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In Brazilian Lizards. Fallisia Effusa Gen.Nov., Sp.Nov. and Fallisia Modesta Gen.Nov., Sp.Nov

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In Brazilian Lizards. Fallisia Effusa Gen.Nov., Sp.Nov. and Fallisia Modesta Gen.Nov., Sp.Nov Paraaitology (1974), 68, 117-121:; 1717 With 4 plates Further parasites of the family Garniidae (Coccidiida: Haemosporidiidea) in Brazilian lizards. Fallisia effusa gen.nov., sp.nov. and Fallisia modesta gen.nov., sp.nov. R. LAINSON,* IRENE LANDAUt and J. J. SHA W* (Receivecl10 August 1973) SUMMARY FaUisia effusa gen.nov., sp.nov. (Haemosporidüdea: Garniidae) is de- scribed iI;Lthe teiid lizard, N eusticurus bicarinatus, from Pará State, north Brazil. Schizogony and gametogony both take place within thrombocytes and white cells of the peripheral blood. There is no development in the red blood cells, by which character the organism is difIerentiated from Garnia, the other known genus within the family. Another species,FaUisiamodesta gen.nov., sp.nov., is described in the iguanid lizard, Tropidurus torquatus hispidus, algo from Pará, Brazil. It is distinguished from F. effusa by its development principally in the lymphocytes, as opposed to the thrombo- . cytes. There are other, striking morphological differences between the two species. It is recommended that great care is needed in the interpretation of so-called 'exo-erythrocytic stages' of saurian plasmodia, which may well really belong to members of the Garniidae, in mixed infections. INTRODUCTION Recently (Lainson, Landau & Shaw, 1971) we erected a new sporozoan fami1y, Garniidae (Coccidiida: Haemosporidiidea), for the inclusion of some newly dis- covered, non-pigmented haemosporidia of South American lizards. These parasites undergo schizogony and dimorphic gametogony in the cells of the peripheral blood of the vertebrate host but, when development is in the redblood cells, there is no production of malarial pigmento By these characters the parasites are differ- entiated from those of the existing three families of the suborder Haemospori- diidea: from the Plasmodüdae and the Haemoproteidae by the absence of pigment, and from the Leucocytozoidae and the Haemoproteidae by the asexual cycle in the circulating blood. The type genus Garnia is characterized by its erythrocytic schizogony and gametogony within immature and mature red blood cells of the peripheral blood, with no production of malarial pigmento In some species there is developed a variable number of azurophilic granules in the cytoplasm, but these are negative to polarized light. Exoerythrocytic schizogony occurs in lymphocytes, monocytes and throm bocytes. Known speciesinclude Garnia gonatodi (Telford, 197O) Lainson, * The WellcomeParasitology Unit, Caixa Postal 3, Belém Pará, Brazil. t MuseumNational d'Histoire Naturelle, Laboratoire de Zoologie(Vers), 57 Rue Cuvier, Paris Ve, France. 8-2 118 R. LAINSON, IRENE LANDAU AND J. J.. SHAW Landau & Shaw, 1971; G. morula (Telford, 1970) Lainson, Landau & Shaw, 1971; G. telfordi Lainson, Landau & Shaw, 1971; and G. utingensis Lainson, Landau & Shaw, 1971. When these parasites were first examined by the electron microscope, minute cytoplasmic inclusions were thought to be sub-light-microscopic grains of pigmento Subsequent opinion, however, is that this was not so, and that we can confidently reiterate our previous definition that the parasites of this family do not produce malarial pigment when developing in the red blood cells. The bulk of the present paper deals with the discovery of an extraordinary new haemosporidian of the teiid lizard Neusticurus bicarinatus (Linnaeus, 1758) Duméril & Bibron, 1839, from Ananindeua and Capanema, Pará State, north Brazil. Although the parasite is clearly related to Garnia, it undergoes both schizo- gony and gametogony in the thrombocytes and, more rarely, the lymphocytes. There is no development in the red blood cells, and we reeI that this peculiarity alone justifies separating the parasite from the genus Garnia within the family Garniidae. The name Fallisia gen.nov. is proposed, in honour of Professor Murray Fallis who has contributed much to our knowledge of other pigmentless haemosporidia, the Leucocytozoidae. A further species of this parasite is also described in the iguanid lizard Tropidurus torquatus hispidus (Spix, 1825) Burt & Burt, 1930, from Belém, Pará, Brazil. MATERIAL AND METHODS N. bicarinatus is a small, semi-aquaticlizard of the family Teiidae. It lives in humid forested aroos, particularly along the banks of the rivers and streamsin which it rapidly seeks protection against would-be predators. The lizards were kept in glass aguaria, with the option of wet or dry conditions, and a number of loose stones under which these animaIs like to hide. T. torquatushispidus, on the other hand, is a sun-Iovingreptile. It prefers open country, waste land, roadsidesor the larger forest clearings.This lizard keepswell in dry aguaria, and water is neededonly for drinking purposes. The lizards were alI bled from the orbital sinus with a glass pipette drawn to a very fine point. Blood illms were dried quickly, fixed at once in absolute methyl alcohol, and stained for 1-2 h in Giemsa stain (30 drops to 15 ml distilled water buffered to pH 7.4). DESORIPTION OF THE PARASITES Fallisia effusa gen.nov.,sp.nov. (PI. 1, figs. 1-15; PI. 2, figs. 16-19; PI. 3, figs. 29-40; PI. 4, figs. 41-46) In spite of a long study of blood films from many infected lizards and the examination of day-to-day developmentin individual animaIs, it has remained difficult to be sure of the complete evolution of the parasite in the peripheral blood. Thus, while it is simple to state that infection starts with a period of schizogonyand endswith an infection consistingalmost entirely of mature gameto- Haemosporidia01 lizards cytes, it is not so easy to interpret the abundance of bizarre forros of the organism in between these two stages. Luckily the infection is extremely common in N. bicarinatus, enabling a large amount of material to be examined. Twenty-six out of thirty-two lizards examined were positive. The youngest parasites seen within the thrombocytes (76 %) or lymphocytes (14 %) measured only ,...,2.0 x 1.0 ",m (PI. 1, figo 3) and are clearly derived from mature, rupturing schizonts (PI. 1, figo 8; PI. 3, figo 34). Some of these merozoites give rise to further schizonts and others apparently develop directly into male and female gametocytes. Schizogony The length of the initial schizogony cycle remains undetermined as, in the absence of experimental transmission, we have been unable to follow the infection from the very beginning. The young trophozoite had an irregular shape, often stellate, with tiny pseudo- podia. Nuclear division (PI. 1, figs. 4-8; PI. 3, figs. 29-34) gives rise to separate chromatin masses (PI. 3, figo 30) or, frequently, t? a network ofnuclear material studded with more intensely stained granules (PI. 1, figs. 5,6). With increasing nuclear division the parasite assumes a smoother outline, the final product being round or oval (PI. 1, figo 7): mature schizonts measure '" 10.0-14.0 p;m in diameter and produce from 50-150 merozoites. These tiny bodies are mostly tear-shaped to triangular (PI. 1, figo 8; PI. 3, figo 34) and measure '" 2.0 x 1.0 p;m: they enter new thrombocytes and lymphocytes to form further schizonts, or gametocytes. Gametogony Young gametocytesbegin to appear early in the infection, when schizogonyis still at a relatively low leveI and when the infection is light. Someof the invading merozoitesdo not undergo nuclear division, but enlarge: whilst still very small they are readily recognizable as young male and female gametocytes (PI. 1, figs. 9, 10). When there is only one parasitein the cell, these early stagesare oval to bean-shaped,but may be distorted by pressure from adjacent parasites in multiple infections. Some already show a variable number of tiny azurophilic granules, such as seenin speciesof Garnia. The microgametocytehas the usual diffusely scattered nuclear material, often with a distinct nucleolus, and has an overall pink hue which contrasts strongly with the vivid blue cytoplasm of the macrogametocyte.The nucleus of the female parasite forros a well-defined area, often band-form: again, there is usually a conspicuousnucleolus. The normal shape of the mature gametocytesappears to be an elongateoval (PI. 1, figs. 11, 12)but they are frequently deformedby pressure againstthe host-cellnucleus, or againstone anotherin cells with multiple infections (PI. 1, figo 14; PI. 3, figo 40). The mature female measures'" 10.0x 6.0 p,m: the males are smaller, '" 8.0 x 5,0 p,m. Both sexesmay have 1-3 prominent vacuoles (PI. 3, figs. 39, 40); more usually there is a single vacuole, measuring up to 1.0 p,m. Counts of gametocytes,in three different lizards which showed only mature 119 R. LAINSON, IRENE LANDAU AND J. J. SHAW forms, gave the proportions ofmales to females as 107: 136,119: 149 and 121: 181. A iourth lizard, however, initially showed only 2: 98; and 1 week later it appeared to have only female parasites in the peripheral blood (out of 230 gametocytes counted). We are at a 10ssto explain this strange disappearance of the micro- gametocytes. As far as we could determine, the development time of the gametocytes is 7 to 10 days. M ultiple infection of tM host cells We have previousIy stressed that muItipIe invasion of the ce11sis a prominent feature ofthe genus Garnia. It is even more pronounced in Fallisia effusa. Thus, a study of lizards with light and presumabIy recent infections showed non-infected and infected thrombocytes in the proportion of 25:29. DoubIe infections were very rare and the parasites were aImost aIways trophozoites and young schizonts, derived from a sma11number of mature schizonts of the tyPe described above. Two days Iater the number of mature schizonts had increased and the proportion of non-infectedjinfected ce11swas now 13: 41. Numerous ce11salready contained two or three parasites and, on subsequent days, the number of infected ce11ssteadiIy increased until virtua.lIy 100% ofthe thrombocytes were parasitized with variabIe mixtures oftrophozoites, schizonts and deveIoping gametocytes (PI. 1, figo 15; PI. 2, 16-19; PI.
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