Animalbased Food

Like a large number of herbivores and detritivores (e.g., Hoffman and Payne, 1969), many cockroaches incorporate animal tissue into their diet when the opportunity arises. Parcoblatta uhleriana has been observed feeding on mammalian cartilage (Gorton, 1980), but most records of cockroaches feeding on living and dead vertebrates come from species that dwell in caves (discussed below) and from pest cockroaches. The latter can eat a great deal of flesh, particularly of human corpses. They also nibble on the calluses, wounds, fingernails and toe-nails, eyelashes, eyebrows, earwax, dandruff, eye crust, and the nasal mucus of sleeping individuals, particularly children. At times they "bite savagely," leaving permanent scars (Roth and Willis, 1957; Denic et al., 1997). Most reports are from ships, nursing homes, unhygienic urban settings, and primitive tropical living quarters. See Roth and Willis (1957) for a full roster of these horror stories.

Many cockroaches are equipped for predation: they are agile, are aggressive in other contexts, have powerful mandibles, and possess spined forelegs to help secure prey. The recorded victims of cockroaches include ants, parasitic wasps, Polistes larvae, centipedes, dermestids, aphids, leafhoppers, mites, and insect eggs (Roth and Willis, 1960). Both B. vaga and B. asahinai eat aphids and are considered generalist predators (Flock, 1941; Persad and Hoy, 2004). Periplaneta americana has been observed both catching and eating blowflies in a laboratory setting (Cooke, 1968), and pursuing and capturing termite de-alates in and around dwellings. They pounced on termites from a distance of 5 cm, and followed them into crevices in the floor (Annandale, 1910; Bowden and Phipps, 1967). Cockroaches that feed on guano, leaf litter, or epi-phylls also ingest the invertebrate microfauna that inhabit their primary food source (WJB, pers. obs). Dead invertebrates are scavenged by Blattella karnyi (Roth and Willis, 1954b), Parcoblatta pennsylvanica (Blatchley, 1920), and P.fuliginosa (Appel and Smith, 2002), among others. "The insect collector will often find that cockroaches, particularly in the tropics, will play sad havoc with his dead specimens" (Froggatt, 1906).

There are a few instances of cockroaches harvesting the secretions and exudates of heterospecific insects. Several are known to feed on honeydew (e.g., Eurycotis spp. sipping it from fulgorids—Naskrecki, 2005). Parcoblatta pennsylvanica has been observed feeding on cercopid spittle (Gorton, 1980). Recently two species of Costa Ri-

can Macrophyllodromia were observed grazing the white, waxy secretion on the tegmina of at least two species of Fulgoridae (Fig. 4.6) (Roth and Naskrecki, 2001).

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