FUNGI ASSOCIATED WITH COCKROACHES
The use of the asterisk (*) is explained in footnote [3], page [4].
Phylum THALLOPHYTA
Class FUNGI IMPERFECTI
Order MONILIALES
Family PSEUDOSACCHAROMYCETACEAE
Candida zeylanoides (Castellani) Langeron and Guerra
Natural host.—Oötheca of Blatta orientalis, Italy (Ronzoni, 1949).
Torulopsis sp.
Natural host.—Oötheca of Blatta orientalis, Italy (Ronzoni, 1949).
Family MONILIACEAE
Spicaria prasina (Maublanc) Sawada
Natural host.—Ischnoptera rufa rufa, Puerto Rico (Wolcott, 1950): A dead specimen of this cockroach was found stuck to a leaf and covered with this fungus.
Aspergillus flavus Link
Natural hosts.—Oöthecae of Blattella germanica and Eurycotis floridana, U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1952): On outer surface. Determination by Miss Mary Downing.
Oöthecae of Periplaneta americana, U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1952): Inside oöthecae. Determination by Miss Mary Downing.
* Aspergillus fumigatus Fresenius
Natural vector.—Blatta orientalis, France (Sartory and Clerc, 1908): From intestine.
* Aspergillus niger van Tieghem
Natural vector.—Periplaneta americana, U.S.A., Texas (McShan in Roth and Willis, 1957a): From feces.
Experimental vector.—Blatta orientalis, Italy (Cao, 1898): Organism passed unchanged through the gut of the insects.
Aspergillus sydowi (Bainier and Sartory) Thom and Church
Natural host.—Oötheca of Eurycotis floridana, U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1952): On outer surface. Determination by Miss Mary Downing.
Aspergillus tamarii Kita
Natural host.—Oöthecae of Blattella germanica, U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1952): On exterior surface. Determination by Miss Mary Downing.
Aspergillus sp.?
Natural and experimental vector.—Blattella germanica, on shipboard (Morrell, 1911): Isolated from feces. Experimentally Morrell also showed that the spores of the fungus could be recovered from feces of cockroaches that had fed on them.
Aspergillus sp.
Natural vector.—Periplaneta americana, England (Bunting, 1956): The fungus was isolated mostly from imperfectly excreted feces.
Beauveria bassiana (Balsamo) Vuillemin
Experimental host.—Blattella germanica and Periplaneta americana, U.S.A. (Dresner, 1949, 1950): The nymphs of American cockroaches became infected when they (1) were injected with a 1-percent suspension of spores, (2) ate rat pellets sprayed with the spore suspension, or (3) were dusted with the fungus spores. The symptoms of the fungus infection were paralysis followed by death; some of the infected insects liquefied, others dried up after the appearance of a subcuticular blackening.
Cephalosporium sp.
Natural vector.—Periplaneta americana, U.S.A., Texas (McShan, unpublished MS., 1953): From feces of cockroaches collected in the basement of a grain elevator at the docks in Galveston.
* Geotrichum candidum Link
Experimental vector.—Blatta orientalis, Italy (Cao, 1898): Organism retained its pathogenicity after passing through the insect's gut.
Penicillium sp.
Natural vector.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): From feces.
Periplaneta americana, England (Bunting, 1956): Mostly from imperfectly excreted feces.
Metarrhizium anisopliae (Metschnikoff) Sorokin
Natural hosts.—Blattidae, Seymour (1929); Charles (1941).
Panesthia australis, U.S.A., Massachusetts (Roth and Willis, unpublished data, 1957): Growing on adult specimens that were found dead in a laboratory colony. Determination by Miss Dorothy Fennell.
Periplaneta americana, England (Bunting, 1956): Growing on genitalia of females where it prevented oöthecal formation.
Cockroach, Puerto Rico (Johnston, 1915): From a "small roach" in the pathological collection at Rio Piedras (no data).
Family DEMATIACEAE
Memnoniella echinata (Rivolta) Galloway
Natural host.—Oötheca of Blattella germanica, U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1952): On material that had oozed from a damaged oötheca. Determination by Miss Mary Downing.
Torula acidophila Owen and Mobley
Natural host.—Periplaneta americana, U.S.A. (Owen and Mobley, 1948): The digestive tract of this cockroach is the normal habitat of this yeast which was transmitted to sirup by the insects. The yeast superimposed a foreign taste, suggestive of malic acid, upon the original flavor of the sirup.
Torula gropengiesseri Lodder
Natural host.—Blatta orientalis, Germany (Gropengiesser, 1925; Lodder, 1934): Isolated from fat body and oöthecae. Gier (1947) is of the opinion that the so-called yeasts that supposedly may displace the bacteroids in the fat body (Mercier, 1907b; Gropengiesser, 1925) may actually represent poorly fixed and insufficiently stained bacteroids.
Torula rosea Preuss
Experimental host.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): Upon repeated feeding of massive doses of this yeast to the cockroach, these workers were able to isolate the organism from the feces up to six days thereafter. There was no evidence that T. rosea was pathogenic for B. craniifer.
Class PHYCOMYCETES
Order MUCORALES
Family MUCORACEAE
Mucor guilliermondii Nadson and Filippov
Natural host.—Periplaneta americana, U.S.S.R. (Nadson and Filippov, 1925; Filippov, 1926): Isolated and cultured from intestine.
Mucor sp.
Natural host.—Oötheca of Periplaneta americana, U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1952): Inside oötheca. Determination by Miss Mary Downing.
Pycnoscelus surinamensis, Germany (Bode, 1936): Isolated from fat body which it had stained red.
Rhizopus nigricans Ehrenberg
Natural vector.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): From feces.
Rhizopus sp.
Natural vector.—Periplaneta americana, U.S.A., Texas (McShan, unpublished MS., 1953): From feces.
Syncephalastrum sp.
Natural vector.—Periplaneta americana, U.S.A., Texas (McShan, unpublished MS., 1953): From feces.
Order ENTOMOPHTHORALES
Family BLASTOCYSTIDACEAE
Blastocystis hominis Brumpt
Natural vector.—Blatta orientalis, U.S.S.R. (Zasukhin, 1930): In hind gut in 40 percent of over 3,000 cockroaches.
Blastocystis sp.
Natural vectors.—Blatta orientalis, U.S.S.R. (Yakimov and Miller, 1922): Found in the intestinal contents of 29 percent of 124 B. orientalis.
Cockroaches, Venezuela (Tejera, 1926).
The placement of the following fungus is problematic.
Coccidioides periplanetae Avrech
Natural host.—Blatta orientalis, Germany (Avrech, 1931): Found in cells lining the lumen of midgut and caeca. The whole upper part of the epithelium was filled with sporangia and spores.
Class ASCOMYCETES
Order ENDOMYCETALES
Family SACCHAROMYCETACEAE
Saccharomyces cerevisiae Hansen
Natural vector.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): In feces.
Saccharomyces sp.
Natural vector.—Blattella germanica, U.S.A. (Janssen and Wedberg, 1952): Found consistently in alimentary tract of B. germanica fed sucrose solutions.
Order HYPOCREALES
Family HYPOCREACEAE
Cordyceps amazonica Hennings
Natural host.—Cockroaches, British Honduras (Mains, 1940).
Cordyceps blattae Petch
Natural host.—Blattella germanica, Ceylon (Petch, 1924): Collected at Hakgala twice. A slight covering of brown mycelium overran the insect and fastened it to the underside of a living leaf.
Order LABOULBENIALES
Family LABOULBENIACEAE
Herpomyces amazonicus Thaxter
Natural host.—Nyctibora obscura, Brazil, Natal (Thaxter, 1931): On antennae.
Herpomyces anaplectae Thaxter
Natural hosts.—Anaplecta sp., Venezuela, Caracas (Thaxter, 1905, 1908); Trinidad (Thaxter, 1931): On antennae.
Cockroach, Sumatra (Thaxter, 1931).
Herpomyces appendiculatus Thaxter
Natural host.—Platyzosteria scabra, Australia, N.S.W. (Thaxter, 1931): On antennae.
Herpomyces arietinus Thaxter
Natural hosts.—Ischnoptera sp., U.S.A., Georgia (Thaxter, 1908).
Parcoblatta uhleriana, U.S.A., Massachusetts (Roth, unpublished data, 1957): The nymphs were in a culture of Parcoblatta virginica which was infected with this fungus; it is possible that these P. uhleriana became infected by contact with P. virginica. Fungus identified by Dr. R. K. Benjamin.
Parcoblatta virginica, U.S.A., Massachusetts (Roth, unpublished data, 1957): Fungus determined by Dr. R. K. Benjamin. Fungus found on antennae, palpi, legs, body surface (pl. [27], A).
Parcoblatta sp., U.S.A., Kentucky, Massachusetts (Thaxter, 1902, 1908): On antennae.
It is likely that Thaxter's host records (certainly those assigned to Temnopteryx and possibly those assigned to Ischnoptera) were species of Parcoblatta. Hebard (1917) has shown that all the species referred to Ischnoptera in the United States, except I. deropeltiformis, now belong in the genus Parcoblatta. All species originally referred to the genus Temnopteryx in the United States are now synonymized with species of Parcoblatta.
Herpomyces chaetophilus Thaxter
Natural hosts.—Periplaneta americana, Brazil (Thaxter, 1931).
Periplaneta sp., Zanzibar and Mauritius (Thaxter, 1902, 1908): On spines of legs, antennae, and cerci.
Herpomyces chilensis Thaxter
Natural host.—Cockroach, Chile (Thaxter, 1918): On antennae.
Herpomyces diplopterae Thaxter
Natural hosts.—Diploptera punctata, Ascension Island (Thaxter, 1902, 1908): On antennae. This species also was infected experimentally (Richards and Smith, 1954).
Cockroach, Fiji (Thaxter, 1931).
Herpomyces ectobiae Thaxter
Natural hosts.—Blattella germanica, U.S.A., Massachusetts (Thaxter, 1902, 1908); Burma, Tenasserim (Spegazzini, 1915); Argentina, Buenos Aires (Spegazzini, 1917): On antennae. U.S.A., Minnesota (Richards and Smith, 1955): Scattered over entire body, wings. France? (Picard, 1913): On tibial spines. Chile and Philippine Islands (Thaxter, 1931).
"Ectobia" spp., Zanzibar and Saint Kitts, B.W.I. (Thaxter, 1902, 1908): Possibly on species that are now in the genus Blattella rather than in the genus Ectobius as it is known today, because Thaxter also used the synonym Ectobia germanica for the German cockroach, Blattella germanica.
Experimental hosts.—Blattella germanica and Blattella vaga, U.S.A. (Richards and Smith, 1954).
Herpomyces forficularis Thaxter
Natural hosts.—Cockroaches, Mauritius? and Fiji (Thaxter, 1902, 1908, 1931): On antennae.
Herpomyces gracilis Thaxter
Natural host.—Blattella humbertiana, Philippine Islands, Luzon (Thaxter, 1931): On antennae.
Herpomyces grenadinus Thaxter
Natural host.—Cockroach, Grenada, B.W.I. (Thaxter, 1931): On antennae of a "brown wingless blattid."
Herpomyces leurolestis Thaxter
Natural host.—Leurolestes pallidus, British Guiana and Trinidad (Thaxter, 1931): On antennae.
Herpomyces lobopterae Thaxter
Natural host.—Loboptera sp., Argentina (Thaxter, 1931): On antennae.
Herpomyces macropus Spegazzini
Natural host.—Blaberus sp.?, Argentina (Spegazzini, 1917).
Cockroaches, Peru, Puerto Rico, Ecuador, and Haiti (Spegazzini, 1915, 1917): Material previously assigned by Spegazzini (1915) to H. paranensis was also placed by him in this new species. However, Thaxter (1931) believed that H. macropus may be synonymous with H. paranensis, but he provisionally retained H. macropus because he had not seen Spegazzini's material.
Herpomyces nyctoborae Thaxter
Natural hosts.—Nyctibora tomentosa, U.S.A., Texas (Thaxter, 1905, 1908): On antennae. This cockroach is not established in Texas, and the specimen may have been misidentified (Gurney, personal communication, 1958).
Nyctibora sp., Argentina (Spegazzini, 1917): On antennae.
Herpomyces panchlorae Thaxter
Natural hosts.—Panchlora nivea, Trinidad (Thaxter, 1931): On antennae.
Herpomyces panesthiae Thaxter
Natural host.—Panesthia lobipennis, Ceylon (Thaxter, 1915): On antennae.
Herpomyces paranensis Thaxter
Natural hosts.—Blaberus sp.? Brazil (Thaxter, 1902, 1908): On antennae.
Blaberus sp., Brazil and Argentina (Spegazzini, 1917): On antennae.
Cockroaches, Trinidad and Argentina (Thaxter, 1931).
Herpomyces periplanetae Thaxter
Natural hosts.—Blaberus sp.?, Argentina (Spegazzini, 1917).
Blatta orientalis, U.S.A., Massachusetts (Thaxter, 1902, 1908); Locality? (Spegazzini, 1915); France? (Picard, 1913).
Periplaneta americana, Bermuda and U.S.A., Massachusetts (Thaxter, 1902, 1908); Plains of Biajar, Italian Somaliland, and Argentina (Spegazzini, 1915, 1917).
Periplaneta australasiae, Bermuda (Thaxter, 1902, 1908).
Periplaneta brunnea, Brazil (Thaxter, 1931).
Periplaneta sp., Mexico, West Indies, Panama, Brazil, Africa, South Seas, and China (Thaxter, 1902, 1908).
Cockroaches, Belgium (Collart, 1947).
Additional locality records: Grenada, Trinidad, B.W.I., and Tangier (Thaxter, 1931).
The fungus was found growing on spines, tegmina, integument, and antennae, at times abundantly.
Experimental hosts.—All the following data are from Richards and Smith (1954):
Blatta orientalis: A few plants matured.
Neostylopyga rhombifolia: Some development but no mature plants.
Periplaneta americana: Fungus developed prolifically with a density equal to that on original host.
Periplaneta australasiae: Some development but no mature plants.
Periplaneta brunnea: Fungus developed prolifically with a density equal to that on original host.
Herpomyces phyllodromiae Thaxter
Natural host.—"Phyllodromia" sp., Abyssinia (Thaxter, 1905, 1908): On antennae.
Herpomyces platyzosteriae Thaxter
Natural host.—"Eurycotis floridana," Mexico (Thaxter, 1905, 1908): On antennal setae.
Since this cockroach is not found in Mexico (J. A. G. Rehn, personal communication, 1957), E. floridana is undoubtedly not the host for this fungus. W. B. Brown (personal communication, 1957) searched the cockroach collection at the Museum of Comparative Zoology but was unable to find Thaxter's insect for reidentification.
Herpomyces stylopygae Spegazzini
Natural hosts.—Blatta orientalis, Argentina (Spegazzini, 1917); U.S.A. (Richards and Smith, 1955a).
Experimental hosts.—Neostylopyga rhombifolia (Richards and Smith, 1954): A few plants matured.
Pycnoscelus surinamensis (Richards and Smith, 1954): Some development but no mature plants.
The fungus (fig. 1) is found on antennae (pl. [27], B, C), palpi, cerci, and femurs. Thaxter (1931) believed H. stylopygae to be synonymous with H. periplanetae. However, Richards and Smith (1954) concluded that H. stylopygae would not grow on P. americana under their laboratory conditions although H. periplanetae would grow on B. orientalis. This indicated a strain or species difference between the two fungi. Gunn and Cosway (1938) reported a species of Stigmatomyces on the antennae of B. orientalis; this fungus was probably H. stylopygae (Richards and Smith, 1956).
Herpomyces supellae (Thaxter)
Natural host.—Supella supellectilium, Trinidad (Thaxter, 1931): On antennal spines.
Herpomyces tricuspidatus Thaxter
Natural hosts.—Blaberus craniifer, U.S.A., Key West (Richards and Smith, 1955).
Blaberus sp. and Epilampra? sp., Panama (Thaxter, 1902, 1908).
Epilampra sp., Saint Kitts, B.W.I., and Haiti (Thaxter, 1902, 1908).
Leucophaea maderae, Fernando Po (Spegazzini, 1915).
Nauphoeta cinerea, Brazil (Thaxter, 1931).
Cockroaches, China? (Thaxter, 1902); Philippine Islands, Mindanao (Thaxter, 1931).
Experimental hosts.—Blaberus craniifer, U.S.A. (Richards and Smith, 1955).
Infections on the antennae. Richards and Smith (1954) were unable to secure experimental infections in L. maderae with H. tricuspidatus. Experiments with N. cinerea showed some development but no mature plants although identification of the growing fungus was uncertain because of simultaneous exposure to H. ectobiae, H. stylopygae, and H. tricuspidatus.
Herpomyces zanzibarinus Thaxter
Natural hosts.—Eurycotis manni, Brazil (Thaxter, 1931): On antennae.
Gyna sp.?, Isle of Nias (Spegazzini, 1915): On antennae.
Cockroach, Zanzibar (Thaxter, 1902): On antennae.