The wide geographic distribution of Schistosoma mansoni, a digenetic trematode and parasite of humans, is determined by the occurrence of its intermediate hosts, freshwater snails of the genus Biomphalaria (Preston 1910). We present phylogenetic analyses of 23 species of Biomphalaria, 16 Neotropical and seven African, including the most important schistosome hosts, using partial mitochondrial ribosomal 16S and complete nuclear ribosomal ITS1 and ITS2 nucleotide sequences. A dramatically better resolution was obtained by combining the data sets as opposed to analyzing each separately, indicating that there is additive congruent signal in each data set. Neotropical species are basal, and all African species are derived, suggesting an American origin for the genus. We confirm that a proto-Biomphalaria glabrata gave rise to all African species through a trans-Atlantic colonization of Africa. In addition, genetic distances among African species are smaller compared with those among Neotropical species, indicating a more recent origin. There are two species-rich clades, one African with B. glabrata as its base, and the other Neotropical. Within the African clade, a wide-ranging tropical savannah species, B. pfeifferi, and a Nilotic species complex, have both colonized Rift Valley lakes and produced endemic lacustrine forms. Within the Neotropical clade, two newly acquired natural hosts for S. mansoni (B. straminea and B. tenagophila) are not the closest relatives of each other, suggesting two separate acquisition events. Basal to these two species-rich clades are several Neotropical lineages with large genetic distances between them, indicating multiple lineages within the genus. Interesting patterns occur regarding schistosome susceptibility: (1) the most susceptible hosts belong to a single clade, comprising B. glabrata and the African species, (2) several susceptible Neotropical species are sister groups to apparently refractory species, and (3) some basal lineages are susceptible. These patterns suggest the existence of both inherent susceptibility and resistance, but also underscore the ability of S. mansoni to adapt to and acquire previously unsusceptible species as hosts. Biomphalaria schrammi appears to be distantly related to other Biomphalaria as well as to Helisoma, and may represent a separate or intermediate lineage.
The validity of Biomphalaria kuhniana (Clessin, 1883) is confirmed through morphological study of specimens from Surinam (type locality) and the area of Tucurui (Tocantins river, state of Pará, Brazil) in comparison with B. straminea (Dunker, 1848), and throught crossing experiments which revealed complete reproductive isolation between the two species. The full-grown shell of kuhniana is smaller (about 7.5 mm) than that of straminea (11 mm to 16.5 mm). Anatomically they differ in the degree of corrugation of the vaginal wall (little developed in kuhniana, conspicuous in straminea), number and shape of prostatic diverticula (kuhniana 4 to 9, shorter and less branched; straminea 9 to 18, longer and more branched),number of muscle layers at the middle of the penis (two in kuhniana, three in straminea), distal segment of the spermiduct usually straight or slightly wavy in kuhniana, more or less curly in straminea. Differences between B. kuhniana and B. intermedia (paraense & Deslandes, 1962) are less marked. The latter has a shell up to about 12 mm in diameter, 7 to 15 prostatic diverticula, two muscle layers at the middle of the penis, and a vaginal wall with a combination of a more or less developed corrugation (or sometimes a mere swelling) on the left of the spermathecal duct and a rudimentary pouch on the right of the duct. A Biomphalaria straminea complex is proposed to include that species as well as B. kuhniana and B. intermedia.
A new species of South American planorbid snail, Biomphalaria occidentalis, is described. It is indistinguishable from B. tenagophila (Orbigny, 1835), by the characteristics of the shell and of most organs of the genital system. In B. tenagophila there is a pouch on the ventral wall of the vagina (Fig. 4A, vp), absent in B. occidentalis (Fig. 3A), and on the ventral wall of the vagina (Fig. 4A, vp), absent in B. occidentalis (Fig. 3A), and the prepuce is much wider than the penial sheath, its width increasing distalward (Fig. 4, ps,pp), whereas in B. occidentalis the prepuce is wider than the penial sheath but keeps about the same width all along (Fig.3, ps, pp). The two species are biologically separate by absolute reproductive isolation. The geographical distribution of B. occidentalis is shown in Fig. 14. So far it has been found in the Brazilian states of Acre, Amazonas (?), Mato Grosso, Mato Grosso do Sul and Paraná, and in Paraguay. Its type-locality is Campo Grande, state of Mato Grosso do sul, where it was collected from several biotopes related to affluents of the Aquiduana river, chiefly Córrego Prosa and Córrego Ceroula. Specimens were deposited in the following malacological collections: Instituto Oswaldo Cruz, Rio de Janeiro; Academy of Natural Sciences, Philadelphia; Museum of Zoology, University of Michigan; and British Museum (Natural History).
É descrita uma nova espécie de molusco planorbídeo sul-americano, Biomphalaria occidentalis, indistinguível da b. tenagophila (Orbigny, 1835) pelos caracteres da concha e da maioria dos órgãos do sistema genital. Na B. tenagophila existe uma bolsa na parede ventral da vagina (Fig. 4A, vp), ausente na B. occidentalis (Fig. 3A), e o prepúcio é muito mais grosso que a bainha do pênis, aumentando de largura na direção distal (Fig.4, ps, pp), ao passo que na B. occidentalis o prepúcio é mais grosso que a bainha do pênis porém seu diâmetro pouco se altera em toda sua extensão (Fig.3, ps, pp). As duas espécies são biologicamente separadas por isolamento reprodutivo absoluto. A Fig. 14 mostra a distribuição geográfica da B. occidentalis, que até agora foi encontrada nos Estados brasileiros do Acre, Amazonas (?), Mato Grosso, Mato Grosso do Sul e Paraná, e no Paraguai. Sua localidade-tipo é Campo Grande, Estado de Mato Grosso do Sul, onde foi coletada em vários biótopos relacionados e afluentes do rio Aquidauana, principalemente nos córregos Prosa e Ceroula. Foram depositados exemplares nas seguintes coleç~eos malacológicas: Instituto Oswaldo Cruz, Rio de Janeiro; Academy of Natural Sciences, Philadelphia; Museum of Zoology, University of Michigan; British Museum (Natural History)
A description of the species Lymnaea diaphana King, 1830 is presented, on the basis of material collected at its type-locality, San Gregorio, on the north coast of the Strait of Magellan, in the Chilean province of Magallanes. It may be identified by the following characters taken together: adult shell over 10 mm in length, whorls inflated, regularly convex, separated by a well-marked suture, aperture ovate occupying about half the shell length; renal organ forming an approximately right angle with the ureter; pouch of the oviduct well noticeable high on the right ventral surface and on the right side of the nidamental gland; uterus bent to the right into an approximately right angle; body of the spermatheca projected into the pulmonary cavity and adhered to the pericardium and to the roof of the pulmonary cavity; spermiduct highly sinuous, folding dorsalward between the left half of the oviduct and the left shoulder of the nidamental gland, and then winding on ventralward to reach the prostate on the middle line; prostate voluminous, convex on the left, pushed in on the right, with a deep dorsal furrow corresponding to a fold which projects into the prostatic lumen and is more developed at the fore half of the organ; apical end of the penial sheath with about six minute protuberances corresponding to inner chambers; prepuce from about as long about twice as long as the penial sheath, with some variation beyond those limits; lateral teeth of the radula basically tricuspid, with a usually simple ectocone which may show a bifid or trifid point. A diagnosis between lymnaea diaphana and three other lymnaeids which also occur in South America and were previously studied by the author - L. columella, L. viatrix and L. rupestris - is presented.
The historical phylogeography of the two most important intermediate host species of the human blood fluke Schistosoma mansoni, B. glabrata in the New World, and B. pfeifferi in the Old World, was investigated using partial 16S and ND1 sequences from the mitochondrial genome. Nuclear sequences of an actin intron and internal transcribed spacer (ITS)-1 were also obtained, but they were uninformative for the relationships among populations. Phylogenetic analyses based on mtDNA revealed six well-differentiated clades within B. glabrata: the Greater Antilles, Venezuela and the Lesser Antilles, and four geographically overlapping Brazilian clades. Application of a Biomphalaria-specific mutation rate gives an estimate of the early Pleistocene for their divergence. The Brazilian clades were inferred to be the result of fragmentation, due possibly to climate oscillations, with subsequent range expansion producing the overlapping ranges. Within the Venezuela and Lesser Antilles clade, lineages from each of these areas were estimated to have separated approximately 740 000 years ago. Compared to B. glabrata, mitochondrial sequences of B. pfeifferi are about 4x lower in diversity, reflecting a much younger age for the species, with the most recent common ancestor of all haplotypes estimated to have existed 880 000 years ago. The oldest B. pfeifferi haplotypes occurred in southern Africa, suggesting it may have been a refugium during dry periods. A recent range expansion was inferred for eastern Africa less than 100 000 years ago. Several putative species and subspecies, B. arabica, B. gaudi, B. rhodesiensis and B. stanleyi, are shown to be undifferentiated from other B. pfeifferi populations.
A review of lymnaeid samples collected by the author from 106 localities in Mexico, Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico, Martinique, Saint Lucia, Guatemala, Costa Rica, Panamá, Ecuador, Peru, Bolivia, Chile, Argentina, Uruguay andBrazil showed that one of them (from Ecuador) belonged to Lymnaea cousini Jousseaume, 1887, and all the others to either L. viatrix Orbigny, 1835 or l. columella Say, 1817. The ranges of L. viatrix and L. columella overlap in Middle America, and in northern and southern South America (Venezuela-Colombia-Ecuador and northeastern Argentina-Uruguay-southernmost Brazil, respectively). L. viatrix was the only species found in Peru west of the Andes and in Chile, and is supposed to have migrated eastward to Argentina via the Negro river basin. The range of L. columella in South America is discontinuous. The species has been recorded from Venezuela, Colombia and Ecuador and, east of the Andes, from latitudes 15º S (central-west Brazil) to 35º S (La Plata, Argentina). Such a gap may be attributed to either introduction from the northern into the southern area, or migration along the unsampled region on the eastern side of the Andes, or extinction in the now vacant area. No lymnaeids have been found so far in Brazil north of latitude 15º S and in the Guianas.
A revisão de amostras de limneídeos coletados pelo autor em 106 localidades no México, Cuba, Jamaica, Haiti, República Dominicana, Porto Rico, Martinica, Santa Lúcia, Guatemala, Costa Rica, Panamá, Equador, Peru, Bolívia, Chile, Argentina, Uruguai e Brasil mostrou que uma delas (do Equador) pertencia a Lymnaea cousini Jousseaume, 1887, e que todas as outras pertenciam a L. viatrix Orbigny, 1835 ou a L. columella Say, 1817. As distribuições da L. viatrix e da l. columella sobrepõem-se na Meso-América e na América do Sul setentrional (Venezuela-Colômbia-Equador) e meridional (nordeste da Argentina-Uruguai-extremo sul do Brasil). A L. viatrix foi a única espécie encontrada no Peru, a oeste dos Andes, e no Chile,parecendo ter migrado na direção leste para a Argentina através da bacia do rio Negro. A distribuição de L. columella na América do Sul é descontínua. Tem sido encontrada na Venezuela, Colômbia e Equador e, a leste dos Andes, entre as latitudes 15º S (Centro-Oeste do Brasil) e 35º S (La Plata, Argentina). Esse hiato na distribuição pode ser atribuído à introdução da espécie do norte para o sul, ou à migração ao longo da faixa não estudada no lado leste dos Andes, ou à extinção na área atualmente vaga. Até agora não foi registrada a presença de limneídeos no Brasil ao norte da latitude 15º S e nas Guianas
Susceptibility experiments were carried out with a Biomphalaria straminea-like planorbid snail (Biomphalaria aff. straminea, species inquirenda) from Espinillar, near Salto (Uruguay), in the area of the Salto Grande reservoir, exposed individually to 5 miracidia of Schistosoma mansoni (SJ2 and BH2 strains). Of 130 snails exposed to the SJ2 strain, originally infective to Biomphalaria tenagophila, 30 became infected (23%). The prepatent (precercarial) period ranged from 35 to 65 days. The cercarial output was irregular, following no definite pattern, varying from 138 to 76,075 per snail (daily average 4.3 to 447.5) and ending up with death. Three specimens that died, without having shed cercariae, on days 69 (2) and 80 after exposure to miracidia, had developing secondary sporocysts in their tissues, justifying the prospect of a longer precercarial period in these cases. In a control group of 120 B. tenagophila, exposed to the SJ2 strain, 40 became infected, showing an infection rate (33.3%) not significantly different from that of the Espinillar snail (chi 2 = 3.26). No cercariae were produced by any of the Espinillar snails exposed to miracidia of the BH2 strain, originally infective to Biomphalaria glabrata. Four specimens showed each a primary sporocyst in one tentacle, which disappeared between 15 and 25 days post-exposure, and two others died with immature, very slender sporocysts in their tissues on days 36 and 54. In a control group of 100 B. glabrata exposed to BH2 miracidia, 94 shed cercariae (94%) and 6 remained negative. Calculation of Frandsen's (1979a, b) TCP/100 index shows that "Espinillar Biomphalaria-SJ2 S. mansoni" is a vector-parasite "compatible" combination.(ABSTRACT TRUNCATED AT 250 WORDS)
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