{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T12:48:20Z","timestamp":1792586900916,"version":"4.5.0"},"reference-count":111,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T00:00:00Z","timestamp":1791417600000},"content-version":"vor","delay-in-days":7,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"content-domain":{"domain":["cell.com","clinicalkey.com","clinicalkey.com.au","clinicalkey.es","clinicalkey.fr","clinicalkey.jp","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Trends in Parasitology"],"published-print":{"date-parts":[[2026,10]]},"DOI":"10.1016\/j.pt.2026.09.018","type":"journal-article","created":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T11:27:23Z","timestamp":1792582043000},"update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["Friend or foe? Functional roles of Ixodes ricinus microbes"],"prefix":"10.1016","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8608-0387","authenticated-orcid":false,"given":"Aleksandra I.","family":"Krawczyk","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Olivier","family":"Duron","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Judith M.","family":"H\u00fcbschen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.pt.2026.09.018_bb0005","doi-asserted-by":"crossref","first-page":"1946","DOI":"10.1016\/S0140-6736(12)61151-9","article-title":"Drivers, dynamics, and control of emerging vector-borne zoonotic diseases","volume":"380","author":"Kilpatrick","year":"2012","journal-title":"Lancet"},{"key":"10.1016\/j.pt.2026.09.018_bb0010","doi-asserted-by":"crossref","first-page":"S150","DOI":"10.1016\/j.actatropica.2013.11.004","article-title":"Harnessing mosquito\u2013Wolbachia symbiosis for vector and disease control","volume":"132","author":"Bourtzis","year":"2014","journal-title":"Acta Trop."},{"key":"10.1016\/j.pt.2026.09.018_bb0015","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1016\/j.pt.2019.12.001","article-title":"Mosquito microbiota and implications for disease control","volume":"36","author":"Gao","year":"2020","journal-title":"Trends Parasitol."},{"key":"10.1016\/j.pt.2026.09.018_bb0020","article-title":"Dengue virus infection modifies mosquito blood-feeding behavior to increase transmission to the host","volume":"119","author":"Xiang","year":"2022","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.pt.2026.09.018_bb0025","doi-asserted-by":"crossref","first-page":"1050","DOI":"10.1016\/j.pt.2021.06.007","article-title":"Wolbachia as translational science: controlling mosquito-borne pathogens","volume":"37","author":"Caragata","year":"2021","journal-title":"Trends Parasitol."},{"key":"10.1016\/j.pt.2026.09.018_bb0030","doi-asserted-by":"crossref","first-page":"1357","DOI":"10.3389\/fmicb.2020.01357","article-title":"Dynamics of insect-microbiome interaction influence host and microbial symbiont","volume":"11","author":"Gupta","year":"2020","journal-title":"Front. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0035","doi-asserted-by":"crossref","first-page":"1798","DOI":"10.1038\/s41396-023-01500-2","article-title":"Impact of intraspecific variation in insect microbiomes on host phenotype and evolution","volume":"17","author":"Lange","year":"2023","journal-title":"ISME J."},{"key":"10.1016\/j.pt.2026.09.018_bb0040","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1186\/1741-7007-6-27","article-title":"The diversity of reproductive parasites among arthropods: Wolbachia do not walk alone","volume":"6","author":"Duron","year":"2008","journal-title":"BMC Biol."},{"key":"10.1016\/j.pt.2026.09.018_bb0045","doi-asserted-by":"crossref","first-page":"2905","DOI":"10.1111\/mec.14094","article-title":"Evolutionary changes in symbiont community structure in ticks","volume":"26","author":"Duron","year":"2017","journal-title":"Mol. Ecol."},{"key":"10.1016\/j.pt.2026.09.018_bb0050","doi-asserted-by":"crossref","first-page":"1016","DOI":"10.1111\/mec.15373","article-title":"Microbial community structure reveals instability of nutritional symbiosis during the evolutionary radiation of Amblyomma ticks","volume":"29","author":"Binetruy","year":"2020","journal-title":"Mol. Ecol."},{"key":"10.1016\/j.pt.2026.09.018_bb0055","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1146\/annurev.ecolsys.110308.120206","article-title":"The ecology and evolution of microbes that manipulate host reproduction","volume":"40","author":"Engelst\u00e4dter","year":"2009","journal-title":"Annu. Rev. Ecol. Evol. Syst."},{"key":"10.1016\/j.pt.2026.09.018_bb0060","doi-asserted-by":"crossref","first-page":"R485","DOI":"10.1016\/j.cub.2019.03.032","article-title":"The life of an insect endosymbiont from the cradle to the grave","volume":"29","author":"McCutcheon","year":"2019","journal-title":"Curr. Biol."},{"key":"10.1016\/j.pt.2026.09.018_bb0065","doi-asserted-by":"crossref","first-page":"816","DOI":"10.1016\/j.pt.2020.07.007","article-title":"Convergence of nutritional symbioses in obligate blood feeders","volume":"36","author":"Duron","year":"2020","journal-title":"Trends Parasitol."},{"key":"10.1016\/j.pt.2026.09.018_bb0070","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1111\/1365-2435.12133","article-title":"Defensive symbiosis in the real world-advancing ecological studies of heritable, protective bacteria in aphids and beyond","volume":"28","author":"Oliver","year":"2014","journal-title":"Funct. Ecol."},{"key":"10.1016\/j.pt.2026.09.018_bb0075","doi-asserted-by":"crossref","first-page":"1747","DOI":"10.1111\/brv.12417","article-title":"The role of defensive symbionts in host-parasite coevolution","volume":"93","author":"Vorburger","year":"2018","journal-title":"Biol. Rev."},{"key":"10.1016\/j.pt.2026.09.018_bb0080","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1011856","article-title":"Diversity and spread of cytoplasmic incompatibility genes among maternally inherited symbionts","volume":"21","author":"Amoros","year":"2025","journal-title":"PLoS Genet."},{"key":"10.1016\/j.pt.2026.09.018_bb0085","doi-asserted-by":"crossref","first-page":"1830","DOI":"10.3390\/ijerph17061830","article-title":"Management options for Ixodes ricinus-associated pathogens: a review of prevention strategies","volume":"17","author":"\u010cern\u00fd","year":"2020","journal-title":"Int. J. Environ. Res. Public Health"},{"key":"10.1016\/j.pt.2026.09.018_bb0090","doi-asserted-by":"crossref","DOI":"10.1111\/pim.12813","article-title":"Update on the intricate tango between tick microbiomes and tick-borne pathogens","volume":"43","author":"Bonnet","year":"2021","journal-title":"Parasite Immunol."},{"key":"10.1016\/j.pt.2026.09.018_bb0095","doi-asserted-by":"crossref","first-page":"696","DOI":"10.1016\/j.pt.2024.06.006","article-title":"Nutritional symbiosis in ticks: singularities of the genus Ixodes","volume":"40","author":"Duron","year":"2024","journal-title":"Trends Parasitol."},{"key":"10.1016\/j.pt.2026.09.018_bb0100","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1186\/s40168-022-01276-1","article-title":"Quantitative microbial population study reveals geographical differences in bacterial symbionts of Ixodes ricinus","volume":"10","author":"Krawczyk","year":"2022","journal-title":"Microbiome"},{"key":"10.1016\/j.pt.2026.09.018_bb0105","doi-asserted-by":"crossref","first-page":"236","DOI":"10.3389\/fcimb.2017.00236","article-title":"The tick microbiome: why non-pathogenic microorganisms matter in tick biology and pathogen transmission","volume":"7","author":"Bonnet","year":"2017","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0110","doi-asserted-by":"crossref","DOI":"10.3389\/fimmu.2021.704621","article-title":"Anti-microbiota vaccines modulate the tick microbiome in a taxon-specific manner","volume":"12","author":"Mateos-Hern\u00e1ndez","year":"2021","journal-title":"Front. Immunol."},{"key":"10.1016\/j.pt.2026.09.018_bb0115","doi-asserted-by":"crossref","first-page":"e02713","DOI":"10.1128\/AEM.02713-20","article-title":"Long-term survival of virulent tularemia pathogens outside a host in conditions that mimic natural aquatic environments","volume":"87","author":"Golovliov","year":"2021","journal-title":"Appl. Environ. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0120","doi-asserted-by":"crossref","DOI":"10.3389\/fvets.2022.1068129","article-title":"Coxiella burnetii in ticks, livestock, pets and wildlife: a mini-review","volume":"9","author":"Celina","year":"2022","journal-title":"Front. Vet. Sci."},{"key":"10.1016\/j.pt.2026.09.018_bb0125","doi-asserted-by":"crossref","first-page":"536","DOI":"10.1016\/j.pt.2015.06.014","article-title":"The importance of ticks in Q fever transmission: what has (and has not) been demonstrated?","volume":"31","author":"Duron","year":"2015","journal-title":"Trends Parasitol."},{"key":"10.1016\/j.pt.2026.09.018_bb0130","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1111\/j.1749-6632.1987.tb40616.x","article-title":"Transmission modes and evolution of the parasitism-mutualism continuum","volume":"503","author":"Ewald","year":"1987","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"10.1016\/j.pt.2026.09.018_bb0135","doi-asserted-by":"crossref","DOI":"10.24072\/pcjournal.633","article-title":"Comparative genomics of Rickettsiella bacteria reveal variable metabolic pathways potentially involved in symbiotic interactions with arthropods","volume":"5","author":"Floriano","year":"2025","journal-title":"Peer Community J."},{"key":"10.1016\/j.pt.2026.09.018_bb0140","doi-asserted-by":"crossref","first-page":"1070","DOI":"10.1016\/j.ttbdis.2019.05.019","article-title":"Tissue tropism and metabolic pathways of Midichloria mitochondrii suggest tissue-specific functions in the symbiosis with Ixodes ricinus","volume":"10","author":"Olivieri","year":"2019","journal-title":"Ticks Tick Borne Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0145","doi-asserted-by":"crossref","DOI":"10.3389\/fcimb.2022.1081666","article-title":"Ixodes ricinus ticks have a functional association with Midichloria mitochondrii","volume":"12","author":"Guizzo","year":"2023","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0150","article-title":"Midichloria mitochondrii stimulates the sylvatic cycle of Borrelia burgdorferi (sensu lato) in Ixodes ricinus and contributes to Lyme disease risk","volume":"8","author":"K\u00f6hler","year":"2025","journal-title":"Curr. Res. Parasitol. Vector Borne Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0155","doi-asserted-by":"crossref","first-page":"1280","DOI":"10.1111\/j.1462-2920.2006.01024.x","article-title":"Widespread distribution and high prevalence of an alpha-proteobacterial symbiont in the tick Ixodes ricinus","volume":"8","author":"Lo","year":"2006","journal-title":"Environ. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0160","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1007\/s00248-021-01869-7","article-title":"The genetic diversity of Rickettsiella symbionts in Ixodes ricinus throughout Europe","volume":"84","author":"Garcia-Vozmediano","year":"2022","journal-title":"Microb. Ecol."},{"key":"10.1016\/j.pt.2026.09.018_bb0165","doi-asserted-by":"crossref","first-page":"17","DOI":"10.52004\/2054930x-20241011","article-title":"Ixodes ricinus (Acari, Ixodidae) ticks and tick-borne pathogen prevalence on the Isle of Wight \u2013 an island perspective in southern England","volume":"43","author":"Medlock","year":"2024","journal-title":"J. Eur. Mosq. Control. Assoc."},{"key":"10.1016\/j.pt.2026.09.018_bb0170","article-title":"Spatial and temporal variation of five different pathogens and symbionts in Ixodes ricinus nymphs in the Netherlands","volume":"6","author":"Hartemink","year":"2024","journal-title":"Curr. Res. Parasitol. Vector Borne Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0175","doi-asserted-by":"crossref","DOI":"10.1128\/msystems.00507-25","article-title":"Deep sequencing of 16 Ixodes ricinus ticks unveils insights into their interactions with endosymbionts","volume":"10","author":"Lesiczka","year":"2025","journal-title":"mSystems"},{"key":"10.1016\/j.pt.2026.09.018_bb0180","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1186\/s13071-023-06025-3","article-title":"Differential associations of horizontally and vertically transmitted symbionts on Ixodes ricinus behaviour and physiology","volume":"16","author":"Bakker","year":"2023","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0185","doi-asserted-by":"crossref","first-page":"3285","DOI":"10.1093\/molbev\/msr159","article-title":"Phylogenomic evidence for the presence of a flagellum and cbb3 oxidase in the free-living mitochondrial ancestor","volume":"28","author":"Sassera","year":"2011","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.pt.2026.09.018_bb0190","doi-asserted-by":"crossref","first-page":"1681","DOI":"10.3201\/eid2808.212524","article-title":"Spiroplasma ixodetis infections in immunocompetent and immunosuppressed patients after tick exposure, Sweden","volume":"28","author":"Eimer","year":"2022","journal-title":"Emerg. Infect. Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0195","doi-asserted-by":"crossref","DOI":"10.1016\/j.isci.2026.115233","article-title":"A human-associated Spiroplasma ixodetis lineage responsible for infantile cataracts and adult febrile illness","volume":"29","author":"Buysse","year":"2026","journal-title":"Iscience"},{"key":"10.1016\/j.pt.2026.09.018_bb0200","doi-asserted-by":"crossref","DOI":"10.1093\/ismeco\/ycaf202","article-title":"Genetic and genomic variability of Spiroplasma and Midichloria endosymbionts associated with the tick Ixodes frontalis","volume":"5","author":"Melis","year":"2025","journal-title":"ISME Commun."},{"key":"10.1016\/j.pt.2026.09.018_bb0205","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1186\/1756-3305-7-128","article-title":"Birds as potential reservoirs of tick-borne pathogens: first evidence of bacteraemia with Rickettsia helvetica","volume":"7","author":"Hornok","year":"2014","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0210","first-page":"357","article-title":"Ixodes ricinus: vector of a hitherto undescribed spotted fever group agent in Switzerland","volume":"36","author":"Burgdorfer","year":"1979","journal-title":"Acta Trop."},{"key":"10.1016\/j.pt.2026.09.018_bb0215","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/1756-3305-6-1","article-title":"Driving forces for changes in geographical distribution of Ixodes ricinus ticks in Europe","volume":"6","author":"Medlock","year":"2013","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0220","doi-asserted-by":"crossref","first-page":"19","DOI":"10.2478\/s11686-018-00004-w","article-title":"Simultaneous occurrence of Borrelia miyamotoi, Borrelia burgdorferi sensu lato, Anaplasma phagocytophilum and Rickettsia helvetica in Ixodes ricinus ticks in urban foci in Bratislava, Slovakia","volume":"64","author":"Vaculov\u00e1","year":"2019","journal-title":"Acta Parasitol."},{"key":"10.1016\/j.pt.2026.09.018_bb0225","doi-asserted-by":"crossref","first-page":"19376","DOI":"10.1038\/s41598-020-76334-5","article-title":"Spatial patterns of pathogen prevalence in questing Ixodes ricinus nymphs in southern Scandinavia, 2016","volume":"10","author":"Kj\u00e6r","year":"2020","journal-title":"Sci. Rep."},{"key":"10.1016\/j.pt.2026.09.018_bb0230","doi-asserted-by":"crossref","first-page":"1042","DOI":"10.1055\/s-2003-39103","article-title":"Fever after a tick bite: clinical manifestations and diagnosis of acute tick bite-associated infections in northeastern Switzerland","volume":"128","author":"Baumann","year":"2003","journal-title":"Dtsch. Med. Wochenschr."},{"key":"10.1016\/j.pt.2026.09.018_bb0235","doi-asserted-by":"crossref","first-page":"389","DOI":"10.3201\/eid0604.000412","article-title":"Evidence of Rickettsia helvetica infection in humans, eastern France","volume":"6","author":"Fournier","year":"2000","journal-title":"Emerg. Infect. Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0240","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1093\/jmedent\/30.1.295","article-title":"Efficient transmission of tick-borne encephalitis virus between cofeeding ticks","volume":"30","author":"Labuda","year":"1993","journal-title":"J. Med. Entomol."},{"key":"10.1016\/j.pt.2026.09.018_bb0245","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1186\/1756-3305-5-177","article-title":"Prevalence of tick borne encephalitis virus in tick nymphs in relation to climatic factors on the southern coast of Norway","volume":"5","author":"Andreassen","year":"2012","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0250","doi-asserted-by":"crossref","first-page":"356","DOI":"10.1016\/j.ttbdis.2015.02.010","article-title":"Risk of exposure to ticks (Ixodidae) and the prevalence of tick-borne encephalitis virus (TBEV) in ticks in Southern Poland","volume":"6","author":"Cuber","year":"2015","journal-title":"Ticks Tick Borne Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0255","doi-asserted-by":"crossref","first-page":"126","DOI":"10.3390\/pathogens11020126","article-title":"New and confirmed foci of tick-borne encephalitis virus (TBEV) in Northern Germany determined by TBEV detection in ticks","volume":"11","author":"Topp","year":"2022","journal-title":"Pathogens"},{"key":"10.1016\/j.pt.2026.09.018_bb0260","doi-asserted-by":"crossref","DOI":"10.1016\/j.ttbdis.2020.101594","article-title":"Prevalence of tick-borne encephalitis virus in questing Dermacentor reticulatus and Ixodes ricinus ticks in Lithuania","volume":"12","author":"Sidorenko","year":"2021","journal-title":"Ticks Tick Borne Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0265","doi-asserted-by":"crossref","DOI":"10.2807\/1560-7917.ES.2018.23.45.1800201","article-title":"Tick-borne encephalitis in Europe, 2012 to 2016","volume":"23","author":"Beaut\u00e9","year":"2018","journal-title":"Euro Surveill."},{"key":"10.1016\/j.pt.2026.09.018_bb0270","doi-asserted-by":"crossref","DOI":"10.1016\/j.ttbdis.2024.102388","article-title":"Publicly available surveillance data on tick-borne encephalitis in Europe, 2023","volume":"15","author":"Angulo","year":"2024","journal-title":"Ticks Tick Borne Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0275","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1016\/j.ttbdis.2012.05.005","article-title":"Different tick-borne encephalitis virus (TBEV) prevalences in unfed versus partially engorged ixodid ticks \u2013 evidence of virus replication and changes in tick behavior","volume":"3","author":"Belova","year":"2012","journal-title":"Ticks Tick Borne Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0280","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1186\/s13071-016-1389-5","article-title":"Larvae of Ixodes ricinus transmit Borrelia afzelii and B. miyamotoi to vertebrate hosts","volume":"9","author":"van Duijvendijk","year":"2016","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0285","doi-asserted-by":"crossref","first-page":"1079","DOI":"10.1017\/S0950268814001691","article-title":"Borrelia miyamotoi in host-seeking Ixodes ricinus ticks in England","volume":"143","author":"Hansford","year":"2015","journal-title":"Epidemiol. Infect."},{"key":"10.1016\/j.pt.2026.09.018_bb0290","doi-asserted-by":"crossref","first-page":"e772","DOI":"10.1016\/S2666-5247(22)00157-4","article-title":"Prevalence and clinical manifestation of Borrelia miyamotoi in Ixodes ticks and humans in the northern hemisphere: a systematic review and meta-analysis","volume":"3","author":"Hoornstra","year":"2022","journal-title":"Lancet Microbe"},{"key":"10.1016\/j.pt.2026.09.018_bb0295","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1017\/S0031182006001545","article-title":"Transstadial and transovarial persistence of Babesia divergens DNA in Ixodes ricinus ticks fed on infected blood in a new skin-feeding technique","volume":"134","author":"Bonnet","year":"2007","journal-title":"Parasitology"},{"key":"10.1016\/j.pt.2026.09.018_bb0300","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1051\/vetres\/2009004","article-title":"Experimental in vitro transmission of Babesia sp. (EU1) by Ixodes ricinus","volume":"40","author":"Bonnet","year":"2009","journal-title":"Vet. Res."},{"key":"10.1016\/j.pt.2026.09.018_bb0305","first-page":"228","article-title":"A study on the transmission (transstadial, transovarial) of Babesia microti, strain \u201cHannover i\u201d, in its tick vector, Ixodes ricinus (author\u2019s transl)","volume":"32","author":"Walter","year":"1981","journal-title":"Tropenmed. Parasitol."},{"key":"10.1016\/j.pt.2026.09.018_bb0310","doi-asserted-by":"crossref","first-page":"230","DOI":"10.3390\/pathogens10020230","article-title":"Global distribution of Babesia species in questing ticks: a systematic review and meta-analysis based on published literature","volume":"10","author":"Onyiche","year":"2021","journal-title":"Pathogens"},{"key":"10.1016\/j.pt.2026.09.018_bb0315","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1089\/vbz.2012.1278","article-title":"Diversity of Babesia and Rickettsia species in questing Ixodes ricinus: a longitudinal study in urban, pasture, and natural habitats","volume":"13","author":"Overzier","year":"2013","journal-title":"Vector-Borne Zoonotic Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0320","doi-asserted-by":"crossref","first-page":"386","DOI":"10.3390\/pathogens10040386","article-title":"Circulation of Babesia species and their exposure to humans through Ixodes ricinus","volume":"10","author":"Azagi","year":"2021","journal-title":"Pathogens"},{"key":"10.1016\/j.pt.2026.09.018_bb0325","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1017\/S0031182000060285","article-title":"The effect of Babesia microti on feeding and survival in its tick vector, Ixodes trianguliceps","volume":"102","author":"Randolph","year":"1991","journal-title":"Parasitology"},{"key":"10.1016\/j.pt.2026.09.018_bb0330","doi-asserted-by":"crossref","first-page":"1165","DOI":"10.3390\/pathogens10091165","article-title":"Human babesiosis in Europe","volume":"10","author":"Hildebrandt","year":"2021","journal-title":"Pathogens"},{"key":"10.1016\/j.pt.2026.09.018_bb0335","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1093\/jmedent\/33.1.189","article-title":"Efficient transmission of Borrelia burgdorferi between cofeeding Ixodes ricinus ticks (Acari: Ixodidae)","volume":"33","author":"Gern","year":"1996","journal-title":"J. Med. Entomol."},{"key":"10.1016\/j.pt.2026.09.018_bb0340","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1186\/s13071-016-1929-z","article-title":"Comparison of the lifetime host-to-tick transmission between two strains of the Lyme disease pathogen Borrelia afzelii","volume":"9","author":"Jacquet","year":"2016","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0345","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1080\/00365540110080313","article-title":"Three major Lyme Borrelia genospecies (Borrelia burgdorferi sensu stricto, B. afzelii and B. garinii) identified by PCR in cerebrospinal fluid from patients with neuroborreliosis in Sweden","volume":"34","author":"Ornstein","year":"2002","journal-title":"Scand. J. Infect. Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0350","doi-asserted-by":"crossref","first-page":"19906","DOI":"10.2807\/ese.16.27.19906-en","article-title":"Lyme borreliosis in Europe","volume":"16","author":"Rizzoli","year":"2011","journal-title":"Eurosurveillance"},{"key":"10.1016\/j.pt.2026.09.018_bb0355","doi-asserted-by":"crossref","first-page":"147","DOI":"10.14411\/fp.2000.029","article-title":"Peculiarities of behaviour of taiga (Ixodes persulcatus) and sheep (Ixodes ricinus) ticks (Acarina: Ixodidae) determined by different methods","volume":"47","author":"Alekseev","year":"2000","journal-title":"Folia Parasitol."},{"key":"10.1016\/j.pt.2026.09.018_bb0360","doi-asserted-by":"crossref","first-page":"1196","DOI":"10.1603\/ME10111","article-title":"Survival of Ixodes ricinus (Acari: Ixodidae) under challenging conditions of temperature and humidity is influenced by Borrelia burgdorferi sensu lato infection","volume":"47","author":"Herrmann","year":"2010","journal-title":"J. Med. Entomol."},{"key":"10.1016\/j.pt.2026.09.018_bb0365","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1017\/S0031182000066336","article-title":"The effect of infection with Lyme disease spirochetes (Borrelia burgdorferi) on the phototaxis, activity, and questing height of the tick vector Ixodes scapularis","volume":"113","author":"Lefcort","year":"1996","journal-title":"Parasitology"},{"key":"10.1016\/j.pt.2026.09.018_bb0370","first-page":"422","article-title":"A Borrelia afzelii infection increases larval tick burden on Myodes glareolus (Rodentia: Cricetidae) and nymphal body weight of Ixodes ricinus (Acari: Ixodidae)","volume":"54","author":"van Duijvendijk","year":"2017","journal-title":"J. Med. Entomol."},{"key":"10.1016\/j.pt.2026.09.018_bb0375","doi-asserted-by":"crossref","first-page":"903","DOI":"10.1016\/j.jinsphys.2012.04.004","article-title":"The correlation between tick (Ixodes persulcatus Sch.) questing behaviour and synganglion neuronal responses to odours","volume":"58","author":"Romashchenko","year":"2012","journal-title":"J. Insect Physiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0380","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1007\/s10493-008-9131-4","article-title":"Tick infestation risk and Borrelia burgdorferi s.l. infection-induced increase in host-finding efficacy of female Ixodes ricinus under natural conditions","volume":"44","author":"Faulde","year":"2008","journal-title":"Exp. Appl. Acarol."},{"key":"10.1016\/j.pt.2026.09.018_bb0385","doi-asserted-by":"crossref","first-page":"40","DOI":"10.3389\/fvets.2020.00040","article-title":"Experimental Ixodes ricinus-sheep cycle of Anaplasma phagocytophilum NV2Os propagated in tick cell cultures","volume":"7","author":"Almaz\u00e1n","year":"2020","journal-title":"Front. Vet. Sci."},{"key":"10.1016\/j.pt.2026.09.018_bb0390","doi-asserted-by":"crossref","first-page":"288","DOI":"10.1186\/s12879-020-04993-w","article-title":"Molecular diagnosis of autochthonous human anaplasmosis in Austria \u2013 an infectious diseases case report","volume":"20","author":"Hoepler","year":"2020","journal-title":"BMC Infect. Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0395","doi-asserted-by":"crossref","first-page":"599","DOI":"10.1186\/s13071-019-3852-6","article-title":"A review on the eco-epidemiology and clinical management of human granulocytic anaplasmosis and its agent in Europe","volume":"12","author":"Matei","year":"2019","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0400","doi-asserted-by":"crossref","first-page":"3179","DOI":"10.1172\/JCI42868","article-title":"Anaplasma phagocytophilum induces Ixodes scapularis ticks to express an antifreeze glycoprotein gene that enhances their survival in the cold","volume":"120","author":"Neelakanta","year":"2010","journal-title":"J. Clin. Invest."},{"key":"10.1016\/j.pt.2026.09.018_bb0405","article-title":"Comparative genomics of emerging human ehrlichiosis agents","volume":"2","author":"Hotopp","year":"2006","journal-title":"PLoS Genet."},{"key":"10.1016\/j.pt.2026.09.018_bb0410","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1111\/j.1365-2915.2011.00973.x","article-title":"Expression of heat shock proteins and subolesin affects stress responses, Anaplasma phagocytophilum infection and questing behaviour in the tick, Ixodes scapularis","volume":"26","author":"Busby","year":"2012","journal-title":"Med. Vet. Entomol."},{"key":"10.1016\/j.pt.2026.09.018_bb0415","doi-asserted-by":"crossref","first-page":"3154","DOI":"10.1074\/mcp.M115.051938","article-title":"Integrated metabolomics, transcriptomics and proteomics identifies metabolic pathways affected by Anaplasma phagocytophilum infection in tick cells","volume":"14","author":"Villar","year":"2015","journal-title":"Mol. Cell. Proteomics"},{"key":"10.1016\/j.pt.2026.09.018_bb0420","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0133038","article-title":"Nuclease Tudor-SN is involved in tick dsRNA-mediated RNA interference and feeding but not in defense against flaviviral or Anaplasma phagocytophilum rickettsial infection","volume":"10","author":"Ayll\u00f3n","year":"2015","journal-title":"PLoS One"},{"key":"10.1016\/j.pt.2026.09.018_bb0425","doi-asserted-by":"crossref","first-page":"23","DOI":"10.3389\/fcimb.2017.00023","article-title":"Anaplasma phagocytophilum infection subverts carbohydrate metabolic pathways in the tick vector, Ixodes scapularis","volume":"7","author":"Cabezas-Cruz","year":"2017","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0430","first-page":"563","article-title":"Candidatus Neoehrlichia mikurensis and Anaplasma phagocytophilum: prevalences and investigations on a new transmission path in small mammals and ixodid ticks","volume":"7","author":"Obiegala","year":"2014","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0435","doi-asserted-by":"crossref","first-page":"1956","DOI":"10.1128\/JCM.02423-09","article-title":"First case of human \u201cCandidatus Neoehrlichia mikurensis\u201d infection in a febrile patient with chronic lymphocytic leukemia","volume":"48","author":"Welinder-Olsson","year":"2010","journal-title":"J. Clin. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0440","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1016\/j.nmni.2017.12.011","article-title":"\u2018Candidatus Neoehrlichia mikurensis\u2019 in Europe","volume":"22","author":"Portillo","year":"2018","journal-title":"New Microbes New Infect."},{"key":"10.1016\/j.pt.2026.09.018_bb0445","doi-asserted-by":"crossref","first-page":"1100","DOI":"10.3390\/ijms24021100","article-title":"Revealing the tick microbiome: insights into midgut and salivary gland microbiota of female Ixodes ricinus ticks","volume":"24","author":"Wiesinger","year":"2023","journal-title":"Int. J. Mol. Sci."},{"key":"10.1016\/j.pt.2026.09.018_bb0450","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1007\/s00248-025-02571-8","article-title":"Diversity of the Ixodes ricinus microbiome across Belgian ecoregions and its association with pathogen and symbiont presence","volume":"88","author":"Philippe","year":"2025","journal-title":"Microb. Ecol."},{"key":"10.1016\/j.pt.2026.09.018_bb0455","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1111\/j.1749-6632.1975.tb35099.x","article-title":"Vectors and vertical transmission: an epidemiologic perspective","volume":"266","author":"Fine","year":"1975","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"10.1016\/j.pt.2026.09.018_bb0460","doi-asserted-by":"crossref","first-page":"638","DOI":"10.1128\/iai.27.2.638-642.1980","article-title":"Sexual transmission of spotted fever group rickettsiae by infected male ticks: detection of rickettsiae in immature spermatozoa of Ixodes ricinus","volume":"27","author":"Hayes","year":"1980","journal-title":"Infect. Immun."},{"key":"10.1016\/j.pt.2026.09.018_bb0465","doi-asserted-by":"crossref","first-page":"7433","DOI":"10.1073\/pnas.1402476111","article-title":"Intrasperm vertical symbiont transmission","volume":"111","author":"Watanabe","year":"2014","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.pt.2026.09.018_bb0470","doi-asserted-by":"crossref","first-page":"17007","DOI":"10.1038\/nmicrobiol.2017.7","article-title":"A Wolbachia deubiquitylating enzyme induces cytoplasmic incompatibility","volume":"2","author":"Beckmann","year":"2017","journal-title":"Nat. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0475","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1186\/1756-3305-6-350","article-title":"Molecular and serological evidence for the circulation of the tick symbiont Midichloria (Rickettsiales: Midichloriaceae) in different mammalian species","volume":"6","author":"Bazzocchi","year":"2013","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0480","doi-asserted-by":"crossref","first-page":"251","DOI":"10.3389\/fpubh.2014.00251","article-title":"Ixodes ricinus and its transmitted pathogens in urban and peri-urban areas in Europe: new hazards and relevance for public health","volume":"2","author":"Rizzoli","year":"2014","journal-title":"Front. Public Health"},{"key":"10.1016\/j.pt.2026.09.018_bb0485","doi-asserted-by":"crossref","first-page":"434","DOI":"10.1186\/s13071-019-3700-8","article-title":"Impact of vertebrate communities on Ixodes ricinus-borne disease risk in forest areas","volume":"12","author":"Takumi","year":"2019","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0490","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1186\/s13071-020-3902-0","article-title":"Effect of rodent density on tick and tick-borne pathogen populations: consequences for infectious disease risk","volume":"13","author":"Krawczyk","year":"2020","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0495","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1038\/s41579-020-0400-5","article-title":"Interactions between Borrelia burgdorferi and ticks","volume":"18","author":"Kurokawa","year":"2020","journal-title":"Nat. Rev. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0500","article-title":"The genus Anaplasma: drawing back the curtain on tick-pathogen interactions","volume":"79","author":"O\u2019Neal","year":"2021","journal-title":"Pathog. Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0505","doi-asserted-by":"crossref","first-page":"2370","DOI":"10.1099\/ijsem.0.002848","article-title":"Burkholderia insecticola sp. nov., a gut symbiotic bacterium of the bean bug Riptortus pedestris","volume":"68","author":"Takeshita","year":"2018","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0510","doi-asserted-by":"crossref","first-page":"2057","DOI":"10.1099\/ijsem.0.003431","article-title":"Reclassification of Burkholderia insecticola as Caballeronia insecticola comb. nov. and reliability of conserved signature indels as molecular synapomorphies","volume":"69","author":"Dobritsa","year":"2019","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0515","doi-asserted-by":"crossref","first-page":"17554","DOI":"10.1038\/s41598-017-17309-x","article-title":"A Coxiella mutualist symbiont is essential to the development of Rhipicephalus microplus","volume":"7","author":"Guizzo","year":"2017","journal-title":"Sci. Rep."},{"key":"10.1016\/j.pt.2026.09.018_bb0520","doi-asserted-by":"crossref","first-page":"1896","DOI":"10.1016\/j.cub.2018.04.038","article-title":"Tick-bacteria mutualism depends on B vitamin synthesis pathways","volume":"28","author":"Duron","year":"2018","journal-title":"Curr. Biol."},{"key":"10.1016\/j.pt.2026.09.018_bb0525","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1016\/j.ttbdis.2018.08.008","article-title":"Midichloria mitochondrii, endosymbiont of Ixodes ricinus: evidence for the transmission to the vertebrate host during the tick blood meal","volume":"10","author":"Cafiso","year":"2019","journal-title":"Ticks Tick Borne Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0530","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.00036-12","article-title":"The Black Queen Hypothesis: evolution of dependencies through adaptive gene loss","volume":"3","author":"Morris","year":"2012","journal-title":"mBio"},{"key":"10.1016\/j.pt.2026.09.018_bb0535","doi-asserted-by":"crossref","first-page":"804","DOI":"10.1038\/nature06244","article-title":"The human microbiome project","volume":"449","author":"Turnbaugh","year":"2007","journal-title":"Nature"},{"key":"10.1016\/j.pt.2026.09.018_bb0540","doi-asserted-by":"crossref","first-page":"211","DOI":"10.3389\/fcimb.2020.00211","article-title":"Poor unstable midgut microbiome of hard ticks contrasts with abundant and stable monospecific microbiome in ovaries","volume":"10","author":"Guizzo","year":"2020","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"10.1016\/j.pt.2026.09.018_bb0545","doi-asserted-by":"crossref","DOI":"10.1016\/j.ttbdis.2020.101489","article-title":"Ticks climb the mountains: Ixodid tick infestation and infection by tick-borne pathogens in the Western Alps","volume":"11","author":"Garcia-Vozmediano","year":"2020","journal-title":"Ticks Tick Borne Dis."},{"key":"10.1016\/j.pt.2026.09.018_bb0550","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1186\/1756-3305-5-8","article-title":"Changes in the geographical distribution and abundance of the tick Ixodes ricinus during the past 30 years in Sweden","volume":"5","author":"Jaenson","year":"2012","journal-title":"Parasit. Vectors"},{"key":"10.1016\/j.pt.2026.09.018_bb0555","doi-asserted-by":"crossref","first-page":"380","DOI":"10.1186\/s13071-022-05519-w","article-title":"Tick microbial associations at the crossroad of horizontal and vertical transmission pathways","volume":"15","author":"Krawczyk","year":"2022","journal-title":"Parasit. Vectors"}],"container-title":["Trends in Parasitology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1471492226003028?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1471492226003028?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T12:29:54Z","timestamp":1792585794000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S1471492226003028"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10]]},"references-count":111,"alternative-id":["S1471492226003028"],"URL":"https:\/\/doi.org\/10.1016\/j.pt.2026.09.018","relation":{},"ISSN":["1471-4922"],"issn-type":[{"value":"1471-4922","type":"print"}],"subject":[],"published":{"date-parts":[[2026,10]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Friend or foe? Functional roles of Ixodes ricinus microbes","name":"articletitle","label":"Article Title"},{"value":"Trends in Parasitology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.pt.2026.09.018","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Authors. Published by Elsevier Ltd.","name":"copyright","label":"Copyright"}]}}