Comparative analysis of the intrinsic leg musculature of the American cockroach,Periplaneta americana (L.)

The American cockroach has a total of 368 muscles inserting on the post-coxal segments of its legs. By using a narrow morphological definition for delimiting individual muscles, it is shown (i) that the protrochanteral musculatures (23 muscles/leg) differ from the essentially identical meso- and metatrochanteral musculatures (24 and 26 muscles/leg) in number and disposition of extensors and in having a completely different flexor composition, and (ii) that the musculatures of the more distal segments of the legs are completely serially homologous, there being 2 muscles for moving each femur, 23 for each tibia, 7 for each first tarsomere, and 5 for each of the paired pretarsal claws. In all six legs, the trochanteral and tibial musculatures each contain single slender muscles that may be acting proprioceptively to measure the angular displacements between, respectively, the coxas and trochanters, and the femurs and tibias. Neurological and phylogenetic considerations are used to demonstrate why a narrow morphological definition should be employed, and why the widely used functional definition of Snodgrass ('35) is not only fallacious on evolutionary grounds, but also leads to making erroneous conclusions regarding the manner in which insect musculature is controlled by the insect central nervous system. Finally, it is hypothesized that the physiological limitations imposed by having an open circulatory system and the problems inherent in the neural control of large muscles may have been major evolutionary factors in forcing insects to use many slender muscles to control their body movements.