This study presents new findings concerning the evolution of the human pathogens, Trypanosoma brucei and T. cruzi, which suggest that these parasites have divergent origins and fundamentally different patterns of evolution. Phylogenetic analysis of 18S rRNA sequences places T. brucei in a clade comprising exclusively mammalian trypanosomes of African origin, suggesting an evolutionary history confined to Africa. T. cruzi (from humans and sylvatic mammals) clusters with trypanosomes specific to Old and New World bats, T. rangeli and a trypanosome species isolated from an Australian kangaroo. The origins of parasites within this clade, other than some of those from bats, lie in South America and Australia suggesting an ancient southern super-continent origin for T. cruzi, possibly in marsupials; the only trypanosomes from this clade to have spread to the Old World are those infecting bats, doubtless by virtue of the mobility of their hosts. Viewed in the context of palaeogeographical evidence, the results date the divergence of T. brucei and T. cruzi to the mid-Cretaceous, around 100 million years before present, following the separation of Africa, South America and Euramerica. The inclusion in this study of a broad range of trypanosome species from various different hosts has allowed long phylogenetic branches to be resolved, overcoming the limitations of many previous studies. Moreover, T. brucei and the other mammalian tsetse-transmitted trypanosomes appear, from these data, to be evolving several times faster than T. cruzi and its relatives.

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The ancient and divergent origins of the human pathogenic trypanosomes, Trypanosoma brucei and T. cruzi.

The presence of haemoparasites from the Order Piroplasmida and the genera Bartonella and Trypanosoma was assessed in the blood of 60 bats, belonging to 7 species, inhabiting sites across Cornwall in southwest England. DNA extracted from macerated heart tissue was incorporated into taxon-specific polymerase chain reactions (PCRs) and amplification products were sequenced as a means of identifying, or assigning an identity, to detected haemoparasites. A Piroplasmida species was detected in 6 Pipistrellus spp., whereas Bartonella infections were detected in 5 bats belonging to 4 different species. Trypanosoma dionisii was detected in 1 Pipistrellus spp. Phylogenetic inference from alignment of a partial 18S rRNA-encoding gene sequence of the pipistrelle-associated Piroplasmida species with homologous sequences available for other members of the Order indicated that this organism was unique but specifically related to members of the genus Babesia, a phylogeny that would be in keeping with the organism being Babesia vesperuginis. Alignment of partial citrate synthase gene sequences from the bat-associated bartonellae revealed 5 distinct genotypes that were probably derived from 2 distinct Bartonella species. The study demonstrates the utility of molecular methods for detecting haemoparasites in dead bats and provides, for the first time, tangible identities for bat-associated Babesia and Bartonella species.

Molecular characterization of haemoparasites infecting bats (Microchiroptera) in Cornwall, UK.

Based on phylogenetic analysis of 18S rRNA sequences and clade taxon composition, this paper adopts a biogeographical approach to understanding the evolutionary relationships of the human and primate infective trypanosomes, Trypanosoma cruzi, T. brucei, T. rangeli and T. cyclops. Results indicate that these parasites have divergent origins and fundamentally different patterns of evolution. T. cruzi is placed in a clade with T. rangeli and trypanosomes specific to bats and a kangaroo. The predominantly South American and Australian origins of parasites within this clade suggest an ancient southern super-continent origin for ancestral T. cruzi, possibly in marsupials. T. brucei clusters exclusively with mammalian, salivarian trypanosomes of African origin, suggesting an evolutionary history confined to Africa, while T. cyclops, from an Asian primate appears to have evolved separately and is placed in a clade with T. (Megatrypanum) species. Relating clade taxon composition to palaeogeographic evidence, the divergence of T. brucei and T. cruzi can be dated to the mid-Cretaceous, around 100 million years before present, following the separation of Africa, South America and Euramerica. Such an estimate of divergence time is considerably more recent than those of most previous studies based on molecular clock methods. Perhaps significantly, Salivarian trypanosomes appear, from these data, to be evolving several times faster than Schizotrypanum species, a factor which may have contributed to previous anomalous estimates of divergence times.

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The Evolution of Trypanosomes Infecting Humans and Primates

A trypanosome identified as Trypanosoma (Schizotrypanum) dionisii Bettencourt & Franca, 1905, has been isolated from Pipistrellus pipistrellus (Chiroptera) in England. At least five out of eight P. pipistrelhis were infected.
 2. In the blood of P. pipistrelhis, the parasite closely resembled T. (S.) cruzi. When grown in vitro in monophasic or diphasic media at 28°C, epimastigotes and trypomastigotes developed. The latter were of two types — very long, thin forms and less numerous shorter individuals.
 3. The trypanosomes multiplied as amastigotes within HeLa and mouse L cells in vitro. After 6–9 days in HeLa cells at 37°C, they transformed into small trypomastigotes and emerged from the cells. Higher infection rates (up to about 4%) were obtained in cell cultures inoculated with flagellates from older monophasic cultures, which contained more of the long, slender trypomastigotes.
 4. Trypanosomes also entered (or were phagocytosed by) up to 60% or more of mouse peritoneal macrophages in vitro. Multiplication occurred by binary and multiple fission of amastigotes in at least some of the parasitized macrophages and transformation into trypomastigotes was seen after 7 or more days at 37°C. Development in macrophages was less synchronous than in HeLa cells.

Trypanosoma (Schizotrypanum) dionisii of pipistrellus pipistrellus (chiroptera): intra- and extracellular development in vitro.

Bats are unique flying mammalians that occur worldwide feeding either on insects, fruits or even on blood depending on the species. Since they are active during the night, many myths have been launched during the last 500 years. All these facets are shortly considered here and prepare for a hopefully intense look into their hidden life.

Gordon B. Thompson

Papers

Two species of Trypanosoma from British bats.