A Redescription of the Genus Parathelohania
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Agricultural Research Service U.S. DEPARTMENT OF AGRICULTURE CONTENTS Page Abstract 1 Introduction ............................................. 2 Methods ................................................. 3 Light microscopy ..................................... 3 Scanning electron microscopy . .. .. 4 Transmission electron microscopy ..................... 4 Results .... , ....................... , ..................... , . 5 Parathelohania Codreanu 1966 ........................ 8 Parathelohania legeri (Hesse 1904) ................... 8 Parathelohania anophelis (Kudo 1924) ................ 10 Pa1'(l.thelohania obesa (Kudo 1924) .................... 14 Parathd:hania illinoisensis (Kudo 1921) .............. 16 Pa'rathelohania anornala (Sen 1941) ................... 18 Parathelohaniaindica (Kudo 1929) .................... 18 Pamthelohania obscura (Kudo 1929) .................. 19 Parathelohania pericu.losa (Kellen and Wills 1962) ..... 19 Parathelohania ajl"icanus sp. n ......................... 20 Par'2thelohania octolagenella sp. n ..................... 22 Literat\,re cited ........................................... 24 Achllowledgments ........................................ 26 Trade names are used in this publication solely for the purpose of providing specific information. Mention of a trade name does not constitute a guarantee or warranty of the product by the U.S. Department of Agriculture or an endorsement by the Department over other products not mentioned. Washington, D.C. Issued December 1974 For sale by the Superintendent of Documents. U.S. Government Printing Office, Washington, D.C.- P<ice 40 cents Stock Number (j 100 - 03364 A Redescription of the Genus Parathelohania Codreanu 1966 (Microsporida: Protozoa) With a Reexamination of Previously Described Species of Thelohania Henneguy 1892 and Descriptions of Two New Species of Parathelohania From Anopheline Mosquitoes By EDWIN I. HAZARD and DARRELL \Y. ANTHONY, 1'esea1'ch entomologists, Insects Affecting iVlan Research Labomt01'Y, Agricultuml Research Serv ice, U.S. Department of Agriculture, Gainesville, Fla. ABSTRACT The genus Parathelohania is redescribed as representing species that, in mosquito larvae, have eight characteristically ridged spores enclosed in a persisting sporont membrane and that, in adult female mosquitoes, have smooth-walled spores not enclosed in a sporont membrane. The type species is Parathelohania legM'i (Hesse) from Anopheles rnaculipennis Meigen. Anopheline mosquitoes are the most common hosts of these microsporidian parasites. Two new species of Paratheloha,nia from Anopheles garnbiae Giles and Ano pheles pretoriensis (Theobald) al'e described. The species name "illinoisensis Kudo" is restored to represent the Parathelohania species in Anopheles punctipennis (Say). All representative species of Parathelohania and their synonymies are listed. The revised or ganization of Parathelohania was made possible by new biological information and new diagnostic characters provided by h'ansmis sion electron microscopy and confirmed by scanning electron micro scopy. Photomicrographs of spores from host larvae (made with light microscopy and scanning and transmission electron micro scopy) are presented to show the characters that distinguish the two new species and three species described earlier. 1 2 TECHNICAL BULLETIN 1505, U.S. DEPT. OF AGRICULTURE INTRODUCTION Thelohania legeri Hesse (1904a) 1 was the first microsporidium known from anopheline mosquitoes and was reportedly taken from larvae of Anopheles rnaculipennis Meigen collected in France. A second species, from the larvae of Anopheles lJUnctipennis (Say) collected in Illinois, U.S.A., was described by Kudo (1921) and named Thelohania illinoisensis. After examining what he believed were two additional host species, Anopheles cTucians Wiedemann and Anopheles quadrimaculatus Say, Kudo (1924a) made T. illin oisensis a synonym of T. legeri. In the same year Kudo (1924b) described three additional microsporidia from N ore1. American anophelines, which he named Nosema anophelis, Thelohania obesa, and Thelohania pyriformis. The host of N. anophelis was A. quadri maculatus, and both late instal' larvae and adults were found to have stages of the microsporidium. The Anopheles species that were hosts of the two Thelohania species were undetermined. Missiroli (1929) described a second species from A. maculipennis, but unlike T. lege?'i, which occurs in the fat body of larvae, this species was found in the ovaries and eggs of adult females. Missiroli named it Thelohania grassii after Grassi (1901), who first observed the spores in adult females collected in Italy. Kudo (1929) described two new Thelohania species from ano pheline larvae collected in India: Thelohania indica from Anopheles hyrcanus (Pallas) and Thelohania obscum from Anopheles varuna Iyengar. In the same paper he added the mosquitoes Anopheles barbi1'ostris Van del' Wulp, Anopheles annularis Van der Wulp (=A. fuliginosus of Kudo), Anopheles ramsayi Covell, and Ano pheles subpictus Grassi, as well as both A. varuna and A. hyrcanus, to the list of hosts of T.legeri. Sen (1941) reported another species, Thelohania anomala, from an Indian anopheline, A. ramsayi. Sen also added Anopheles vagus Donitz to the list of hosts of T. Zegeri, and Tour et al. (1971) included Anopheles labmnchiae atroparvus Van Thiel. The last Thelohania to be described from anopheline mosquitoes was Thelohania periculosa Kellen and Wills (1962), from the adipose tissue of the larvae of Anopheles franciscanus McCracken collected in California, U.S.A. Later, Kellen et al. (1967) described Nosema chapmani, from adult female A. francis canus, but Hazard and Weiser (1968) have since demonstrated that the sporonts and cylindrical spores are the stages ofthe transovarial cycle of T. periculosa. Codreanu (1966), believing that Thelohania octospom Henne guy (1892) was the type species of Thelohania and noting that it J Dates in parentheses refer to "Literature Cited," p. 24. REDESCRIPTION OF PARATHELOHANIA 3 has spores with filamentous appendages, chose Thelohania legeri Hesse as the type species for a new genus he named Parathelohania. Codreanu was not aware that Gurley (1893) many years before had designated Thelohania giardi Henneguy (1892), which does not have spore appendages, as the type species of Thelohania. Cod reanu's mistake, however, did not produce complications for the taxonomy of Microsporida because those Thelohania found in Ano pheles mosquitoes are sufficiently different from the Thelohania species described from decapod crustaceans by Henneguy. The spores of T. legeri and other species in anopheline mosquitoes havn a characteristic constricted posterior end, and this difference, as well as other characteristics, justifies their placement in a genus separated from Thelohania. The original problem Codreanu at tempted to resolve, however, remains untouched by his changes in the generic placement of T. legeri. T. octospora should now be placed in a new genus if Codreanu or other investigators continue to believe that spore appendages are important generic characters. Pleistophora and Nosema diseases have been reported in colon ized anopheline mosquitoes by Canning (1957), Fox and Weiser (1959), Vavra and Undeen (1970), Canning and Hulls (1970), and Hazard (1970) ; these are well-established Nosema and Pleistophora species and do not need further discussion in this paper. Some spe cies formerly placed in the genus Thelohania, however, must be re viewed here because of the new information presented by Hazard and Weiser (1968) concerning the unusual sporonts that produce spores in adult female mosquitoes unlike those commonly known in male mosquito larvae and because of new diagnostic characters found on and in the spores from male larvae in scanning and trans mission electron photomicrographs. The sporonts of all Parathelo hania species in male larvae are similar and do not pl·esent diag nostic characters for the separation of species. The spores from these larvae, however, and the shapes and tissue sites of the spores in adult females have many structural characters that can be used as criteria for species differentiation. A complete description of the genus Parathelohania is presented, and two new species are described. Synonymies are also listed for existing species. METHODS Light Microscopy Infected mosquito larvae were cut into small pieces for preparing Giemsa-stained and Heidenhain's hematoxylin-stained smears. Giemsa smears were prepared by smearing the infected tissue on a 4 TECHNICAL BULLETIN 1505, U.S. DEPT. OF AGRICULTURE microscope slide, fixing with 95 % methanol for 1 min, staining with a 10 % Giemsa solution, made with buffered