INTRODUCTION
An article1 on Wellcome Trust funded research at Instituto Evandro Chagas, in Pará State, Brazil, did not give details of the new organisms that were discovered. A total of 119 new species were named and, due to taxonomic revisions, five species named previously by other authors were transferred (comb. nov.) to newly created genera. These taxa are listed in table 1 together with their hosts and authors. The aim of the present article is to briefly mention how their discovery influenced future research and to show how they relate to present trends in the different areas.
Ralph Lainson was supported for the longest period and his contributions were dominant, being an author/co-author of 108 new species and five new combinations. Jeffrey Shaw authored/co-authored 60 new species and two new combinations. One hundred and one of these new species were discovered in Brazil and two names were given to Leishmania found in Panama. The descriptions of 13 published individually by Ralph Lainson (10) and Jeffrey Shaw (three) were of material that they had collected while respectively working in Belize and Central America.
SPECIES OF PROTOZOA
PROTOZOA: EUCOCCIDIIDA & MICROSPORIDIA
The taxonomic ranks (family, genus, and subgenus) that were created for some of the species are not listed in table 1 and are as follows: the haemosporidian genus Saurocytozoon Lainson & Shaw, 19692 ; the haemosporidian family Garnidae Lainson, Landau & Shaw, 19713 containing the genera Garnia Lainson, Landau & Shaw, 1971, Fallisia Lainson, Landau & Shaw, 19744, and Progarnia Lainson, 19955; two hemogregarine genera Cyrilia Lainson, 19816 and Hemolivia Petit, Landau, Baccam & Lainson, 19907; and the microsporidian genus Alloglugea Paperna & Lainson, 19958. A simple question is why were they found? All are from cold-blooded vertebrates and the majority is from lizards. While searching for lizard Leishmania that to this date have never been found in the Americas, a wealth of parasites was discovered in different blood cells, opening new research areas. However, besides this, it was obvious that an enormous number of parasites was waiting to be discovered. Thus, any available animal was examined for parasites, irrespective of it being a potential reservoir of a human disease.
Their discoveries stimulated others to look for them around the world. Garnia is only known in the Western Hemisphere; Fallisia species have been recorded in the Americas, South East Asia, and Australasia; and Saurocytozoon occurs in both North and South America and in Asia9. Molecular methods are now showing possible evolutionary pathways that reflect the above distributions. The 60 Eimeridae species listed in table 1 were found in either warm or cold blooded vertebrates. A recent paper10 showed that species of Cyclospora, Eimeria, and Isospora from warm blooded vertebrates form a well-supported clade, but organisms identified as Eimeria or Isospora species from cold-blooded vertebrates and marsupials do not fall within this group, suggesting that these are polyphyletic genera. For example, based on 18S rRNA sequences10, two Eimeria species, one from a snake and the other from an anuran, were phylogenetically closer to European Schellackia species within the family Schellackiidae. These observations clearly indicate that future revisions will be required to resolve the polyphyletic genera.
Six species of Sarcocystis were described based on distinctive cyst wall morphology. Except for one, their definitive hosts are unknown. The sexual cycle of Sarcocystis ameivamastigodryasi Lainson & Paperna, 200011 of the teiid lizard, Ameiva ameiva occurs in the intestine of the colubrid snake, Mastigodryas bifossatus. The oocysts are like those of Isospora species having two sporocysts that contain four sporozoites. In the case of the teiid parasite, the oocyst wall ruptures inside the snake's intestine so mature sporocysts are liberated in the faeces. The cystic stage normally occurs in herbivores and the sexual stage in carnivores ranging from canids, corvid birds to snakes. It is possible that some Isospora species could in fact be Sarcocystis species.
PROTOZOA: KINETOPLASTIDA
Given the medical importance of Leishmania species, it is not surprising that the most well-known taxa and taxonomic rank, established in Belém, Pará State, under Wellcome Trust funding, belong to this genus. The creation of the subgenus Leishmania (Viannia) Lainson & Shaw, 198712 was a fundamental step forward in understanding the disease known as leishmaniasis and its phylogeny. Parasites belonging to this subgenus are only found in the Americas, while those of the subgenus Leishmania (Leishmania) are found in the Old World and the Americas. Three previously named organisms, L. braziliensis, L. guyanensis, and L. peruviana were assigned to this subgenus and eight other species were described (Table 1). All but two of these belong to the subgenus L. (Viannia). One of these, L. (L.) amazonensis Lainson & Shaw, 197213, grows profusely in culture and readily infects a range of laboratory animals, producing in hamsters a pathology like diffuse cutaneous leishmaniasis. Because of these attributes this parasite is used extensively in immunological and chemotherapeutic studies. Molecular methods14 have shown that the subgenus L. (Viannia) separated from the basal stock some 80 MYA while the subgenus L. (Leishmania) separation was later being around 50 MYA. There are distinct differences between the immunological response and pathology of the two subgenera in man. Somewhere along the evolutionary line, the subgenus L. (Leishmania) seems to have opted to depress the cell mediated response, while the subgenus L. (Viannia) opted to stimulate it. Perhaps this was due to the very different mammalian hosts in which the two subgenera evolved, one possible being rodents and the other Xenarthra. Today we see this reflected in the extreme immunological differences between diffuse cutaneous leishmaniasis caused by L. (L.) amazonensis and mucocutaneous by L. (V.) braziliensis15.
Leishmania hertigi deanei Lainson & Shaw 197716, later raised to specific status by Lainson and Shaw12 in 1987, is very different from all other Leishmania. The amastigotes are very large and do not appear to be intracellular. Latterly studies have shown that it is phylogenetically outside the genus Leishmania being closer to Endotrypanum. This led to the creation of the genus Porcisia Shaw, Camargo & Teixeira 201617 to accommodate it and the Panamanian porcupine parasite, P. hertigi.
Great advances in our understanding of the taxonomy of Trypanosoma (Schizotrypanum) cruzi were made by Miles et al.18 while working at the Instituto Evandro Chagas that confirmed three genetically distinct lineages that were denoted as Type I, II & III. In the opinion of the author these represent distinct species but have never been named so they do not appear in table 1. Subsequently these lineages have been shown to be distinct by with different molecular markers.
Protozoa | Host |
---|---|
Eucoccidiida: Haemosporida: Plasmodiidae | |
Plasmodium vacuolatum Lainson, Shaw & Landau, 1975 | Lizard |
Plasmodium neusticuri Lainson & Paperna, 1996 | Lizard |
Plasmodium kentropyxi Lainson, Landau & Paperna, 2001 | Lizard |
Plasmodium carmelinoi Lainson, Franco & Matta, 2010 | Lizard |
Eucoccidiida: Haemosporida: Leucocytozoidae | |
Saurocytozoan tupinambi Lainson & Shaw, 1969 | Lizard |
Saurocytozoan mabuyi Lainson, Landau & Shaw, 1974 | Lizard |
Eucoccidiida: Haemosporida: Haemoproteidae | |
Polychromophilus deanei Garnham, Lainson & Shaw, 1971 | Bat |
Haemoproteus peltocephali Lainson & Naiff, 1998 | Lizard |
Haemoproteus geochelonis Lainson & Naiff, 1998 | Lizard |
Eucoccidiida: Haemosporida: Garniidae | |
Garnia gonatodi (Telford, 1970) Lainson, Landau & Shaw, 197l, comb. nov. | Lizard |
Garnia telfordi Lainson, Landau & Shaw, 1971 | Lizard |
Garnia utingensis Lainson, Landau & Shaw, 1971 | Lizard |
Garnia multiformis Lainson, Shaw & Landau, 1975 | Lizard |
Garnia uranoscodoni Lainson, Shaw & Landau, 1975 | Lizard |
Garnia morula (Telford, 1970) Lainson, Landau & Shaw, 1971, comb. nov. | Lizard |
Garnia karyolytica Lainson & Naiff, 1999 | Lizard |
Fallisia effusa Lainson, Landau & Shaw, 1974 | Lizard |
Fallisia modesta Lainson, Landau & Shaw, 1974 | Lizard |
Fallisia audaciosa Lainson, Shaw & Landau, 1975 | Lizard |
Fallisia simplex Lainson, Shaw & Landau, 1975 | Lizard |
Progarnia archosauriae Lainson, 1995 | Lizard |
Eucoccidiida: Adeleina: Haemogregarinidae | |
Cyrilia lignieresi (Laveran, 1906) Lainson, l992, comb. nov. (Syns Haemogregarina lignieresi Laveran, 1906; H. gomesi Neiva & Pinto, 1926; Cyrilia gomesi Lainson, 1981) | Fish |
Hemolivia stellata Petit, Landau, Baccam & Lainson, 1990 | Anuran |
Eucoccidiida: Eimeriina: Lankesterellidae | |
Lankesterella petiti Lainson & Paperna, 1995 | Anuran |
Eucoccidiida: Eimeriina: Schellackiidae | |
Schellackia landauae Lainson, Shaw & Ward, 1976 | Lizard |
Eucoccidiida: Eimeriina: Eimeridae | |
Tyzzeria boae Lainson & Paperna, 1994 | Snake |
Cyclospora niniae Lainson, 1965 | Snake |
Cyclospora schneideri Lainson, 2005 | Snake |
Caryospora pseustesi Lainson, Nascimento & Shaw, 1991 | Snake |
Caryospora micruri Lainson, Nascimento & Shaw, 1991 | Snake |
Caryospora constanciae Lainson, Nascimento & Shaw, 1991 | Snake |
Caryospora paraensis Lainson, Nascimento & Shaw, 1991 | Snake |
Caryospora carajasensis Lainson, Nascimento & Shaw, 1991 | Snake |
Caryospora epicratesi Lainson, Nascimento & Shaw, 1991 | Snake |
Isospora albicolis Lainson & Shaw, 1989 | Bird |
Isospora wilkiei Lainson, 1968 | Crocodile |
Isospora basilisci Lainson, 1968 | Lizard |
Isospora tucuruiensis Lainson & Shaw, 1989 | Bird |
Protozoa | Host |
Isospora saimiri Lainson & Shaw, 1989 | Mammal |
Isospora cacici Lainson, 1994 | Bird |
Isospora thraupis Lainson, 1994 | Bird |
Isospora capanemaensis Lainson, 2003 | Mammal |
Isospora rodriguesae Lainson, Da Silva, Franco & De Souza, 2008 | Chelonia |
Eimeria orthogeomyos Lainson, 1968 | Mammal |
Eimeria tamanduae Lainson, 1968 | Mammal |
Eimeria rhynchonycteridis Lainson, 1968 | Mammal |
Eimeria pseudemydis Lainson, 1968 | Turtle |
Eimeria bothrops Lainson, 1968 | Snake |
Eimeria ameivae Lainson, 1968 | Lizard |
Eimeria crocodyli Lainson, 1968 | Crocodile |
Eimeria poti Lainson, 1968 | Mammal |
Eimeria micruri Lainson & Shaw, 1973 | Snake |
Eimeria liophi Lainson & Shaw, 1973 | Snake |
Eimeria leimadophi Lainson & Shaw, 1973 | Snake |
Eimeria cyclopei Lainson & Shaw, 1982 | Mammal |
Eimeria choloepi Lainson & Shaw, 1982 | Mammal |
Eimeria trichechi Lainson, Naiff, Best & Shaw, 1983 | Mammal |
Eimeria philanderi Lainson & Shaw, 1989 | Mammal |
Eimeria caluromydis Lainson & Shaw, 1989 | Mammal |
Eimeria vitellini Lainson, Costa & Shaw, 1990 | Bird |
Eimeria corticulata Lainson & Shaw, 1990 | Mammal |
Eimeria zygodontomyis Lainson & Shaw, 1990 | Mammal |
Eimeria lagunculata Lainson, Costa & Shaw, 1990 | Chelonia |
Eimeria mammiformis Lainson, Costa & Shaw, 1990 | Chelonia |
Eimeria podocnemis Lainson, Costa & Shaw, 1990 | Chelonia |
Eimeria carinii Lainson, Costa & Shaw, 1990 | Chelonia |
Eimeria marajoensis Lainson & Shaw, 1991 | Mammal |
Eimeria porphyrulae Lainson, 1994 | Bird |
Eimeria crypturelli Lainson, 1994 | Bird |
Eimeria bufomarini Paperna & Lainson, 1995 | Anuran |
Eimeria peltocephali Lainson & Naiff, 1998 | Turtle |
Eimeria molossi Lainson & Naiff, 1998 | Bat |
Eimeria bragancaensis Lainson & Naiff, 2000 | Bat |
Eimeria carmelinoi Lainson, 2002 | Lizard |
Eimeria damnosa Lainson, Brigido & Silveira, 2005 | Mammal |
Eimeria lepidosirenis Lainson & Ribeiro, 2006 | Fish |
Eimeria amazonensis Lainson, Da Silva, Franco & De Souza, 2008 | Chelonia |
Eimeria carbonaria Lainson, Da Silva, Franco, & De Souza, 2008 | Chelonia |
Eimeria carajasensis Lainson, Da Silva, Franco & De Souza, 2008 | Chelonia |
Eimeria wellcomei Lainson, Da Silva, Franco & De Souza, 2008 | Chelonia |
Acroeimeria paraensis Lainson, 2002 | Lizard |
Acroeimeria cnemidophori (Carini, 1941) Lainson, 2002, comb. nov. | Mammal |
Choleoeimeria rochalima (Carini & Pinto, 1926) Lainson & Paperna, 1999, comb. nov. | Lizard |
Choleoeimeria carinii Lainson & Paperna, 1999 | Lizard |
Choleoeimeria amphisbaenae Lainson, 2003 | Lizard |
Eucoccidiida: Eimeriina: Sarcocystidae | |
Sarcocystis kinosterni Lainson & Shaw, 1972 | Mammal |
Sarcocystis azevedoi Shaw & Lainson, 1969 | Mammal |
Sarcocystis marmosae Shaw & Lainson, 1969 | Mammal |
Sarcocystis oryzomyos Shaw & Lainson, 1969 | Mammal |
Sarcocystis proechimyos Shaw & Lainson 1969 | Mammal |
Sarcocystis ameivamastigodryasi Lainson & Paperna, 2000 | Lizard/Snake |
Protozoa | Host |
Piroplasmida: Theileriidae | |
Theileria electrophori Lainson, 2007 | Fish |
Microspora: Glugeidae | |
Alloglugea bufonis Paperna & Lainson, 1995 | Anuran |
Kinetoplastida: Trypanosomatidae | |
Endotrypanum monterogeii Shaw, 1969 | Mammal |
Leishmania (Leishmania) amazonensis Lainson & Shaw, 1972 | Mammal* |
Leishmania (Viannia) panamensis Lainson & Shaw, 1972 | Mammal* |
Leishmania (Leishmania) aristidesi Lainson & Shaw, 1979 | Mammal |
Leishmania (Viannia) lainsoni Silveira, Shaw, Braga & Ishikawa, 1987 | Mammal* |
Leishmania (Viannia) naiffi Lainson & Shaw, 1989 | Mammal* |
Leishmania (Viannia) shawi Lainson, Braga, de Souza, Póvoa & Ishikawa, 1989 | Mammal* |
Leishmania (Viannia) lindenbergi Silveira, Ishikawa, de Souza & Lainson, 2002 | Mammal* |
Leishmania (Viannia) utingensis Braga, Lainson, Ishikawa & Shaw, 2003 | Mammal |
Porcisia deanei (Lainson & Shaw, 1977) Espinosa et al., 2018 | Mammal |
Trypanosoma leuwenhoeki Shaw, 1969 | Mammal |
Trypanosoma preguici Shaw, 1969 | Mammal |
Trypanosoma plicae Lainson, Shaw & Landau, 1975 | Lizard |
Trypanosoma cecili Lainson, 1977 | Crocodile |
Trypanosoma (Megatrypanum) saloboense Lainson, Da Silva & Franco, 2008 | Mammal |
Insecta | |
Diptera: Psychodidae: Phlebotominae | |
Bruptomyia orlandoi Fraiha, Shaw & Lainson, 1970 | |
Psychodopygus wellcomei Fraiha, Shaw & Lainson, 1971† | |
Psychodopygus lainsoni Fraiha & Ward, 1974 | |
Nyssomyia umbratilis (Ward & Fraiha, 1977) Galati, 2003‡ | |
Psychodopygus llanosmartinsi Fraiha & Ward, 1980† | |
Nyssomyia shawi (Fraiha, Ward & Ready, 1981) Galati, 2003† | |
Nyssomyia richardwardi (Ready & Fraiha, 1981) Galati, 2003 | |
Trichopygomyia ratcliffei (Arias, Ready & Freitas, 1983) Galati, 2003 | |
Psychodopygus leonidasdeanei Fraiha, Ryan, Ward, Lainson & Shaw, 1986 | |
Evandromyia carmelinoi (Ryan, Fraiha, Lainson & Shaw, 1986) Galati, 2003 | |
Trichopygomyia readyi (Ryan, 1986) Galati, 2003 |
* Infections found in man; † Found infected with Leishmania (Viannia) braziliensis; ‡ Found infected with Leishmania (Viannia) guyanensis.
SPECIES OF INSECTS
INSECTA: DIPTERA: PSYCHODIDAE: PHLEBOTOMINAE
Unravelling the epidemiologies of the different Leishmania species inevitable led to the discovery of new phlebotomine species. It also showed how the epidemiological importance of different groups varies in different biomes. The discovery of Psychodopygus wellcomei Fraiha, Shaw & Lainson 197119 was the first indication of the vectorial importance of this genus for L. (Viannia) species in Amazonia. Six Psychodopygus species are associated with L. (V.) braziliensis and five with L. (V.) naiffi20. It is the predominant genus in south of the Amazon River extending to virgin Atlantic rain forest. In north of the river, L. (V.) guyanensis, transmitted by N. umbratilis, is the dominant leishmania in man. The number of Nyssomyia sand flies is significantly greater in this biome, but there is no significant difference in the variety of species of Nyssomyia and Psychodopygus21.
Continuous environmental variations related to global warming and man's activities modulate the sand fly fauna. Understanding and documenting this is the challenge.
CONCLUSION
Unforeseen benefits have resulted from the description of the species that form the subject of this paper. They range from a clearer understanding of the taxonomic groups to which they belong to how some are transmitted to man with resultant contrasting pathologies and treatments. For example, in 1965 it was accepted that Leishmania braziliensis was the etiological agent of all forms of cutaneous leishmaniasis in Brazil. Studies of the parasites from wild animals, man and sand flies showed that this was wrong. Many scientific doors were opened, giving just a glimpse to the amazing variety of protozoal parasites that occur in Amazonian vertebrates.