Molecular characterization of haemococcidia genus Schellackia (Apicomplexa) reveals the polyphyletic origin of the family Lankesterellidae RODRIGO MEG  IA-PALMA JAVIER MART  INEZ &SANTIAGO MERINO Submitted: 15 January 2014 Accepted: 2 February 2014 doi:10.1111/zsc.12050 Meg  ıa-Palma, R., Mart  ınez, J. & Merino, S. (2014). Molecular characterization of haemococcidia genus Schellackia (Apicomplexa) reveals the polyphyletic origin of the family Lankesterellidae. — Zoologica Scripta, 00, 000–000. The current taxonomy on the haemococcidia establishes that the two genera of protozoan parasites that integrate the family Lankesterellidae are Lankesterella and Schellackia. How- ever, the phylogeny of these genera, as well as the other coccidia, remains unresolved. In this sense, the use of type and described species is essential for the resolution of systematic conflicts. In this study, we molecularly characterize the type species of the genus Schellackia, that is, S. bolivari from Europe and also a described species of the same genus from Asia. At the same time, we contribute with the molecular characterization of another species of the genus Lankesterella. All this put together supports the polyphyly of the family Lankesterelli- dae. Therefore, we propose the resurrection of the zoological family, Schellackiidae Grass e 1953, to include species within the genus Schellackia. Corresponding author: Rodrigo Meg  ıa-Palma, Departamento de Ecolog  ıa Evolutiva, Museo Nacional de Ciencias Naturales-CSIC, Jos  e Guti  errez Abascal 2, E-28006 Madrid, Spain. E-mail: rodrigo.megia@mncn.csic.es Rodrigo Meg  ıa-Palma, Departamento de Ecolog  ıa Evolutiva, Museo Nacional de Ciencias Naturales- CSIC, J. Guti  errez Abascal, 2, Madrid, E-28006, Spain. E-mail: rodrigo.megia@mncn.csic.es Javier Mart  ınez, Departamento de Microbiolog  ıa y Parasitolog  ıa, Facultad de Farmacia, Universidad de Alcal a de Henares, Alcal a de Henares, Madrid, E-28871, Spain. E-mail: francisco. martinez@uah.es Santiago Merino, Departamento de Ecolog  ıa Evolutiva, Museo Nacional de Ciencias Naturales-CSIC, J. Guti  errez Abascal, 2, Madrid, E-28006, Spain. E-mail: santiagom@mncn.csic.es Introduction In 1899, Labb e described the genus Lankesterella in a frog species. This is a genus of apicomplexan parasites that occur primarily in amphibians around the world (Upton 2000), although there are some species within the genus Lankesterella described in lizards from Europe (  Alvarez- Calvo 1975; Chiriac & Steopoe 1977), and recent molecu- lar studies have reported lankesterellids infecting birds (Merino et al. 2006; Biedrzycka et al. 2013). This genus is characterized by endogenous oocysts containing 32, or more, naked sporozoites. Later on, in 1920, Reichenow described the genus Schellackia in the blood cells of Acanthodactylus vulgaris (=erythrurus) and Psammodromus hispanicus, both of the family Lacertidae, in a population from Madrid, Spain. After carrying out some cross-infec- tion experiments among individuals of both species of lizards, he concluded the conspecificity of the parasite (Reichenow 1920). The main characteristic of the genus is the formation of thin-walled oocysts in the lamina propia each containing eight naked sporozoites (Upton 2000; Telford 2008). In 1920, N€ oller coined the name of the family Lankesterellidae that include both genera, Lankesterella and Schellackia. All species of this family are heteroxenous, but sexual and asexual reproduction (i.e. merogony, gamogony and sporogony) occur in the vertebrate host’s gut. The oocysts are not expelled outside, and the sporozoites are released in situ and pass through gut to the blood stream where they penetrate into blood cells. Thereafter, the spor- ozoites are ingested by haematophagous invertebrate hosts (i.e. mites, dipterans or leeches) where they became dormant stages (Upton 2000). In 1926, Wenyon described the subfamilies Schellackinae and Lankesterellinae within the family Lankesterellidae. Some years after, Grass e (1953) reclassify these two ª 2014 Royal Swedish Academy of Sciences 1 Zoologica Scripta