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This study aimed to fill a gap of knowledge by providing a quantitative measure of molecularly identified species and genotypes belonging to Echinococcus granulosus sensu lato (s.l.) causing human cystic echinococcosis (CE) in Europe during the period 2000-2021. As these species and genotypes are characterized by genetic, animal host and geographical differences, studying the E. granulosus s.l. complex is epidemiologically relevant. A systematic review (SR) was conducted on the basis of both scientific and grey literature considering primary studies between 2000 and 2021 in four databases. From a total of 1643 scientific papers, 51 records were included in the SR. The main inclusion criterion for this study was the molecular confirmation of E. granulosus s.l. at the genotype/species level as a causative agent of human CE cases in selected European countries. Relevant data were obtained from 29 out of 39 eligible European countries. This SR identified 599 human molecularly confirmed echinococcal cysts: 460 (76.8%) identified as E. granulosus sensu stricto (s.s.), 130 (21.7%) as E. canadensis cluster (G6/7 and G10), 7 (1.2%) as E. ortleppi (G5), and 2 as E. vogeli (0.3%). Three geographical hotspots of human CE caused by different species of the E. granulosus s.l. complex were identified: (1) E. granulosus s.s. in Southern and South-eastern Europe (European-Mediterranean and Balkan countries); (2) E. canadensis (G6/7) in Central and Eastern Europe; (3) E. ortleppi in Central and Western Europe. This SR also identified data gaps that prevented a better definition of the geographical distribution of the Echinococcus granulosus s.l. species complex in Europe: western Balkan countries, part of Central Europe, and Baltic countries. These results mandate longitudinal, multi-centre, intersectoral and transdisciplinary studies which consider both molecular and clinical epidemiology in animals and humans. Such studies would be valuable for a better understanding of the transmission of the E. granulosus s.l. species complex and their potential clinical impact on humans.

The island of Cyprus was a historical endemic area for cystic echinococcosis (CE) in the Mediterranean. During the last decades, Cyprus has been an open-air laboratory and a model for testing and implementing control measures aiming to eliminate CE as a public health problem. Despite control and surveillance measures implemented during last 50 years, molecular characterization of Echinococcus granulosus sensu lato specimens has been never provided. In February 2023, the carcass of a stray dog collected in the Nicosia district was examined by the Veterinary Services and found infected with Echinococcus spp. worms. The worms were sent to the European Union Reference Laboratory (EURLP) for species/genotype identification. The sequences analyses of nad2 and nad5 genes allowed us to identify the tapeworms as Echinococcus canadensis, genotype G7b. In November 2023, a parasitic liver cyst was observed during the post-mortem examination of a mouflon from the same area of the dog's finding. The cyst sample was also referred to EURLP for identification and comparison with tapeworms previously collected from the dog. The sequences analysis of cox1 gene allowed to identify the cyst as E. granulosus sensu stricto, genotype G1. The finding of 2 different species of E. granulosus s.l. in a limited area raises epidemiological questions on the origin of the samples: whether distinct transmission cycles are present or a recent introduction event have occurred. From a public health perspective, it will be essential to conduct further molecular epidemiology studies to clarify the recent transmission dynamics of Echinococcus species in Cyprus.

Background The Three-River-Source Region of the Qinghai-Tibet Plateau is a hyperendemic focus for echinococcosis, with Echinococcus granulosus , E. multilocularis , and E. shiquicus circulating between definitive canid hosts (dogs and foxes) and intermediate hosts (livestock and rodents). However, the extent of environmental contamination by Echinococcus eggs remains understudied and poses significant risks to human and animal health. Methods From 2019 to 2021, we collected 631 canid fecal samples (296 from dogs and 335 from foxes) and 398 adjacent soil samples across endemic counties in the Three-River-Source Region of the Qinghai-Tibet Plateau. Multiplex real-time PCR was employed to detect Echinococcus species DNA in feces and soil samples. Results The overall Echinococcus prevalence in canid feces was 7.13% (45/631), with 3.01% in E. multilocularis (19/631), 2.06% in E. granulosus (13/631), and 2.06% in E. shiquicus (13/631). Foxes presented increased E. multilocularis (3.88%, 13/335) and E. shiquicus (2.69%, 9/335) infections, whereas dogs presented increased E. granulosus prevalence (2.70%, 8/296). Soil contamination with  Echinococcus  species was detected in 2.51% (10/398) of the samples. The primary contaminants were  E. multilocularis  and  E. shiquicus  (1.01% each, 4/398), whereas  E. granulosus  was less frequent (0.50%, 2/398). Moreover, the soil near fox feces was contaminated with both E. multilocularis and E. shiquicus , whereas the dog-associated soil was contaminated with all three species. Conclusions This study suggests widespread environmental deposition of Echinococcus eggs on the Qinghai-Tibet Plateau, driven by canid defecation. If these eggs remain viable, their persistence in soil would indicate a potential zoonotic transmission risk, highlighting the need for integrated control strategies targeting both domestic and wild canids. Graphical abstract

The parasitic species of the Echinococcus granulosus sensu lato (sl) complex are the causative agents of cystic echinococcosis in humans. The lifecycle of E. granulosus sl is essentially domestic, and is based on the consumption by dogs of hydatid cysts in viscera of livestock species. The aim of this study was to survey E. granulosus sensu lato in livestock in France. A 1-year national survey of E. granulosus sl in livestock at the slaughterhouse was organized in 2012 in France, with systematic molecular confirmation. The prevalence of E. granulosus ss nationally was 0.002% in sheep, mainly focused in the Alpine area, and 0.001% in cattle, with the distribution of cases throughout the country. Echinococcus canadensis G6/7 was observed only in Corsica in pigs, with a prevalence of nearly 1% in the island. A national prevalence of 0.0002% was estimated for E. ortleppi in cattle, due to seven cases distributed in two foci. The results of this survey are of particular interest because of the zoonotic risk associated with the presence of these parasite species, for which systematic control at the slaughterhouse should enable their elimination.

Background Echinococcosis, which is caused by the larvae of cestodes of the genus Echinococcus , is a parasitic zoonosis that poses a serious threat to the health of humans and animals globally. Albendazole is the drug of choice for the treatment of echinococcosis, but it is difficult to meet clinical goals with this chemotherapy due to its low cure rate and associated side effects after its long-term use. Hence, novel anti-parasitic targets and effective treatment alternatives are urgently needed. A previous study showed that verapamil (Vepm) can suppress the growth of Echinococcus granulosus larvae; however, the mechanism of this effect remains unclear. The aim of the present study was to gain insight into the anti-echinococcal effect of Vepm on Echinococcus with a particular focus on the regulatory effect of Vepm on calcium/calmodulin-dependent protein kinase II (Ca 2+ /CaM-CaMKII) in infected mice. Methods The anti-echinococcal effects of Vepm on Echinococcus granulosus protoscoleces (PSC) in vitro and Echinococcus multilocularis metacestodes in infected mice were assessed. The morphological alterations in Echinococcus spp. induced by Vepm were observed by scanning electron microscopy (SEM), and the changes in calcium content in both the parasite and mouse serum and liver were measured by SEM-energy dispersive spectrometry, inductively coupled plasma mass spectrometry and alizarin red staining. Additionally, the changes in the protein and mRNA levels of CaM and CaMKII in infected mice, and in the mRNA levels of CaMKII in E. granulosus PSC, were evaluated after treatment with Vepm by immunohistochemistry and/or real-time quantitative polymerase chain reaction. Results In vitro , E. granulosus PSC could be killed by Vepm at a concentration of 0.5 μg/ml or higher within 8 days. Under these conditions, the ultrastructure of PSC was damaged, and this damage was accompanied by obvious calcium loss and downregulation of CaMKII mRNA expression. In vivo , the weight and the calcium content of E. multilocularis metacestodes from mice were reduced after treatment with 40 mg/kg Vepm, and an elevation of the calcium content in the sera and livers of infected mice was observed. In addition, downregulation of CaM and CaMKII protein and mRNA expression in the livers of mice infected with E. multilocularis metacestodes was found after treatment with Vepm. Conclusions Vepm exerted a parasiticidal effect against Echinococcus both in vitro and in vivo through downregulating the expression of Ca 2+ /CaM-CaMKII, which was over-activated by parasitic infection. The results suggest that Ca 2+ /CaM-CaMKII may be a novel drug target, and that Vepm is a potential anti-echinococcal drug for the future control of echinococcosis. Graphical Abstract

Background Canids are definitive hosts of Echinococcus multilocularis and Echinococcus granulosus. This study aimed to survey these two Echinococcus species in canids of North-Khorasan Province, northeastern Iran, using morphological criteria and genetic characterization of mitochondrial DNA. Methods The carcasses of 106 canids, namely 61 jackals ( Canis aureus ), 23 foxes ( Vulpes vulpes ), 19 dogs ( Canis familiaris ) and three wolves ( Canis lupus ) were collected from the study area in 2013–2014 and examined for Echinococcus species. Morphological features were assessed by microscopy of adult worms. For molecular characterization, DNA was extracted, mostly from the adult worms but also from eggs. DNA fragments of the cytochrome c oxidase subunit 1 ( cox 1) and NADH dehydrogenase subunit 1 ( nad 1) mitochondrial genes were amplified and sequenced. Sequences were aligned and compared with reference sequences. Intraspecific and interspecific diversity were calculated and phylogenetic analysis was performed. Results Overall, 9.4% of the canids (eight jackals and two foxes) were found infected with E. multilocularis by molecular methods, of which seven cases were also confirmed using morphological description of the adult worms. Echinococcus granulosus was found in 6.6% of the canines (four dogs, two jackals and one wolf) as determined by both molecular methods and adult cestode morphology. All E. granulosus isolates were identified as the G1 genotype. Comparative sequence analysis indicated 0–0.7% and 0% intraspecific divergence within E. granulosus isolates and 0% and 0–0.2% within E. multilocularis isolates for cox 1 and nad 1, respectively. Conclusions This study revealed the presence of E. multilocularis and E. granulosus in canids of North-Khorasan Province of Iran. Jackals were found infected with both E. multilocularis and E. granulosus , but infection with the former species was higher.

Cystic echinococcosis is a neglected, zoonotic disease in Turkey. The disease is commonly seen in rural areas where the local population is in close contact with livestock and dogs. This research aimed to molecularly identify of hydatid cysts in cattle and human isolates from Konya, Turkey. Following sample collection, direct microscopy was performed. After direct examination, total DNA was extracted, and positive PCR products of cox 1 mitochondrial gene (~ 875 bp) were sequenced. A total of 83 hydatid cysts (cattle n = 57 and human n = 26), 82 were identified as Echinococcus granulosus sensu stricto (G1–G3 genotypes), and one human isolate was characterized as Echinococcus equinus (G4 genotype). Fertility rates of cysts belonging to cattle for liver and lung cysts were 93.3% and 80%, respectively. Out of 26 human originated isolates, 18 (69.2%) of cysts were found to be fertile. To the best of our knowledge, this is the first report of E. equinus from human host in Turkey.

Cystic and alveolar echinococcosis are considered the second and third most significant foodborne parasitic diseases worldwide. The microscopic eggs excreted in the feces of the definitive host are the only source of contamination for intermediate and dead-end hosts, including humans. However, estimating the respective contribution of the environment, fomites, animals or food in the transmission of Echinococcus eggs is still challenging. Echinococcus granulosus and E. multilocularis seem to have a similar survival capacity regarding temperature under laboratory conditions. In addition, field experiments have reported that the eggs can survive several weeks to years outdoors, with confirmation of the relative susceptibility of Echinococcus eggs to desiccation. Bad weather (such as rain and wind), invertebrates and birds help scatter Echinococcus eggs in the environment and may thus impact human exposure. Contamination of food and the environment by taeniid eggs has been the subject of renewed interest in the past decade. Various matrices from endemic regions have been found to be contaminated by Echinococcus eggs. These include water, soil, vegetables and berries, with heterogeneous rates highlighting the need to acquire more robust data so as to obtain an accurate assessment of the risk of human infection. In this context, it is essential to use efficient methods of detection and to develop methods for evaluating the viability of eggs in the environment and food.

Alveolar echinococcosis (AE) is a life-threatening disease in humans caused by the larval stage of . Domestic animals, dogs, foxes, and small mammals constitute the circular chain of AE. To evaluate the infection, distribution, and genetic polymorphism of AE in the Ili Prefecture (Nilka, Xinyuan and Zhaosu), we conducted this survey. In June and July 2018, 267 small mammals were captured using water-infusion and mousetrap methods. Combined pathogenic and molecular biological methods were used to observe the histopathology of carried by rodents, amplify the mitochondrial 1 gene of the pathogen, and investigate the genotype and haplotype diversity of in rodents in Ili Prefecture. Morphological identification revealed that these captured small mammals belonged to three species, with being the dominant species (183/267). Pathological and molecular biological results confirmed that was the pathogen of echinococcosis in small mammals, with an infection rate of 15.73% (42/267). Among the three areas sampled, the highest infection rate of rodents was 25.45% (14/55) in Nilka County. However, there was no significant difference in the infection rates between regions (χ = 5.119, p > 0.05). Of the three captured rodent species, had the highest infection rate of 17.49% (32/183), but there was no significant difference in infection rates between the rodent species (χ = 1.364, p > 0.05). Phylogenetic analyses showed that the nad1 gene sequences obtained in this study clustered in the same clade as isolates from China. These isolates contained 21 haplotypes (Hap_1-21); Hap_2 was the most common haplotype (9/42). Furthermore, haplotype diversity (0.925 ± 0.027) and nucleotide diversity (0.01139 ± 0.00119) were higher in the Ili Prefecture than in other regions, indicating that population differentiation was high. Tajima's D and Fu's Fs tests were negative (p > 0.10), indicating that the population had expanded. The low fixation index (Fst) ranged from 0.00000 to 0.16945, indicating that the degree of genetic differentiation was different among different populations. In summary, Ili Prefecture is a high incidence area of AE, and Microtus spp. may play an important role in the transmission of AE in this area. The results of this study provide basic data for further study of the molecular epidemiology, genetic differences, and control of in the Ili Prefecture, Xinjiang.

Echinococcus granulosus sensu lato ( s.l. ) is a species complex with the potential to cause cystic echinococcosis (CE). Contact with the feces of domestic dogs ( Canis familiaris ) fed with raw viscera of intermediate livestock hosts is a risk factor for this infection in the southern region of Brazil. Although the region has been considered endemic to CE for many years, molecular data regarding the species of the complex causing CE in humans are scarce. This study aimed to perform a molecular analysis of the biological fluid from a human liver cyst to investigate the species responsible for CE. Genetic material obtained from the hydatid fluid of a hepatic cyst from a human with CE was subjected to PCR to amplify mitochondrial and nuclear DNA sequences. The phylogenetic analysis confirmed the human infection by Echinococcus canadensis G7 in the state of Paraná, Brazil. This is the first molecular record of E. canadensis G7 infecting a human in Brazil, and it is important to reiterate the risk of human CE caused by this species in South America, as reported by a previous study in Patagonia, Argentina. From the epidemiological point of view, this finding is of great relevance for the southern region of Brazil, since this parasite has previously only been detected in pigs in the state of Rio Grande do Sul, neighboring Paraná. The finding points to the importance of this identification in the molecular epidemiology of E. granulosus s.l. , especially in South America.