Fig 1.
Pipeline of the analysis conducted.
Created in BioRender. Mora, J. (2026) https://BioRender.com/is02atg.
Fig 2.
Bayesian inference phylogenetic trees of Leishmania spp. using the RNA polymerase subunit II (a), 18S (b) and kDNA (c).
Posterior probability (PP) values lower than 0.6 are not shown. Node circle size and color are proportional to the PP. Created in BioRender. Mora, J. (2026) https://BioRender.com/is02atg.
Fig 3.
Global-fit and event-based cophylogenetic analysis between Leishmania spp. and its annotated vertebrate hosts including L. gymnodactyli association.
A. Tanglegram showing host-parasite associations according to GenBank metadata. B. Procrustean superimposition plot between the principal coordinates derived from patristic distances of the RNA Polymerase II of Leishmania spp. and their vertebrate host phylogenies. Each parasite and host are denoted as circles and arrow heads, respectively. Leishmania spp. are color coded according to the subgenus. Close host and parasite positions in the PCo may indicate cophylogenetic associations C. Contribution of each Leishmania-vertebrate host link to the global phylogenetic congruence. Each bar represents the squared residual of each association and are color-coded according to the Leishmania subgenus. Error bars correspond to 95% confidence intervals of the squared residuals. The median squared residual is indicated as a dotted line. Asterisks at the top of each bar represent a significant ParaFitLink1 value and daggers to significant ParaFitLink2 values. Squared residual values lower than the median squared value suggests cophylogenetic congruence between that host-parasite association. D. Coevolutionary reconstruction of the host (black lines) and parasite (blue lines) phylogenies with the lowest global cost according to eMPRess. Values in top of nodes correspond to the support of the predicted event. E. Total cost distribution of random solutions. Created in BioRender. Mora, J. (2026) https://BioRender.com/is02atg.
Table 1.
Summary of statistics of the global fit and event-based methods for the analysis of coevolution between Leishmania spp. and its vertebrate and invertebrate hosts.
Fig 4.
Global-fit and event-based cophylogenetic analysis between Leishmania spp. and its invertebrate hosts.
A. Tanglegram showing possible host-parasite associations according to the literature. B. Procrustean superimposition plot between the principal coordinates derived from patristic distances of the RNA Polymerase II of Leishmania spp. and their invertebrate host phylogenies. Each parasite and host are denoted as circles and arrow heads, respectively. Leishmania spp. are color coded according to the subgenus. Close host and parasite positions in the PCo may indicate cophylogenetic associations C. Contribution of each Leishmania-invertebrate host link to the global phylogenetic congruence. Each bar represents the squared residual of each association and are color-coded according to the Leishmania subgenus. Error bars correspond to 95% confidence intervals of the squared residuals. The median squared residual is indicated as a dotted line. Asterisks at the top of each bar represent a significant ParaFitLink1 value and daggers to significant ParaFitLink2 values. Squared residual values lower than the median squared value suggests cophylogenetic congruence between that host-parasite association. D. Coevolutionary reconstruction of the host (black lines) and parasite (blue lines) phylogenies with the lowest global cost obtained with Jane taking into consideration different Leishmania spp. isolates when having multi-host parasites. Values on top of nodes correspond to the support of the predicted event E. Total cost distribution of random solutions obtained with Jane. Red dotted line indicated the cost of the analyzed solution. Animal icons were created with Biorender.com.
Fig 5.
Supercontinent or Multiple origin hypothesis on the origin of the genus Leishmania according to the results obtained in this research, and previous observations made by Steverding (53), Harkins, Schwartz (21), Akhoundi, Kuhls (12) and Momen and Cupolillo (24).
A. During the Mesozoic era in the Jurassic period, before the breakup of the supercontinent Gondwana, Leishmania parasites may have used ancient rodent species to co-diversify, while ancient sand fly species may have emerged earlier. B. After the split of Gondwana in the Cretaceous period, different Leishmania subgenera arose, i.e., Viannia in South America, Leishmania and Sauroleishmania in Africa and Mundinia spread all over the world. With time, speciation in vertebrate hosts occurred by host-switching or duplication, where ancestral parasite species adapted to permissive hosts giving rise to new Leishmania spp. and parasites cospeciated with ancestors of the New and Old World sand flies. C. Dispersal of Leishmania spp. by using different vertebrate and invertebrate host gave rise to the colonization of new geographical regions. For instance, it has been suggested that rodents infected with L. major migrated through the Nearctic by the Bering Land Bridge to North America, originating L. mexicana in autochtonous rodent species, which further migrated to the South giving rise to L. amazonensis, L. venezuelensis or L. waltoni. Moreover, L. aethiopica was established in East Africa with hyraxes, as well as L. (Mundinia) procaviensis, and then switched to humans. Species of the Viannia subgenus spread throughout South and Central America using humans as a main reservoir, but according to other studies, sloths and porcupines may have played an important role in the maintenance of ancestral Viannia spp. In addition, species of the Sauroleishmania subgenus spread with herpetofauna to North Africa, East Asia and Europe, without having been reported in the Americas. Finally, species of the Mundinia subgenus have been described in diverse and widely separated geographical regions, suggesting that their distribution may date back to before the breakup of Gondwana. Created in BioRender. Mora, J. (2026) https://BioRender.com/jzxajbr.