{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,27]],"date-time":"2026-06-27T01:19:21Z","timestamp":1782523161405,"version":"3.54.5"},"reference-count":32,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2022,9,1]],"date-time":"2022-09-01T00:00:00Z","timestamp":1661990400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2022,10,25]],"date-time":"2022-10-25T00:00:00Z","timestamp":1666656000000},"content-version":"vor","delay-in-days":54,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100018818","name":"National Research, Development and Innovation Office","doi-asserted-by":"publisher","award":["TKP2020-NKA-01"],"award-info":[{"award-number":["TKP2020-NKA-01"]}],"id":[{"id":"10.13039\/501100018818","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100018818","name":"National Research, Development and Innovation Office","doi-asserted-by":"publisher","award":["TKP2020-NKA-01"],"award-info":[{"award-number":["TKP2020-NKA-01"]}],"id":[{"id":"10.13039\/501100018818","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100018818","name":"National Research, Development and Innovation Office","doi-asserted-by":"publisher","award":["TKP2020-NKA-01"],"award-info":[{"award-number":["TKP2020-NKA-01"]}],"id":[{"id":"10.13039\/501100018818","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100018818","name":"National Research, Development and Innovation Office","doi-asserted-by":"publisher","award":["TKP2020-NKA-01"],"award-info":[{"award-number":["TKP2020-NKA-01"]}],"id":[{"id":"10.13039\/501100018818","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100018818","name":"National Research, Development and Innovation Office","doi-asserted-by":"publisher","award":["TKP2020-NKA-01"],"award-info":[{"award-number":["TKP2020-NKA-01"]}],"id":[{"id":"10.13039\/501100018818","id-type":"DOI","asserted-by":"publisher"}]},{"name":"University of Veterinary Medicine"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Exp Appl Acarol"],"published-print":{"date-parts":[[2022,9]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>This study was initiated to assess the seasonality and to investigate the morphology of questing ixodid ticks in an urban habitat in Central Europe, Hungary. A neglected part of a large cemetery, with sparse tree covering and dense lower vegetation, was sampled monthly from February 2019 to May 2021. All ticks were analyzed morphologically, and selected specimens by amplifying and sequencing two genetic markers. During the study 3818 ticks were collected, including <jats:italic>Ixodes ricinus<\/jats:italic> (n\u2009=\u20092772), <jats:italic>Ixodes frontalis<\/jats:italic> (n\u2009=\u2009350) and <jats:italic>Haemaphysalis concinna<\/jats:italic> (n\u2009=\u2009696). <jats:italic>Ixodes ricinus<\/jats:italic> adults and nymphs showed year-round activity, whereas <jats:italic>H. concinna<\/jats:italic> was not active during winter months and early spring. Most <jats:italic>I. frontalis<\/jats:italic> nymphs were collected in late winter and early spring, whereas the peak activity of larvae was during late autumn. Interestingly, during the spring, the peak activity of <jats:italic>I. ricinus<\/jats:italic> adults and nymphs was later (in May) when preceded by a warm winter in 2020. In contrast, the 2019 and 2021 spring activity peaks occurred in March and April after sharply rising temperatures in February. This shift in the peak activity of <jats:italic>I. ricinus<\/jats:italic> coincided with the initiation of questing activity of <jats:italic>H. concinna<\/jats:italic>. Three notably different morphotypes and four malformed specimens of <jats:italic>I. ricinus<\/jats:italic> were found. However, these were not significantly different in their mitochondrial haplotypes and phylogenetic clustering from typical specimens of this species. On the other hand, <jats:italic>I. frontalis<\/jats:italic> was represented by two remarkably different haplogroups, between which in the nymph stage there were no recognizable morphological differences, suggesting the status of these as cryptic species.<\/jats:p>","DOI":"10.1007\/s10493-022-00756-1","type":"journal-article","created":{"date-parts":[[2022,10,25]],"date-time":"2022-10-25T10:02:46Z","timestamp":1666692166000},"page":"127-138","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":19,"title":["Shift in the seasonality of ixodid ticks after a warm winter in an urban habitat with notes on morphotypes of Ixodes ricinus and data in support of cryptic species within Ixodes frontalis"],"prefix":"10.1007","volume":"88","author":[{"given":"Ciara","family":"Reynolds","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jen\u0151","family":"Kontsch\u00e1n","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"N\u00f3ra","family":"Tak\u00e1cs","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Norbert","family":"Solymosi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Attila D.","family":"S\u00e1ndor","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gerg\u0151","family":"Keve","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"S\u00e1ndor","family":"Hornok","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2022,10,25]]},"reference":[{"key":"756_CR1","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1016\/j.ttbdis.2018.12.009","volume":"10","author":"A Agoulon","year":"2019","unstructured":"Agoulon A, Hoch T, Heylen D, Chalvet-Monfray K, Plantard O (2019) Unravelling the phenology of Ixodes frontalis, a common but understudied tick species in Europe. Ticks Tick Borne Dis 10:505\u2013512","journal-title":"Ticks Tick Borne Dis"},{"key":"756_CR2","doi-asserted-by":"publisher","first-page":"812","DOI":"10.1093\/jmedent\/45.4.812","volume":"45","author":"AN Alekseev","year":"2008","unstructured":"Alekseev AN, Dubinina HV (2008) Enhancement of risk of tick-borne infection: environmental and parasitological aspects of the problem. J Med Entomol 45:812\u2013815","journal-title":"J Med Entomol"},{"key":"756_CR3","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1017\/S0031182000084432","volume":"42","author":"DR Arthur","year":"1953","unstructured":"Arthur DR (1953) The morphology of the British Prostriata with particular reference to Ixodes hexagonus Leach II. Parasitology 42:161\u2013186","journal-title":"Parasitology"},{"key":"756_CR4","doi-asserted-by":"publisher","first-page":"10034","DOI":"10.1073\/pnas.91.21.10034","volume":"91","author":"WC Black","year":"1994","unstructured":"Black WC, Piesman J (1994) Phylogeny of hard and soft-tick taxa (Acari: Ixodida) based on mitochondrial 16s rDNA sequences. Proc Nat Acad Sci USA 91:10034\u201310038","journal-title":"Proc Nat Acad Sci USA"},{"key":"756_CR5","doi-asserted-by":"publisher","first-page":"359","DOI":"10.1007\/s10493-016-0036-3","volume":"69","author":"SV Bugmyrin","year":"2016","unstructured":"Bugmyrin SV, Belova OA, Bespyatova LA, Ieshko EP, Karganova GG (2016) Morphological features of Ixodes persulcatus and I. ricinus hybrids: nymphs and adults. Exp Appl Acarol 69:359\u2013369","journal-title":"Exp Appl Acarol"},{"key":"756_CR6","unstructured":"C3S: Copernicus Climate Change Service (2020) The boreal winter season 19\/20 was by far the warmest winter season ever recorded in Europe. https:\/\/climate.copernicus.eu\/boreal-winter-season-1920-was-far-warmest-winter-season-ever-recorded-europe-0 [accessed 16 October, 2021]"},{"key":"756_CR7","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1186\/s13071-017-2112-x","volume":"10","author":"C Cayol","year":"2017","unstructured":"Cayol C, Koskela E, Mappes T, Siukkola A, Kallio ER (2017) Temporal dynamics of the tick Ixodes ricinus in northern Europe: epidemiological implications. Parasit Vectors 10:166","journal-title":"Parasit Vectors"},{"key":"756_CR8","doi-asserted-by":"publisher","first-page":"379","DOI":"10.1007\/s10493-017-0163-5","volume":"72","author":"L Chitimia-Dobler","year":"2017","unstructured":"Chitimia-Dobler L, Bestehorn M, Br\u00f6ker M, Borde J, Molcanyi T, Andersen NS, Pfeffer M, Dobler G (2017) Morphological anomalies in Ixodes ricinus and Ixodes inopinatus collected from tick-borne encephalitis natural foci in central Europe. Exp Appl Acarol 72:379\u2013397","journal-title":"Exp Appl Acarol"},{"key":"756_CR9","doi-asserted-by":"publisher","first-page":"478","DOI":"10.1186\/s13071-015-1092-y","volume":"8","author":"M Daniel","year":"2015","unstructured":"Daniel M, Mal\u00fd M, Danielov\u00e1 V, K\u0159\u00ed\u017e B, Nuttall P (2015) Abiotic predictors and annual seasonal dynamics of Ixodes ricinus, the major disease vector of central Europe. Parasit Vectors 8:478","journal-title":"Parasit Vectors"},{"key":"756_CR10","doi-asserted-by":"publisher","first-page":"1017","DOI":"10.1016\/j.pt.2018.09.009","volume":"34","author":"F Dantas-Torres","year":"2018","unstructured":"Dantas-Torres F (2018) Species concepts: what about ticks? Trends Parasitol 34:1017\u20131026","journal-title":"Trends Parasitol"},{"key":"756_CR11","unstructured":"Dautel H, Kahl O (1999) Ticks (Acari: Ixodidea) and their medical importance in the urban environment. In: Robinson W H, Rettich F, Rambo G W (Eds). Proceedings of the 3rd International Conference on Urban Pests, pp 73\u201382"},{"key":"756_CR12","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1007\/s10493-019-00375-3","volume":"78","author":"M Drehmann","year":"2019","unstructured":"Drehmann M, Chitimia-Dobler L, Lindau A, Frank A, Mai S, Fachet K, Hauck D, Knoll S, Strube C, L\u00fchken R, Fischer D, Ziegler L, Mackenstedt U (2019) Ixodes frontalis: a neglected but ubiquitous tick species in Germany. Exp Appl Acarol 78:79\u201391","journal-title":"Exp Appl Acarol"},{"key":"756_CR13","doi-asserted-by":"publisher","first-page":"734","DOI":"10.1016\/j.ttbdis.2014.05.003","volume":"5","author":"A Estrada-Pe\u00f1a","year":"2014","unstructured":"Estrada-Pe\u00f1a A, Nava S, Petney T (2014) Description of all the stages of Ixodes inopinatus n. sp. (Acari: Ixodidae). Ticks Tick Borne Dis 5:734\u2013743","journal-title":"Ticks Tick Borne Dis"},{"key":"756_CR14","doi-asserted-by":"publisher","unstructured":"Estrada-Pe\u00f1a A, Mihalca AD, Petney TN (2017). In: Agust\u00edn Estrada-Pe\u00f1a, Andrei Daniel Mihalca, Trevor N. Petney (eds). Ticks of Europe and North Africa. Cham. Springer International Publishing. https:\/\/doi.org\/10.1007\/978-3-319-63760-0","DOI":"10.1007\/978-3-319-63760-0"},{"key":"756_CR15","first-page":"294","volume":"3","author":"O Folmer","year":"1994","unstructured":"Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of mitochondrial cytochrome C oxidase subunit I from diverse metazoan invertebrates. Mel Marine Biol Biot 3:294\u2013299","journal-title":"Mel Marine Biol Biot"},{"key":"756_CR16","doi-asserted-by":"publisher","DOI":"10.1016\/j.ttbdis.2020.101464","volume":"11","author":"D Hauck","year":"2020","unstructured":"Hauck D, Springer A, Chitimia-Dobler L, Strube C (2020) Two-year monitoring of tick abundance and influencing factors in an urban area (city of Hanover, Germany). Ticks Tick Borne Dis 11:101464","journal-title":"Ticks Tick Borne Dis"},{"key":"756_CR17","doi-asserted-by":"publisher","first-page":"366","DOI":"10.1016\/j.vetpar.2009.03.048","volume":"163","author":"S Hornok","year":"2009","unstructured":"Hornok S (2009) Allochronic seasonal peak activities of Dermacentor and Haemaphysalis spp. under continental climate in Hungary. Vet Parasitol 163:366\u2013369","journal-title":"Vet Parasitol"},{"key":"756_CR18","doi-asserted-by":"publisher","first-page":"785","DOI":"10.1016\/j.ttbdis.2014.05.010","volume":"5","author":"S Hornok","year":"2014","unstructured":"Hornok S, Meli ML, G\u00f6nczi E, Hal\u00e1sz E, Tak\u00e1cs N, Farkas R, Hofmann-Lehmann R (2014) Occurrence of ticks and prevalence of Anaplasma phagocytophilum and Borrelia burgdorferi s.l. in three types of urban biotopes: forests, parks and cemeteries. Ticks Tick Borne Dis 5:785\u2013789","journal-title":"Ticks Tick Borne Dis"},{"key":"756_CR19","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1186\/s13071-016-1365-0","volume":"9","author":"S Hornok","year":"2016","unstructured":"Hornok S, Flaisz B, Tak\u00e1cs N, Kontsch\u00e1n J, Cs\u00f6rg\u0151 T, Csipak \u00c1, Jaksa BR, Kov\u00e1ts D (2016) Bird ticks in Hungary reflect western, southern, eastern flyway connections and two genetic lineages of Ixodes frontalis and Haemaphysalis concinna. Parasit Vectors 9:101","journal-title":"Parasit Vectors"},{"key":"756_CR32","doi-asserted-by":"publisher","first-page":"545","DOI":"10.1186\/s13071-017-2424-x","volume":"10","author":"S Hornok","year":"2017","unstructured":"Hornok S, S\u00e1ndor AD, Beck R, Farkas R, Beati L, Kontsch\u00e1n J, Tak\u00e1cs N, F\u00f6ldv\u00e1ri G, Silaghi C, Meyer-Kayser E, Hod\u017ei\u0107 A, Tomanovi\u0107 S, Abdullah S, Wall R, Estrada-Pe\u00f1a A, Duscher GG, Plantard O (2017) Contributions to the phylogeny of Ixodes (Pholeoixodes) canisuga, I. (Ph.) kaiseri, I. (Ph.) hexagonus and a simple pictorial key for the identification of their females. Parasit Vectors 10:545","journal-title":"Parasit Vectors"},{"key":"756_CR20","doi-asserted-by":"publisher","DOI":"10.1016\/j.ttbdis.2021.101865","volume":"13","author":"S Hornok","year":"2022","unstructured":"Hornok S, Daccord J, Tak\u00e1cs N, Kontsch\u00e1n J, Tuska-Szalay B, S\u00e1ndor AD, Szekeres S, Meli ML, Hofmann-Lehmann R (2022) Investigation on haplotypes of ixodid ticks and retrospective finding of Borrelia miyamotoi in bank vole (Myodes glareolus) in Switzerland. Ticks Tick Borne Dis 13:101865","journal-title":"Ticks Tick Borne Dis"},{"key":"756_CR21","first-page":"159","volume":"28","author":"Z Hub\u00e1lek","year":"2003","unstructured":"Hub\u00e1lek Z, Halouzka J, Juricov\u00e1 Z (2003) Host-seeking activity of ixodid ticks in relation to weather variables. J Vector Ecol 28:159\u2013165","journal-title":"J Vector Ecol"},{"issue":"Suppl","key":"756_CR22","doi-asserted-by":"publisher","first-page":"S3","DOI":"10.1017\/S0031182004005967","volume":"129","author":"F Jongejan","year":"2004","unstructured":"Jongejan F, Uilenberg G (2004) The global importance of ticks. Parasitology 129(Suppl):S3-14","journal-title":"Parasitology"},{"key":"756_CR23","doi-asserted-by":"publisher","first-page":"8363","DOI":"10.1029\/2019GL082062","volume":"46","author":"R Lorenz","year":"2019","unstructured":"Lorenz R, Stalhandske Z, Fischer EM (2019) Detection of a climate change signal in extreme heat, heat stress, and cold in Europe from observations. Geophys Res Lett 46:8363\u20138374","journal-title":"Geophys Res Lett"},{"key":"756_CR24","doi-asserted-by":"publisher","first-page":"456","DOI":"10.2307\/1367725","volume":"79","author":"J Lussenhop","year":"1977","unstructured":"Lussenhop J (1977) Urban cemeteries as bird refuges. Condor 79:456\u2013461","journal-title":"Condor"},{"key":"756_CR25","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1186\/1756-3305-7-93","volume":"7","author":"J Lv","year":"2014","unstructured":"Lv J, Wu S, Zhang Y, Chen Y, Feng C, Yuan X, Jia G, Deng J, Wang C, Wang Q, Mei L, Lin X (2014) Assessment of four DNA fragments (COI, 16S rDNA, ITS2, 12S rDNA) for species identification of the Ixodida (Acari: Ixodida). Parasit Vectors 7:93","journal-title":"Parasit Vectors"},{"key":"756_CR26","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1016\/j.ijpara.2010.08.010","volume":"41","author":"R Noureddine","year":"2011","unstructured":"Noureddine R, Chauvin A, Plantard O (2011) Lack of genetic structure among eurasian populations of the tick Ixodes ricinus contrasts with marked divergence from North-African populations. Int J Parasitol 41:183\u2013192","journal-title":"Int J Parasitol"},{"key":"756_CR27","unstructured":"OMSZ (2021) Hungarian Meteorological Service, retrospective data. https:\/\/www.met.hu\/eghajlat\/magyarorszageghajlata\/eghajlati_visszatekinto\/elmult_evszakok_idojarasa\/ [accessed June 23, 2021]"},{"key":"756_CR28","doi-asserted-by":"publisher","DOI":"10.1016\/j.ttbdis.2020.101625","volume":"12","author":"O Plantard","year":"2021","unstructured":"Plantard O, Hoch T, Daveu R, Rispe C, Stachurski F, Bou\u00e9 F, Poux V, Cebe N, Verheyden H, Ren\u00e9-Martellet M, Chalvet-Monfray K, Cafiso A, Olivieri E, Moutailler S, Pollet T, Agoulon A (2021) Where to find questing Ixodes frontalis ticks? under bamboo bushes! Ticks Tick Borne Dis 12:101625","journal-title":"Ticks Tick Borne Dis"},{"issue":"Pt 1","key":"756_CR29","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1017\/S0031182099006083","volume":"121","author":"SE Randolph","year":"2000","unstructured":"Randolph SE, Green RM, Peacey MF, Rogers DJ (2000) Seasonal synchrony: the key to tick-borne encephalitis foci identified by satellite data. Parasitology 121(Pt 1):15\u201323","journal-title":"Parasitology"},{"key":"756_CR30","doi-asserted-by":"publisher","first-page":"979","DOI":"10.1016\/S0020-7519(02)00030-9","volume":"32","author":"SE Randolph","year":"2002","unstructured":"Randolph SE, Green RM, Hoodless AN, Peacey MF (2002) An empirical quantitative framework for the seasonal population dynamics of the tick Ixodes ricinus. Int J Parasitol 32:979\u2013989","journal-title":"Int J Parasitol"},{"key":"756_CR31","unstructured":"UN DESA (2018) United Nations, Department of Economic and Social Affairs, Population Division. World Urbanization Prospects: The 2018 Revision, Online Edition. Available from https:\/\/population.un.org\/wup\/Publications\/Files\/WUP2018-Report.pdf [accessed December 8, 2021]"}],"container-title":["Experimental and Applied Acarology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10493-022-00756-1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s10493-022-00756-1\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10493-022-00756-1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,11,14]],"date-time":"2022-11-14T14:09:28Z","timestamp":1668434968000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s10493-022-00756-1"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,9]]},"references-count":32,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2022,9]]}},"alternative-id":["756"],"URL":"https:\/\/doi.org\/10.1007\/s10493-022-00756-1","relation":{},"ISSN":["0168-8162","1572-9702"],"issn-type":[{"value":"0168-8162","type":"print"},{"value":"1572-9702","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,9]]},"assertion":[{"value":"11 April 2022","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"12 October 2022","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 October 2022","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"No vertebrate animals were caught or restrained for sample collection therefore no ethical permission was needed.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent to participate"}},{"value":"Not applicable.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent to publish"}},{"value":"The authors have no relevant financial or non-financial interests to disclose.","order":5,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}