Tick species from Africa by migratory birds: a 3-year study in Italy.
Italy
Migratory birds
Ticks
Journal
Experimental & applied acarology
ISSN: 1572-9702
Titre abrégé: Exp Appl Acarol
Pays: Netherlands
ID NLM: 8507436
Informations de publication
Date de publication:
Jan 2021
Jan 2021
Historique:
received:
28
05
2020
accepted:
11
11
2020
pubmed:
19
11
2020
medline:
22
12
2020
entrez:
18
11
2020
Statut:
ppublish
Résumé
The role of resident or migratory birds in dispersal of tick species and tick-borne pathogens is still poorly known in Italy. We report here the results of a 3-year project based on sampling ticks from migratory birds, as well as from the vegetation at three stop-over sites for migrants, namely the islands of Ventotene (Latium), Asinara (Sardinia) and Ustica (Sicily). During the spring seasons from 2017-2019, in total 2681 ticks were collected, 2344 of which were sampled from migratory birds and 337 from the vegetation. Ticks were identified by morphology or by molecular tools when necessary. In total, 16 tick species were identified among which the following were exclusively found on birds: Hyalomma rufipes (43.3%), Hy. truncatum (0.1%), Ixodes frontalis (11.8%), Ix. inopinatus (0.2%), Ix. ricinus (3%), Haemaphysalis punctata (0.08%), Hae. erinacei (0.1%), Amblyomma variegatum (0.08%) and Argas vulgaris 0.1%), whereas five species were exclusively collected from the vegetation: Rhipicephalus bursa (10.5%), Rh. turanicus (5.9%), Rh. sanguineus sensu lato (2%), Rh. pusillus (2.4%), Hae. sulcata (0.08%). Hy. marginatum (10.3%) and Ix. ventalloi (9.3%) were found both on birds and on the vegetation on the island Ustica. It is worth noting that the search for ticks on the vegetation did not detect allochthonous tick species. Although we found several interesting local species and allochthonous ticks like Hy. rufipes, Am. variegatum and Ar. vulgaris on birds, further investigations are needed to better define the possible role of migratory birds in the introduction of ticks and tick-borne diseases in Italy, above all after the evidence of imported ticks positive to Crimean Congo hemorrhagic fever (CCHF) virus in several European countries.
Identifiants
pubmed: 33206312
doi: 10.1007/s10493-020-00573-4
pii: 10.1007/s10493-020-00573-4
doi:
Types de publication
Journal Article
Review
Langues
eng
Sous-ensembles de citation
IM
Pagination
147-164Références
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. https://doi.org/10.1016/j.ttbdis.2018.12.009
doi: 10.1016/j.ttbdis.2018.12.009
pubmed: 30612949
Albanese M, Smiraglia CB, Lavagnino A (1971) Notes on the ticks of Sicily, with descriptions of Hyalomma detritum and Amblyomma variegatum. Riv Parassitol 32(4):273–276
pubmed: 5147924
Apanaskevich DA, Horak IG (2008) The genus Hyalomma Koch, 1844.V. Re-evaluation of the taxonomic rank of taxa comprising the Hy. (Euhyalomma) marginatum Koch complex of species (Acari: Ixodidae) with redescription of all parasitic stages and notes on biology. Int J Acarol 34:13–42
Barker SC, Walker AR (2014) Ticks of Australia. The species that infest domestic animals and humans. Zootaxa 18:1–144
Beati L, Keirans JE (2001) Analysis of the systematic relationships among ticks of the genera Rhipicephalus and Boophilus (acari: Ixodidae) based on mitochondrial 12S ribosomal DNA gene sequences and morphological characters. J Parasitol 87:32–48
pubmed: 11227901
Black WC, Piesman J (1994) Phylogeny of hard- and soft-tick taxa (Acari: Ixodida) based on mitochondrial 16S rDNA sequences. PNAS 21:10034–10038. https://doi.org/10.1073/pnas.91.21.10034
doi: 10.1073/pnas.91.21.10034
Cecere JG, Matricardi C, Frank B, Imperio S, Spina F, Gargallo G, Barboutis C, Boitani L (2010) Nectar exploitation by songbirds at mediterranean stopover sites. Ardeola 57:143–157
Charrier NP, Hermouet A, Hervet C, Agoulon A, Barker SC, Heylen D, Toty C, McCoy KD, Plantard O, Rispe C (2019) A transcriptome-based phylogenetic study of hard ticks (Ixodidae) (2020). Sci Rep. https://doi.org/10.1038/s41598-019-49641-9
doi: 10.1038/s41598-019-49641-9
pubmed: 31506545
pmcid: 6736953
Chastel C, Main AJ, Couatarmanac’h A, Le Lay G, Knudson DL, Quillien MC, Beaucournu JC, (1984) Isolation of Eyach virus (Reoviridae, Colorado tick fever group) from Ixodes ricinus and Ix. ventalloi ticks in France. Arch Virol 82:161–171
pubmed: 6508529
Chitimia-Dobler L, Rieß R, Kahl O, Wölfel S, Dobler G, Nava S, Estrada-Peña A (2018) Ixodes inopinatus—occurring also outside the Mediterranean region. Ticks Tick-Borne Dis 9(2):196–200
pubmed: 28935402
Cicculli V, de Lamballerie X, Charrel R, Falchi A (2019) First molecular detection of Rickettsia africae in a tropical bont tick, Amblyomma variegatum, collected in Corsica, France. Exp Appl Acarol 77:207–214
pubmed: 30656468
Doby JM (1998) Contribution to the knowledge of Ixodes (Trichotoixodes) pari Leach (= I. frontalis (Panzer)) (Acari: Ixodidae), a tick specific to birds. Acarologia 39:315–325
Dabert M, Dabert J, Siuda K (1999) Species validity of the soft-tick Argas polonicus (Acari: Argasidae) based on 16S rDNA sequence analysis. In: Bruin J, van der Geest LPS, Sabelis MW (eds) Ecology and Evolution of the Acari. Series Entomologica, vol 55 Springer, Dordrecht. doi: https://doi.org/10.1007/978-94-017-1343-6_58
de la Fuente J, Estrada-Peña A, Cabezas-Cruz A, Brey R (2015) Flying ticks: anciently evolved associations that constitute a risk of infectious disease spread. Parasites Vectors 8:538. https://doi.org/10.1186/s13071-015-1154-1
doi: 10.1186/s13071-015-1154-1
pubmed: 26467109
pmcid: 4607018
De Liberato C, Frontoso R, Magliano A, Montemaggiori A, Autorino GL, Sala M, Bosworth A, Scicluna MT (2018) Monitoring for the possible introduction of Crimean-Congo haemorrhagic fever virus in Italy based on tick sampling on migratory birds and serological survey of sheep flocks. Prev Vet Med 149:47–52. https://doi.org/10.1016/j.prevetmed.2017.10.014
doi: 10.1016/j.prevetmed.2017.10.014
pubmed: 29290300
Dietrich M, Gomez-Dıaz E, McCoy KD (2011) Worldwide distribution and diversity of seabird ticks: implications for the ecology and epidemiology of tick-borne pathogens. Vector-Borne Zoonotic Dis 11(5):453–470
pubmed: 20874222
Dusbabek F (1976) Argas (Argas) vulgaris Filippova, 1961, A new member of Czechoslovak tick fauna. Folia Parasitologica (Praha) 23:281–283
Estrada-Peña A, Venzal JM (2006) Changes in habitat suitability for the tick Ixodes ricinus (Acari: Ixodidae) in Europe (1900–1999). EcoHealth 3(3):154–162
Estrada-Peña A, Nava S, Petney T (2014) Description of all the stages of Ixodes inopinatus n. sp. (Acari: Ixodidae). Ticks Tick-borne Dis 5(6):734–743
pubmed: 25108790
Estrada-Peña A, Venzal JM, Nava S. Redescription, molecular features, and neotype deposition of Rhipicephalus pusillus Gil Collado and Ixodes ventalloi Gil Collado (Acari, Ixodidae) (2018) Zootaxa 4442(2):262–276. doi: https://doi.org/10.11646/zootaxa.4442.2.4
Estrada-Peña A, Mihalca AD, Petney TN (2017) Ticks of Europe and North Africa—a guide to species identification. Springer International Publishing, New York, p 404
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. Mol Mar Biol Biotechnol 3(5):294–299
pubmed: 7881515
Georgopoulou I, Tsiouris V (2008) The potential role of migratory birds in the transmission of zoonoses. Vet Ital 44:671–677
pubmed: 20411494
Gillingham EL, Hansford KM, Meadows S, Henney J, Wieckowski F, Hernández-Triana LM, Muscat I, Muscat J, Beckerth C, Nikolova NI, Cull B, Medlock JM (2020) Ticks on the Channel Islands and implications for public healtHy. Ticks Tick-borne Dis. https://doi.org/10.1016/j.ttbdis.2020.101405
doi: 10.1016/j.ttbdis.2020.101405
pubmed: 33007668
Gilot B, Beaucournu JC, Chastel C (1997) Fixing on man of Ixodes (Trichotoixodes) frontalis (Panzer, 1795) collected with the flagging method. Parasite 2:197–199
Goymann W, Spina F, Ferri A, Fusani L (2010) Body fat influences departure from stopover sites in migratory birds: evidence from whole island telemetry. Biol Lett 6:478–481. https://doi.org/10.1098/rsbl.2009.1028
doi: 10.1098/rsbl.2009.1028
pubmed: 20164077
pmcid: 2936206
Grandi G, Chitimia-Dobler L, Choklikitumnueya P, Strube C, Springer A, Albihna A, Jaenson TGT, Omazic A (2020) First records of adult Hyalomma marginatum and Hy. rufipes ticks (Acari: Ixodidae) in Sweden. Ticks and Tick-borne Diseases. https://doi.org/10.1016/j.ttbdis.2020.101403
Gray JS, Dautel H, Estrada-Peña A, Kahl O, Lindgren E (2009) Effects of climate change on ticks and tick-borne diseases in Europe. Interdiscip Perspect Infect Dis 2009:1–12
Guglielmone AA, Robbins RG, Apanaskevich DA, Petney TN, Estrada-Peña A, Horak IG (2009) Comments on controversial tick (Acari: Ixodida) species names and species described or resurrected from 2003 to 2008. Exp App Acarol 48:311–327
Guglielmone AA, Robbins RG, Apanaskevich DA, Petney TN, Estrada-Peña A, Horak IG, Renfu S, Barker SC (2010) The Argasidae, Ixodidae and Nuttalliellidae (Acari: Ixodida) of the world: a list of valid species names. Zootaxa 2528:1–28
Hasle G (2013) Transport of ixodid ticks and tick-borne pathogens by migratory birds. Front Cell Infect Microbiol 3:48
pubmed: 24058903
pmcid: 3767891
Hauck D, Springer A, Pachnicke S, Schunack B, Fingerle V, Strube C (2019) Ixodes inopinatus in northern Germany: occurrence and potential vector role for Borrelia spp., Rickettsia spp., and Anaplasma phagocytophilumin comparison with Ixodes ricinus. Parasitol Res 118:3205–3216. https://doi.org/10.1007/s00436-019-06506-4
doi: 10.1007/s00436-019-06506-4
pubmed: 31720842
Heylen D, De Coninck E, Jansen F, Madder M (2014) Differential diagnosis of three common Ixodes spp. ticks infesting songbirds of Western Europe: Ixodes arboricola, I. frontalis and I. ricinus. Tick Tick-borne Dis 5(6):693–700
Heylen D, Fonville M, Docters van Leeuwen A, Sprong H (2016) Co-infections and transmission dynamics in a tick-borne bacterium community exposed to songbirds. Environ Microbiol 18(3):988–996
pubmed: 26627444
Heylen D, Lasters R, Adriaensen F, Fonville M, Sprong H, Matthysen E (2019) Ticks and tick-borne diseases in the city: role of landscape connectivity and green space characteristics in a metropolitan area. Sci Total Environ 670:941–949. https://doi.org/10.1016/j.scitotenv.2019.03.235
doi: 10.1016/j.scitotenv.2019.03.235
pubmed: 30921726
Hildebrandt A, Franke J, Meier F, Sachse S et al (2010) The potential role of migratory birds in transmission cycles of Babesia spp., Anaplasma phagocytophilum and Rickettsia spp. Ticks Tick Borne Dis 1:105–107
pubmed: 21771516
Hornok S, Horváth G (2012) First report of adult Hyalomma marginatum rufipes (vector of Crimean-Congo haemorrhagic fever virus) on cattle under a continental climate in Hungary. Parasit Vectors 5:170. https://doi.org/10.1186/1756-3305-5-170
doi: 10.1186/1756-3305-5-170
pubmed: 22889105
pmcid: 3436687
Hornok S, Flaisz B, Takács N, Kontschán J, Csörgo T, Csipak A, Jaksa BR, Kováts D (2016) Bird ticks in Hungary reflect western, southern, eastern flyway connections and two genetic lineages of Ixodes frontalis and Haemaphysalis concinna. Parasite Vector 9:1–9. https://doi.org/10.1186/s13071-016-1365-0
doi: 10.1186/s13071-016-1365-0
Hubálek Z, Rudolf I (2012) Tick-borne viruses in Europe. Parasitol Res 111:9–36
pubmed: 22526290
Hubálek Z, Sedláček P, Estrada-Peña A, Vojtíšek (2020) First record of Hyalomma rufipes in the Czech Republic, with a review of relevant cases in other parts of Europe. Ticks and Tick-borne Diseases (in press)
Iori A, Di Giulio A, De Felici S (2005) Zecche d’Italia. Cringoli G, Iori A, Rinaldi L, Veneziano V, Genchi C - Mappe parassitologiche: Zecche. Rolando Editore, Napoli, pp 52–163
Jameson LJ, Medlock JM (2011) Tick surveillance in Great Britain. Vector Borne Zoonotic Dis 11(4):403–412. https://doi.org/10.1089/vbz.2010.0079
doi: 10.1089/vbz.2010.0079
pubmed: 20849277
Khasnis AA, Nettleman MD (2005) Global warming and infectious disease. Arch Med Res 36:689–696
pubmed: 16216650
Kolodziejek J, Marinov M, Kiss BJ, Alexe V, Nowotny N (2014) The complete sequence of a West Nile virus lineage 2 strain detected in a Hyalomma marginatum marginatum tick collected from a song thrush (Turdus philomelos) in Eastern Romania in 2013 revealed closest genetic relationship to strain Volgograd 2007. PLoS One. https://doi.org/10.1371/journal.pone.0109905
doi: 10.1371/journal.pone.0109905
pubmed: 25279973
pmcid: 4184904
Lindeborg M, Barboutis C, Ehrenborg C, Fransson T, Jaenson TG, Lindgren PE et al (2012) Migratory birds, ticks, and Crimean-Congo hemorrhagic fever virus. Emerg Infect Dis 8:2095–2097. https://doi.org/10.3201/eid1812.120718
doi: 10.3201/eid1812.120718
Liu X, Yang M, Liu G, Zhao S, Yuan W, Xiao R, Hazihan W, Hornok S, Wang Y (2018) Molecular evidence of Rickettsia raoultii, “Candidatus Rickettsia barbariae” and a novel Babesia genotype in marbled polecats (Vormela peregusna) at the China-Kazakhstan border. Parasit Vectors 11:450. https://doi.org/10.1186/s13071-018-3033-z
doi: 10.1186/s13071-018-3033-z
pubmed: 30075738
pmcid: 6090811
Laakkonen J, Terhivuo J, Huhtamo E, Vapalathi O, Uzcàtegui NY (2009) Memoranda. Soc Fauna Flora Fennica 85:16–19
Latrofa MS, GiannelliA PMF, Pennisi MG, Solano-Gallego L, Brianti E, Parisi A, Wall R, Dantas-Torres F, Otranto D (2017) Ixodes ventalloi: morphological and molecular support for species integrity. Parasitol Res 116:251–258. https://doi.org/10.1007/s00436-016-5286-9
doi: 10.1007/s00436-016-5286-9
pubmed: 27761719
Lwande OW, Venter M, Lutomiah J, Michuki G, Rumberia C, Gakuya F, Obanda V, Tigoi C, Odhiambo C, Nindo F, Symekher S, Sang R (2014) Whole genome phylogenetic investigation of a West Nile virus strain isolated from a tick sampled from livestock in north eastern Kenya. Parasite Vector 7:1–10. https://doi.org/10.1186/s13071-014-0542-2
doi: 10.1186/s13071-014-0542-2
Manilla G (1998) Fauna d’Italia—Ixodida. Edizioni Calderini - Edagricole, Bologna
Mancuso E, Toma L, Polci A, d’Alessio SG, Di Luca M, Orsini M, Di Domenico M, Marcacci M, Mancini G, Spina F, Goffredo M, Monaco F (2019) Crimean-congo hemorrhagic fever virus genome in tick from migratory bird. Italy Emerg Infect Dis 25(7):1419
Mangold AJ, Bargues MD, Mas-Coma S (1998) 18S rRNA gene sequences and phylogenetic relationships of European hard-tick species (Acari: Ixodidae). Parasitol Res 84:31–37
pubmed: 9491423
Marquez FJ (2008) Spotted fever group Rickettsia in ticks from southeastern Spain natural parks. Exp Appl Acarol 45:185–194
pubmed: 18677442
Martyn KP (1988) Provisional atlas of the ticks (Ixodida) of the British Isles. Biological Records Centre—Natural Environment Research Council, Dorset Press, Dorchester, Dorset, UK
Monaco F, Goffredo M, Briguglio P, Pinoni C, Polci A, Iannetti S, Marruchella G, Di Francesco G, Di Gennaro AP, Pais M, Teodori L, Bruno R, Catalani M, Ruiu A, Lelli R, Savini G (2015) The 2011 West Nile disease outbreak in Sardinia region, Italy. Vet Ital 51:5–16. https://doi.org/10.12834/VetIt.260.2386.2
doi: 10.12834/VetIt.260.2386.2
pubmed: 25842208
Monks D, Fisher M, Forbes NA (2006) Ixodes frontalis and avian tick-related syndrome in the United Kingdom. J Small Anim Pract 47(8):451–455
pubmed: 16911113
Norte AC, Lopes de Carvalho I, Ramos JA, Gonçalves M, Gern L, Núncio MS (2012) Diversity and seasonal patterns of ticks parasitizing wild birds in western Portugal. Exp Appl Acarol 58 (3):327-339
pubmed: 22669280
Norte AC, Margos G, Beker NS, (and other Authors), (2019) Host dispersal shapes the population structure of a tick-borne bacterial pathogen. Mol Ecol. https://doi.org/10.1111/mec.15336
doi: 10.1111/mec.15336
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 41:183–192. https://doi.org/10.1016/j.ijpara.2010.08.010
doi: 10.1016/j.ijpara.2010.08.010
pubmed: 20946897
Orkun O, Çakmak A, Nalbantoğlu S, Karaer Z (2019) Molecular detection of a novel Babesia sp. and pathogenic spotted fever group rickettsiae in ticks collected from hedgehogs in Turkey: Haemaphysalis erinacei, a novel candidate vector for the genus Babesia. Inf Gen Evol 69:190–198
Otranto D, Dantas-Torres F, Giannelli A, Latrofa MS, Cascio A, Cazzin S, Ravagnan S, Montarsi F, Zanzani SA, Manfredi MT, Capelli G (2014) Ticks infesting humans in Italy and associated pathogens. Parasit Vectors 7:328
pubmed: 25023709
pmcid: 4223688
Palomar AM, Portillo A, Santibanez P, Mazuelas D, Crespo A, Gutierrez O et al (2013) CCHFV from migratory birds, Morocco. Emerg Infect Dis 19:260–263. https://doi.org/10.3201/eid1902.121193
doi: 10.3201/eid1902.121193
pubmed: 23347801
pmcid: 3559059
Pascucci I, Di Domenico M, Capobianco Dondona G, Di Gennaro A, Polci A, Capobianco Dondona A, Mancuso E, Cammà C, Savini G, Cecere JG, Spina F, Monaco F (2019) Assessing the role of migratory birds in the introduction of ticks and tick-borne pathogens from African countries: an Italian experience. Ticks Tick Borne Dis 10(6):101272. https://doi.org/10.1016/j.ttbdis.2019.101272
doi: 10.1016/j.ttbdis.2019.101272
pubmed: 31481344
Pennisi MG, Persichetti MF, Serrano L, Altet L, Reale S, Gulotta L, Solano-Gallego L (2015) Ticks and associated pathogens collected from cats in Sicily and Calabria (Italy). Parasit Vectors 8:512
pubmed: 26445916
pmcid: 4596469
Petney TN, Pfäffle MP, Skuballa JD (2012) An annotated checklist of the ticks (Acari: Ixodida) of Germany. Syst Appl Acarol 17(2):115–170
Pintore E, Olivieri E, Floriano AM, Sassera D, Sanna N, Garippa G (2021) First detection of Amblyomma variegatum and molecular finding of Rickettsia africae in Sardinia. Ticks and Tick-borne Dis Italy. https://doi.org/10.1016/j.ttbdis.2020.101561
doi: 10.1016/j.ttbdis.2020.101561
Santos Dias JAT, Santos-Reis M (1989) A carraça Ixodes ventalloi Gil Collado, 1936 como principal ectopara-sita de uma população de doninhas (Mustela nivalis Linnaeus, 1766) em Portugal. Garcia de Orta: Série de Zoologia. 14:35–50
Santos-Silva MM, Sousa R, Santos AS, Melo P et al (2006) Ticks parasitizing wild birds in Portugal: detection Rickettsia aeschlimannii, R. helvetica and R. massiliae. Exp Appl Acarol 39:331–338
pubmed: 16897568
Santos-Silva MM, Beati L, Santos AS, De Sousa R, Núncio MS, Melo P, Santos-Reis M, Fonseca C, Formosinho P, Vilela C, Bacellar F (2011) The hard-tick fauna of mainland Portugal (Acari: Ixodidae): an update on geographical distribution and known associations with hosts and pathogens. Exp Appl Acarol 55(1):85–121
pubmed: 21452063
Savini G, Monaco F, Calistri P, Lelli R (2008) Rapid communications phylogenetic analysis of West Nile virus isolated in Italy in 2008. Eurosurveillance 13:260967–260968
Sotelo E, Fernandez-Pinero J, Llorente F, Agüero M, Hoefle U, Blanco JM, Jiménez-Clavero MA (2009) Characterization of West Nile virus isolates from Spain: new insights into the distinct West Nile virus eco-epidemiology in the Western Mediterranean. Virology 395:289–297. https://doi.org/10.1016/j.virol.2009.09.013
doi: 10.1016/j.virol.2009.09.013
pubmed: 19833373
Spina F, Massi A, Montemaggiori A, Baccetti N (1993) Spring migration across central Mediterranean: general results from the “Progetto Piccole Isole.” Die Vogelwarte 37:1–94
Spengler JR, Bente DA, Bray M, Burt F, Hewson R, Korukluoglu G, Mirazimi A, Weber F, Papa A (2018) Second international conference on Crimean-Congo hemorrhagic fever. Antiviral Res 150:137–147
pubmed: 29199036
Tenan S, Spina F (2010) Timing and condition-related effects on recapture probability, mass change and stopover length of spring migrating songbirds on a small Mediterranean island. Ardeola 57:121–132
Toma L, Mancini F, Di Luca M, Cecere JG, Bianchi R, Khoury C, Quarchioni E, Manzia F, Rezza G, Ciervo A (2014) Detection of microbial agents in ticks collected from migratory birds in central Italy. Vector Borne Zoonotic Dis 14(3):199–205. https://doi.org/10.1089/vbz.2013.1458
doi: 10.1089/vbz.2013.1458
pubmed: 24576218
pmcid: 3952585
Torina A, Blanda V, Blanda M, Auteri M, La Russa F, Scimeca S et al (2018) A geographical information system based approach for integrated strategies of tick surveillance and control in the peri-urban natural reserve of Monte Pellegrino (Palermo, Southern Italy). Int J Environ Res Public Health 15:404. https://doi.org/10.3390/ijerph15030404
doi: 10.3390/ijerph15030404
pmcid: 5876949
Walker AR, Bouattour A, Camicas J-L, Estrada-Peña A, Horak IG, Latif AA, Pegram RG, Preston PM (2003) Ticks of domestic animals in Africa: a guide to identification of species. Bioscience Reports, Edinburgh Scotland, UK