In surveying the literature on this subject we have collected a far more extensive list of animals and plants associated with cockroaches than one might have expected from an examination of any one of the previous papers on this subject. In our review of the medically important organisms associated with the Blattaria, we pointed out that in addition to many experimental associations cockroaches have been found to harbor, naturally, 4 strains of poliomyelitis virus, about 40 species of pathogenic bacteria, the eggs of 7 species of pathogenic helminths, and to serve as intermediate hosts of 12 other species of helminths pathogenic for vertebrates; cockroaches have also been found to carry, on occasion, 3 species of Protozoa that are pathogenic to man and 2 species of fungi which are sometimes found associated with pathological conditions.
In addition to the above organisms of medical importance, we have compiled records of other organisms, nonpathogenic to vertebrates, which are naturally associated in some way with cockroaches. None of the following numbers can be considered absolute because some names may be synonyms. However, we believe that these figures are very close to the actual numbers of species that have been isolated because we have attempted to refer all obvious synonyms to the currently accepted name for each organism. On this basis there are about 45 species of bacteria, 40 fungi, 6 yeasts, 90 Protozoa, and 45 helminths that have been found associated naturally with cockroaches. Of the arthropods there are about 2 species of scorpions, 4 spiders, 15 mites, 4 centipedes, and 90 insects. Of vertebrates there are 4 species of fish, 16 amphibians, 12 reptiles, 20 birds, and 27 mammals. Besides these there are many records of experimental associations that have been contrived in the laboratory.
Some idea of the increase in our knowledge of the biotic associations of cockroaches, during the last 70 years, may be gathered from a comparison of the above figures with those of Miall and Denny (1886) who presented " ... a long list of parasites which infest the Cockroach." This list included 2 bacteria, 6 Protozoa (some of the names are synonyms), 7 nematodes (some of these names are also synonyms), 1 mite, 1 wasp, and 1 beetle. In addition, they mentioned as other foes of the cockroach: monkeys, hedgehogs, polecats, cats, rats, birds, chameleons, and frogs.
METHODS
We have listed the organisms known to be associated with cockroaches systematically by phylum, class, order, and family. Within each family the organisms are listed alphabetically by genus and species. Under each organism the associated cockroaches are listed as natural or experimental hosts, vectors, or prey. Identified cockroaches are listed by the currently accepted name. Unidentified cockroaches are indicated by the word "Cockroaches." The name of each cockroach is followed by the country in which the observation was made, the authority for the record, and with a few exceptions[3] pertinent biological information, where this is known. Question marks following the names of organisms or countries indicate tentative or questionable identifications.
Records of predators capturing and feeding on cockroaches in zoos and on shipboard we consider natural, even though it is very likely that these particular predators would not normally have access to this prey in nature.
Experimental prey are cockroaches that were fed to predators in the laboratory. Although these predators may have little, if any, access to these cockroaches in nature, we have included such records to indicate the relative acceptability of cockroaches as food by a wide variety of animals.
Records of presumed or known cockroach associates that give no information about an associated cockroach are not included in this review, even though certain of these (e.g., species of Ampulex, Evania, Podium) probably prey upon or parasitize cockroaches exclusively.
The validity of a host-parasite or predator-prey record is dependent upon the accuracy and knowledge of the observer. In assembling these records we have had to accept, in most instances, the identifications of species made by the original authors. However, as a result of our studies on the biology of various species of cockroaches, including some work on their hymenopterous parasites, we have questioned certain records in the literature. Other dubious records which have been perpetuated from one publication to the next, but which apparently were not based on fact, have also been questioned or have been clarified with the aid of specialists in particular groups.