summarized in Table 1. Unspecified grooming behavior of cockroaches is mentioned by Patton (1941) and Burkholder (1965). Others discuss grooming of particular body parts (Haber, 1920; Mote, Wilcox, and Davis, 1926; Roth and Willis, 1952 and 1954; and Eisner, 1961). A few authors are more detailed: Gangwere (1958), Yoshikawa (1958), Eaton and Farley (1969), and Lipps (1973). The most complete description of roach grooming is by Turner (1913), who describes in excellent detail antennal, palpal, and leg cleaning using the mouth, and the use of a leg to rub the base of the antenna and dorsal surface of the abdomen, all in a roach he called Periplaneta orientalis, now placed in the genus Blatta.