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Background Canine babesiosis is a severe disease caused by several Babesia spp. A number of names have been proposed for the canine-infecting piroplasmid pathogen initially named Theileria annae Zahler, Rinder, Schein & Gothe, 2000. It was shown to be a member of the Babesia ( sensu lato ) group infecting carnivores and is also closely related to the Babesia microti group. Subsequently, the same parasite species was reclassified as a member of the genus Babesia and the name Babesia vulpes Baneth, Florin-Christensen, Cardoso & Schnittger, 2015 was proposed for it. However, both names do not meet the requirements of the International Code of Zoological Nomenclature (no accompanying descriptions, no deposition of type-specimens) and cannot be recognized as available names from the nomenclatural point of view. The purpose of this study was to further characterize this parasite in order to confirm its validity, to provide its description and to introduce zoological nomenclature for it with the name Babesia vulpes n. sp. Results Morphological description of the parasite in canine erythrocytes demonstrated that it takes the shape of small (1.33 × 0.98 µm), round to oval forms reminiscent of the pyriform and ring shapes of other small canine Babesia spp., such as Babesia gibsoni Patton, 1910 and Babesia conradae Kjemtrup, Wainwright, Miller, Penzhorn & Carreno, 2006. However, these parasite forms were overall smaller than those measured for the latter two species and no tetrad (Maltese cross) form was reported. Furthermore, phylogenetic analysis using the cytochrome c oxidase subunit 1 (COX1) amino acid sequences substantiates the species identity of this parasite as previously demonstrated based on phylogenetic analysis of the 18S rRNA and β-tubulin genes. The holotype of the parasite species was designated and deposited in an accessible public collection. Conclusions This study ratifies the name Babesia vulpes n. sp. proposed for the parasite previously referred to as Theileria annae Zahler, Rinder, Schein & Gothe, 2000, Babesia annae (Zahler, Rinder, Schein & Gothe, 2000) or Babesia vulpes Baneth, Florin-Christensen, Cardoso & Schnittger, 2015, or mentioned as “ Babesia microti -like piroplasm”, “ Babesia Spanish dog isolate” and Babesia cf. microti .

BACKGROUND: Theileria annae is a tick-transmitted small piroplasmid that infects dogs and foxes in North America and Europe. Due to disagreement on its placement in the Theileria or Babesia genera, several synonyms have been used for this parasite, including Babesia Spanish dog isolate, Babesia microti-like, Babesia (Theileria) annae, and Babesia cf. microti. Infections by this parasite cause anemia, thrombocytopenia, and azotemia in dogs but are mostly subclinical in red foxes (Vulpes vulpes). Furthermore, high infection rates have been detected among red fox populations in distant regions strongly suggesting that these canines act as the parasite’s natural host. This study aims to reassess and harmonize the phylogenetic placement and binomen of T. annae within the order Piroplasmida. METHODS: Four molecular phylogenetic trees were constructed using a maximum likelihood algorithm based on DNA alignments of: (i) near-complete 18S rRNA gene sequences (n = 76 and n = 93), (ii) near-complete and incomplete 18S rRNA gene sequences (n = 92), and (iii) tubulin-beta gene sequences (n = 32) from B. microti and B. microti-related parasites including those detected in dogs and foxes. RESULTS: All phylogenetic trees demonstrate that T. annae and its synonyms are not Theileria parasites but are most closely related with B. microti. The phylogenetic tree based on the 18S rRNA gene forms two separate branches with high bootstrap value, of which one branch corresponds to Babesia species infecting rodents, humans, and macaques, while the other corresponds to species exclusively infecting carnivores. Within the carnivore group, T. annae and its synonyms from distant regions segregate into a single clade with a highly significant bootstrap value corroborating their separate species identity. CONCLUSION: Phylogenetic analysis clearly shows that T. annae and its synonyms do not pertain to Theileria and can be clearly defined as a separate species. Based on the facts that T. annae and its synonyms have not been shown to have a leukocyte stage, as expected in Theileria, do not infect humans and rodents as B. microti, and cluster phylogenetically as a separate species, this study proposes to name this parasite Babesia vulpes sp. nov., after its natural host, the red fox V. vulpes.

Background Infections with Babesia bovis , Babesia bigemina, Theileria species and Anaplasma marginale are endemic in Kenya yet there is a lack of adequate information on their genotypes. This study established the genetic diversities of the above tick-borne hemoparasites infecting cattle in Kenya. Methods Nested PCR and sequencing were used to determine the prevalence and genetic diversity of the above parasites in 192 cattle blood samples collected from Ngong and Machakos farms. B. bovis spherical body protein 4, B. bigemina rhoptry-associated protein 1a, A. marginale major surface protein 5, Theileria spp. 18S rRNA, T. parva p104 and T. orientalis major piroplasm surface protein were used as the marker genes. Results B. bovis , B. bigemina , T. parva , T. velifera , T. taurotragi , T. mutans and A. marginale were prevalent in both farms, whereas T. ovis, Theileria sp . (buffalo) and T. orientalis were found only in Ngong farm. Co-infections were observed in more than 50 % of positive samples in both farms. Babesia parasites and A. marginale sequences were highly conserved while T. parva and T. orientalis were polymorphic. Cattle-derived T. parva was detected in Machakos farm. However, cattle and buffalo–derived Theileria were detected in Ngong farm suggesting interactions between cattle and wild buffaloes. Generally, the pathogens detected in Kenya were genetically related to the other African isolates but different from the isolates in other continents. Conclusions The current findings reaffirm the endemicity and co-infection of cattle with tick-borne hemoparasites, and the role of wildlife in pathogens transmission and population genetics in Kenya.

For most of the 20th century the causative agent of canine babesiosis, wherever it occurred in the world, was commonly referred to as Babesia canis . Early research, from the 1890s to the 1930s, had shown that there were three distinctly different vector-specific parasite entities occurring in specific geographical regions, that host response to infection ranged from subclinical to acute, and that immunity to one stock of the parasite did not necessarily protect against infection with other stocks. This substantial body of knowledge was overlooked or ignored for 50 years. In this review the first records and descriptions of the disease in four geographical regions were traced: sub-Saharan Africa, Europe, North Africa and Asia. Research leading to identification of the specific tick vector species involved is documented. Evidence is given of the growing realisation that there were substantial biological differences between stocks originating from different geographical regions. Etymological provenance for Babesia vogeli is proposed.

We report a case of autochthonous human babesiosis in Hungary, confirmed by PCR and partial sequencing of the Babesia spp. 18S rRNA gene. Babesiosis should be considered during the differential diagnosis of febrile illnesses, and peripheral blood smears to detect Babesia spp. should be part of the routine clinical workup.

Introduction Babesiosis is a protozoan tick-borne infection associated with anemia and life-threatening disease in humans, domestic and wildlife animals. Dogs are infected by at least six well-characterized Babesia spp. that cause clinical disease. Infection with a piroplasmid species was detected by light microscopy of stained blood smears from five sick dogs from Israel and prompted an investigation on the parasite’s identity. Methods Genetic characterization of the piroplasmid was performed by PCR amplification of the 18S rRNA and the cytochrome c oxidase subunit 1 ( cox 1) genes, DNA sequencing and phylogenetic analysis. Four of the dogs were co-infected with Borrelia persica (Dschunkowsky, 1913), a relapsing fever spirochete transmitted by the argasid tick Ornithodoros tholozani Laboulbène & Mégnin. Co-infection of dogs with B. persica raised the possibility of transmission by O. tholozani and therefore, a piroplasmid PCR survey of ticks from this species was performed. Results The infected dogs presented with fever (4/5), anemia, thrombocytopenia (4/5) and icterus (3/5). Comparison of the 18S rRNA and cox 1 piroplasmid gene sequences revealed 99–100% identity between sequences amplified from different dogs and ticks. Phylogenetic trees demonstrated a previously undescribed species of Babesia belonging to the western group of Babesia ( sensu lato ) and closely related to the human pathogen Babesia duncani Conrad, Kjemtrup, Carreno, Thomford, Wainwright, Eberhard, Quick, Telfrom & Herwalt, 2006 while more moderately related to Babesia conradae Kjemtrup, Wainwright, Miller, Penzhorn & Carreno, 2006 which infects dogs. The piroplasm forms detected included tetrads (Maltese cross), merozoite and trophozoite stages whose average size was larger than stages of other canine Babesia spp. belonging to the Babesia ( s.l .) and B. gibsoni Patton, 1910, and smaller than other canine Babesia ( sensu stricto ) spp. Of 212 O. tholozani ticks surveyed, 11 (5.2%) harbored DNA of the new species of Babesia . Conclusions Babesia negevi n. sp. is described based on morphological and genetic characterization and phylogenetic analyses. The species is named after the Negev desert of southern Israel, where the first infected dog originated from. Despite co-infection in four dogs, the fifth dog had fatal disease attesting that B. negevi n. sp. infection requires clinical attention. Incriminating O. tholozani or another tick species as the vector of Babesia negevi n. sp., would require additional studies.

Background Bovine babesiosis is caused by infection with the protozoal parasite Babesia bovis , which is transmitted by Rhipicephalus ( Boophilus ) spp. It can cause mortality rates up to 90% in immunologically naive Bos taurus cattle. In south Texas, R . ( B .) microplus is known to infest nilgai antelope ( Boselaphus tragocamelus ); however, their susceptibility to infection with B .  bovis and their role in the transmission of the parasite remain unknown. In this study, we challenged nilgai antelope with B .  bovis and evaluated their susceptibility to infection. Methods Nilgai were needle inoculated with ≈10 8 B .  bovis -parasitized erythrocytes (merozoites) or a homogenate of B .  bovis -infected larval ticks (sporozoite) delivered intravenously. Bos taurus beef calves were inoculated in parallel, as this strain of B .  bovis is lethal to cattle. Temperature and hematocrit were monitored daily over the course of each study, and whole blood was collected for molecular [polymerase chain reaction (PCR)] and serological [indirect enzyme-linked immunosorbent assay (ELISA)] diagnostic evaluation. Histological sections of nilgai cerebral tissue were examined for evidence of infection. Recipient bovine calves were sub-inoculated with blood from nilgai challenged with either stage of the parasite, and they were monitored for clinical signs of infection and evaluated by a PCR diagnostic assay. Red blood cells (RBCs) from prechallenged nilgai and B .  taurus beef cattle were cultured with an in vitro B .  bovis merozoite culture to examine colonization of the RBCs by the parasite. Results Nilgai did not display clinical signs of infection upon inoculation with either the merozoite or sporozoite stage of B .  bovis . All nilgai were PCR-negative for the parasite, and they did not develop antibodies to B .  bovis . No evidence of infection was detected in histological sections of nilgai tissues, and in vitro culture analysis indicated that the nilgai RBCs were not colonized by B .  bovis merozoites. Cattle subinoculated with blood from challenged nilgai did not display clinical signs of infection, and they were PCR-negative up to 45 days after transfer. Conclusions Nilgai do not appear to be susceptible to infection with a strain of B .  bovis that is lethal to cattle. Tick control on these alternative hosts remains a critical priority, especially given their potential to disseminate ticks over long distances. Graphical Abstract

We have designed and performed a new PCR method based on the 18S rRNA in order to individuate the presence and the identity of Babesia parasites. Out of 1159 Ixodes ricinus (Acari: Ixodidae) ticks collected in four areas of Switzerland, nine were found to contain Babesia DNA. Sequencing of the short amplicon obtained (411-452 bp) allowed the identification of three human pathogenic species: Babesia microti, B. divergens, for the first time in Switzerland, Babesia sp. EU1. We also report coinfections with B. sp. EU1-Borrelia burgdorferi sensu stricto and Babesia sp. EU1-B. afzelii.

Background Babesiosis is an important haemoparasitic disease, caused by the infection and subsequent intra-erythrocytic multiplication of protozoa of the genus Babesia that impacts the livestock industry and animal health. The distribution, epidemiology and genetic characterization of B. bigemina , B. bovis , and B. ovata in cattle in China as well as the prevalence of these protozoan agents were assessed. Methods A total of 646 blood specimens from cattle, dairy cattle and yaks from 14 provinces were collected and tested for the presence of the three Babesia species via a specific nested PCR assay based on the rap-1 and ama-1 genes. The PCR results were confirmed by DNA sequencing. Gene sequences and the genetic characterization were determined for selected positive samples from each sampling area. Results Of a total of 646 samples, 134 (20.7 %), 60 (9.3 %) and 10 (1.5 %) were positive for B. bovis , B. bigemina and B. ovata infections, respectively. Mixed infections were found in 7 of 14 provinces; 43 (6.7 %) samples were infected with B. bovis and B. bigemina . Three samples (0.5 %) exhibited a co-infection with B. bovis and B. ovata , and 6 (0.9 %) were infected with all three parasites. The rap-1a gene of B. bovis indicated a high degree of sequence heterogeneity compared with other published rap-1a sequences worldwide and was 85–100 % identical to B. bovis rap-1a sequences in Chinese isolates. B. bigemina rap-1c and B. ovata ama-1 genes were nearly identical, with 97.8–99.3 % and 97.8–99.6 % sequence identity, respectively, in GenBank. Conclusions Positive rates of B. bovis and B. bigemina infection are somewhat high in China. The B. bovis infection in yaks was first reported. The significant sequence heterogeneity in different variants of the rap-1a gene from Chinese B. bovis isolates might be a great threat to the cattle industry if RAP-1a protein is used as immunological antigen against Babesia infections in China. The data obtained in this study can be used to plan effective control strategies against babesiosis in China.

Background Babesiosis, a tick-borne disease caused by protozoans of the genus Babesia , is widespread in subtropical and tropical countries. Mitochondria are essential organelles that are responsible for energy transduction and metabolism, calcium homeostasis and cell signaling. Mitochondrial genomes could provide new insights to help elucidate and investigate the biological features, genetic evolution and classification of the protozoans. Nevertheless, there are limited data on the mitochondrial genomes of ovine Babesia spp. in China. Methods Herein, we sequenced, assembled and annotated the mitochondrial genomes of six ovine Babesia isolates; analyzed the genome size, gene content, genome structure and cytochrome b ( cytb ) amino acid sequences and performed comparative mitochondrial genomics and phylogenomic analyses among apicomplexan parasites. Results The mitochondrial genomes range from 5767 to 5946 bp in length with a linear form and contain three protein-encoding genes, cytochrome c oxidase subunit 1 ( cox 1), cytochrome c oxidase subunit 3 ( cox 3) and cytb , six large subunit rRNA genes ( LSU ) and two terminal inverted repeats (TIR) on both ends. The cytb gene sequence analysis indicated the binding site of anti- Babesia drugs that targeted the cytochrome bc 1 complex. Babesia microti and Babesia rodhaini have a dual flip-flop inversion of 184–1082 bp, whereas other Babesia spp. and Theileria spp. have one pair of TIRs, 25–1563 bp. Phylogenetic analysis indicated that the six ovine Babesia isolates were divided into two clades, Babesia sp. and Babesia motasi . Babesia motasi isolates were further separated into two small clades ( B. motasi Hebei/Ningxian and B. motasi Tianzhu/Lintan). Conclusions The data provided new insights into the taxonomic relationships and drug targets of apicomplexan parasites.