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Tick ecology: processes and patterns behind the epidemiological risk posed by ixodid ticks as vectors

  • S. E. RANDOLPH (a1)


The population ecology of ticks is fundamental to the spatial and temporal variation in the risk of infection by tick-borne pathogens. Tick population dynamics can only be fully understood by quantifying the rates of the demographic processes, which are influenced by both abiotic (climatic) factors acting on the free-living tick stages and biotic (host) responses to the tick as a parasite. Within the framework of a population model, I review methods and results of attempts to quantify (1) rates of tick development and the probability of diapause, (2) the probability of questing for hosts by unfed ticks, (3) the probability of ticks attaching to a host, and (4) tick mortality rates. Biologically, these processes involve the physiological and behavioural response of ticks to temperature, moisture stress and day length that result in specific patterns of seasonal population dynamics and host relationships. Temperate and tropical patterns will be illustrated with reference mostly to Ixodes ricinus and Rhipicephalus appendiculatus, respectively.


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ALEKSEEV, A. N., DUBININA, H. V., RIJPKEMA, S. G. T. & SCHOULS, L. M. ( 1999). Sexual transmission of Borrelia garinii by male Ixodes persulcatus ticks (Acari, Ixodidae). Experimental and Applied Acarology 23, 165169.
ALLAN, B. F., KEESING, F. & OSTFELD, R. S. ( 2003). Effect of forest fragmentation on Lyme disease risk. Conservation Biology 17, 267272.
ANDERSON, J. F. ( 1991). Epizootiology of Lyme borreliosis. Scandinavian Journal of Infectious Diseases Supplement 77, 2334.
ANDERSON, R. M. & GORDON, D. M. ( 1982). Processes influencing the distribution of parasite numbers within host populations with special emphasis on parasite-induced host mortalities. Parasitology 85, 373398.
ANDERSON, R. M. & MAY, R. M. ( 1978). Regulation and stability of host–parasite population interactions. I. Regulatory processes. Journal of Animal Ecology 47, 219249.
APPERSON, C. S., LEVINE, J. F., EVANS, T. L., BRASWELL, A. & HELLER, J. ( 1993). Relative utilization of reptiles and rodents as hosts by immature Ixodes scapularis (Acari: Ixodidae) in the coastal plain of North Carolina, USA. Experimental and Applied Acarology 17, 719731.
BEGON, M., HARPER, J. L. & TOWNSEND, C. R. ( 1996). Ecology (3rd edn.). Oxford, Blackwell Scientific Publications.
BELOZEROV, V. N. ( 1967). [Larval diapause in the tick Ixodes ricinus L and its dependence on external conditions. IV. Interactions between exogenous and endogenous factors in the regulation of the larval diapause.] (In Russian). Entomological Review 46, 447451.
BELOZEROV, V. N. ( 1970). [Nymphal diapause in the tick Ixodes ricinus L. (Ixodidae). III. Photoperiodic reaction in unfed nymphs.] (In Russian.) Parazitologiya 4, 139145.
BELOZEROV, V. N. ( 1982). Diapause and biological rhythms in ticks. In Physiology of Ticks ( ed. Obenchain, F. D. & Galun, R.), pp. 469500. Oxford, Pergamon Press.
BELOZEROV, V. N. ( 1998 a). Role of two-step photoperiodic reaction in the control of development and diapause in the nymphs of Ixodes persulcatus. Russian Journal of Zoology 2, 414418.
BELOZEROV, V. N. ( 1998 b). Dynamics of gas exchange during development of Ixodid ticks: III. Dynamics of gas exchange in nymphs of the Ixodid ticks Hyalomma anatolicum Koch (Acari, Ixodidae) during active development and developmental diapause. Entomological Review 78, 197205.
BELOZEROV, V. N. & NAUMOV, R. L. ( 2001). Photoperiodic control of nymphal diapause in the North American tick, Ixodes (Ixodes) scapularis Say (Acari: Ixodidae). In IVth European Workshop of Invertebrate Ecophysiology ( ed. Kipyatkov, V. E.), p. 69. St Petersburg, Russia.
BERKVENS, D. L., PEGRAM, R. G. & BRANDT, J. R. A. ( 1995). A study of the diapausing behaviour of Rhipicephalus appendiculatus and R. zambesiensis under quasi-natural conditions in Zambia. Medical and Veterinary Entomology 9, 307315.
BERTRAND, M. R. & WILSON, M. L. ( 1997). Microhabitat-independent regional differences in survival of unfed Ixodes scapularis nymphs (Acari: Ixodidae) in Connecticut. Journal of Medical Entomology 34, 167172.
BRANAGAN, D. ( 1973). Observations on the development and survival of the Ixodid tick Rhipicephalus appendiculatus Neumann, 1901 under quasi-natural conditions in Kenya. Tropical Animal Health and Production 5, 153165.
CAMPBELL, J. A. ( 1948). The life history and development of the sheep tick Ixodes ricinus Linaeus in Scotland, under natural and controlled conditions. PhD thesis, University of Edinburgh.
CERNY, M., DANIEL, M. & ROSICKY, B. ( 1974). Some features of the developmental cycle of the tick Ixodes ricinus (L.) (Acarina: Ixodidae). Folia Parasitologica 21, 8587.
CHIERA, J. W., NEWSON, R. M. & CUNNINGHAM, M. P. ( 1985). Cumulative effects of host resistance on Rhipicephalus appendiculatus Neumann (Acarina: Ixodidae) in the laboratory. Parasitology 90, 401408.
CHMELA, J. ( 1969). On the developmental cycle of the common tick (Ixodes ricinus L.) in the north-Moravian natural focus of tick-borne encephalitis. Folia Parasitologica 16, 313319.
CLARK, D. D. ( 1995). Lower temperature limits for activity of several ixodid ticks (Acari: Ixodidae): effects of body size and rate of temperature change. Journal of Medical Entomology 32, 449452.
clayton, d. h. & moore, j. (ed.) ( 1997). Host–Parasite Evolution. Oxford, Oxford University Press.
CRAINE, N. G., RANDOLPH, S. E. & NUTTALL, P. A. ( 1995). Seasonal variation in the role of grey squirrels as hosts of Ixodes ricinus, the tick vector of the Lyme disease spirochaete, in a British woodland. Folia Parasitologica 42, 7380.
CROFTON, H. D. ( 1971 a). A quantitative approach to parasitism. Parasitology 62, 179193.
CROFTON, H. D. ( 1971 b). A model of host–parasite relationships. Parasitology 63, 343364.
CUMMING, G. S. ( 1996). The evolutionary ecology of African ticks. DPhil thesis, University of Oxford.
CUMMING, G. S. ( 1998). Host preference in African ticks (Acari: Ixodidae): a quantitative data set. Bulletin of Entomological Research 88, 379406.
CUMMING, G. S. ( 1999). Host distributions do not limit the species ranges of most African ticks (Acari: Ixodida). Bulletin of Entomological Research 89, 303327.
CUMMING, G. S. ( 2002). Comparing climate and vegetation as limiting factors for species ranges of African ticks. Ecology 83, 255268.
DANIEL, M., CERNY, V., DUSBABEK, F., HONZAKOVA, E. & OLEJNICEK, J. ( 1976). Influence of microclimate on the life cycle of the common tick Ixodes ricinus (L.) in thermophilic oak forest. Folia Parasitologica 23, 327342.
DANIEL, M., CERNY, V., DUSBABEK, F., HONZAKOVA, E. & OLEJNICEK, J. ( 1977). Influence of microclimate on the life cycle of the common tick Ixodes ricinus (L.) in an open area in comparison with forest habitats. Folia Parasitologica 24, 149160.
DANIEL, M. & KOLAR, J. ( 1990). Using satellite data to forecast the occurrence of the common tick Ixodes ricinus (L.). Journal of Hygiene, Epidemiology, Microbiology and Immunology 45, 243252.
DANIEL, M., KOLAR, J., ZEMAN, P., PAVELKA, K. & SADLO, J. ( 1998). Predictive map of Ixodes ricinus high-incidence habitats and a tick-borne encephalitis risk assessment using satellite data. Experimental and Applied Acarology 22, 417433.
DANIELOVÁ, V. ( 2002). Natural foci of tick-borne encephalitis and prerequisites for their existence. International Journal of Medical Microbiology 291, S183S186.
DANIELOVÁ, V. & HOLUBOVÁ, J. ( 1991). Transovarial transmission rate of tick-borne encephalitis virus in Ixodes ricinus ticks. In Modern Acarology ( ed. Dusbabek, F. & Bukva, V.), pp. 710. The Hague, Academia, Prague and SPB Academic Publishing.
DANIELOVÁ, V., HOLUBOVA, J., PEJCOCH, M. & DANIEL, M. ( 2002). Potential significance of transovarial transmission in the circulation of tick-borne encephalitis virus. Folia Parasitologica 49, 323325.
DANIELS, T. J., FALCO, R. C., CURRAN, K. L. & FISH, D. ( 1996). Timing of Ixodes scapularis (Acari: Ixodidae) oviposition and larval activity in southern New York. Journal of Medical Entomology 33, 140147.
DANIELS, T. J., FALCO, R. C. & FISH, D. ( 2000). Estimating population size and drag sampling efficiency for the blacklegged tick (Acari: Ixodidae). Journal of Medical Entomology 37, 357363.
DANIELS, T. J. & FISH, D. ( 1995). Effect of deer exclusion on the abundance of immature Ixodes scapularis (Acari: Ixodidae) parasitizing small and medium-sized mammals. Journal of Medical Entomology 32, 511.
DISTER, S. W., FISH, D., BROS, S. M., FRANK, D. H. & WOOD, B. L. ( 1997). Landscape characterization of peridomestic risk for Lyme disease using satellite imagery. American Journal of Tropical Medicine and Hygiene 57, 687692.
DIZIJ, A. & KURTENBACH, K. ( 1995). Clethrionomys glareolus, but not Apodemus flavicollis, acquires resistance to Ixodes ricinus L., the main European vector of Borrelia burgdorferi. Parasite Immunology 17, 177183.
EISEN, L., EISEN, R. J. & LANE, R. S. ( 2002). Seasonal activity patterns of Ixodes pacificus nymphs in relation to climatic conditions. Medical and Veterinary Entomology 16, 235244.
ESTRADA-PEÑA, A. ( 1998). Geostatistics and remote sensing as predictive tools of tick distribution: a cokriging system to estimate Ixodes scapularis (Acari: Ixodidae) habitat suitability in the United States and Canada from Advanced Very High Resolution Radiometer satellite imagery. Journal of Medical Entomology 35, 989995.
ESTRADA-PEÑA, A. ( 1999). Geostatistics as predictive tools to estimate Ixodes ricinus (Acari: Ixodidae) habitat suitability in the western Paleartic from AVHRR satellite imagery. Experimental and Applied Acarology 23, 337349.
FALCO, R. C., DANIELS, T. J. & FISH, D. ( 1995). Increase in abundance of immature Ixodes scapularis (Acari: Ixodidae) in an emergent Lyme disease endemic area. Journal of Medical Entomology 32, 522526.
FIELDEN, L. J. & LIGHTON, J. R. B. ( 1996). Effects of water stress and relative humidity on ventilation in the tick Dermacentor andersoni (Acari: Ixodidae). Physiological Zoology 69, 599617.
FILIPPOVA, N. A. (ed.) ( 1985). [Taiga Tick Ixodes persulcatus Schulze (Acarina, Ixodidae).] (In Russian). Leningrad: Nauka.
FIVAZ, B. H. & NORVAL, R. A. I. ( 1990). Immunity of the ox to the brown ear tick Rhipicephalus appendiculatus. Experimental and Applied Acarology 8, 5163.
FOURIE, L. J., BELOZEROV, V. N. & NEEDHAM, G. R. ( 2001). Ixodes rubicundus nymphs are short-day diapause-induced ticks with thermolabile sensitivity and desiccation resistance. Medical and Veterinary Entomology 15, 335341.
GAEDE, K. & KNÜLLE, W. ( 1997). On the mechanism of water vapour sorption from unsaturated atmospheres by ticks. Journal of Experimental Biology 200, 14911498.
GIGON, F. ( 1985). Biologie d'Ixodes ricinus L. sur le Plateau Suisse – une contribution a l'écologie de ce vecteur. MS thesis, Université de Neuchâtel, Switzerland.
GILOT, B., BONNEFILLE, M., DEGEILH, B., BEAUCOURNU, J.-C., PICHOT, J. & GUIGUEN, C. ( 1994). La colonisation des massifs forestiers par Ixodes ricinus (Linne, 1758) en France: utilisation du chevreuil, Capreolus capreolus (L. 1758) comme marqueur biologique. Parasite 1, 8186.
GLASS, G. E., SCHWARZ, B. S., MORGAN III, J. M., JOHNSON, D. T., NOY, P. M. & ISRAEL, E. ( 1995). Environmental risk factor for Lyme disease identified with geographic information systems. American Journal of Public Health 85, 944948.
GRAY, J. S. ( 1981). The fecundity of Ixodes ricinus (L.) (Acarina: Ixodidae) and the mortality of its developmental stages under field conditions. Bulletin of Entomological Research 71, 533542.
GRAY, J. S. ( 1982). The development and questing activity of Ixodes ricinus (L.) (Acari: Ixodidae) under field conditions in Ireland. Bulletin of Entomological Research 72, 263270.
GRAY, J. S. ( 1985). Studies on the larval activity of the tick Ixodes ricinus L. in Co. Wicklow, Ireland. Experimental and Applied Acarology 1, 307316.
GRAY, J. S., KAHL, O., JANETZKI, C. & STEIN, J. ( 1992). Studies on the ecology of Lyme disease in a deer forest in County Galway, Ireland. Journal of Medical Entomology 29, 915920.
GRAY, J. S., VON STEDINGK, L. V. & GURTELSCHMID, M. E. A. ( 2002). Transmission studies of Babesia microti in Ixodes ricinus ticks and gerbils. Journal of Clinical Microbiology 40, 12591263.
GUERRA, M., WALKER, E., JONES, C. J., PASKEWITZ, S., CORTINAS, M. R., SANCIL, A., BECK, L., BOBO, M. & KITRON, U. ( 2002). Predicting the risk of Lyme disease: habitat suitability for Ixodes scapularis in the north central United States. Emerging Infectious Diseases 8, 289297.
HASIBEDER, G. & DYE, C. M. ( 1988). Population dynamics of mosquito-borne disease: persistence in a completely heterogeneous environment. Theoretical Population Biology 33, 3153.
HOODLESS, A. N., KURTENBACH, K., NUTTALL, P. A. & RANDOLPH, S. E. ( 2002). The impact of ticks on pheasant territoriality. Oikos 96, 245250.
HOOGSTRAAL, H. ( 1981). Changing patterns of tick-borne diseases in modern society. Annual Review of Entomology 26, 7599.
HUDSON, P. J., RIZZOLI, A., ROSA, R., CHEMINI, C., JONES, L. D. & GOULD, E. A. ( 2001). Tick-borne encephalitis virus in northern Italy: molecular analysis, relationships with density and seasonal dynamics of Ixodes ricinus. Medical and Veterinary Entomology 15, 304313.
HUGH-JONES, M., BARRE, N., NELSON, G., WEHNES, K., WARNER, J., GARVIN, J. & GARRIS, G. ( 1992). Landsat-TM identification of Amblyomma variegatun (Acari: Ixodidae) habitats in Guadeloupe. Remote Sensing of the Environment 40, 4355.
HUGHES, V. L. & RANDOLPH, S. E. ( 2001 a). Testosterone depresses innate and acquired resistance to ticks in natural rodent hosts: a force for aggregated distributions of parasites. Journal of Parasitology 87, 4954.
HUGHES, V. L. & RANDOLPH, S. E. ( 2001 b). Testosterone increases the transmission potential of tick-borne parasites. Parasitology 123, 365371.
HUTCHINSON, M. F., NIX, H. A., McMAHON, J. P. & ORD, K. D. ( 1995). Africa. A topographic and climate database. In Centre for Resources and Environmental Studies, Canberra, The Australian National University.
JENSEN, P. M. & FRANDSEN, F. ( 2000). Temporal risk assessment for Lyme borreliosis in Denmark. Scandinavian Journal of Infectious Diseases 35, 539544.
JENSEN, P. M., HANSEN, H. & FRANDSEN, F. ( 2000). Spatial risk assessment for Lyme borreliosis in Denmark. Scandinavian Journal of Infectious Diseases 35, 545550.
JONGEJAN, F., PEGRAM, R. G., ZIVKOVIC, D., HENSEN, E. J., MWASE, E. T., THIELEMANS, M. J. C., COSSE, A., NIEWOLD, T. A., SAID, A. E. & UILENBERG, G. ( 1989). Monitoring of naturally acquired and artificially induced immunity to Amblyomma variegatum and Rhipicephalus appendiculatus ticks under field and laboratory conditions. Experimental and Applied Acarology 7, 181189.
KAISER, M. N., SUTHERST, R. W. & BOURNE, A. S. ( 1982). Relationship between ticks and zebu cattle in southern Uganda. Tropical Animal Health Production 14, 6374.
KAISER, M. N., SUTHERST, R. W. & BOURNE, A. S. ( 1991). Tick (Acarina: Ixodidae) infestations on zebu cattle in northern Uganda. Bulletin of Entomological Research 81, 257262.
KAISER, M. N., SUTHERST, R. W., BOURNE, A. S., GORISSEN, L. & FLOYD, R. B. ( 1988). Population dynamics of ticks on Ankole cattle in five ecological zones in Burundi and strategies for their control. Preventive Veterinary Medicine 6, 199222.
KEMP, D. H. ( 1968). Physiological studies on hard ticks Ixodidae. PhD thesis, University of Edinburgh.
KITRON, U., JONES, C. J., BOUSEMAN, J. K., NELSON, J. A. & BAUMGARTNER, D. L. ( 1992). Spatial analysis of the distribution of Ixodes dammini (Acari: Ixodidae) on white-tailed deer in Ogle county, Illinois. Journal of Medical Entomology 29, 259266.
KITRON, U. & KAZMIERCZAK, J. J. ( 1997). Spatial analysis of the distribution of Lyme disease in Wisconsin. American Journal of Epidemiology 145, 558566.
KLOMPEN, J. S. H., BLACK, W. C., KEIRANS, J. E. & OLIVER, J. H. ( 1996). Evolution of ticks. Annual Review of Entomology 41, 141161.
KNULLE, W. & RUDOLPH, D. ( 1982). Humidity relationships and water balance of ticks. In Physiology of Ticks ( ed. Obenchain, F. D. & Galun, R.), pp. 4370. Oxford, Pergamon Press.
KORENBERG, E. I. ( 2000). Seasonal population dynamics of Ixodes ticks and tick-borne encephalitis virus. Experimental and Applied Acarology 24, 665681.
KORENBERG, E. I. & KOVALEVSKII, Y. V. ( 1994). A model for relationships among the tick-borne encephalitis virus, its main vectors, and hosts. Advances in Disease Vector Research 10, 6592.
KURTENBACH, K., PEACEY, M. F., RIJPKEMA, S. G. T., HOODLESS, A. N., NUTTALL, P. A. & RANDOLPH, S. E. ( 1998). Differential transmission of the genospecies of Borrelia burgdorferi sensu lato by game birds and small rodents in England. Applied and Environmental Microbiology 64, 11691174.
LABUDA, M., AUSTYN, J. M., ZUFFOVA, E., KOZUCH, O., FUCHSBERGER, N., LYSY, J. & NUTTALL, P. A. ( 1996). Importance of localized skin infection in tick-borne encephalitis virus transmission. Virology 219, 357366.
LABUDA, M., JONES, L. D., WILLIAMS, T., DANIELOVA, V. & NUTTALL, P. A. ( 1993 a). Efficient transmission of tick-borne encephalitis virus between cofeeding ticks. Journal of Medical Entomology 30, 295299.
LABUDA, M., KOZUCH, O., ZUFFOVA, E., ELECKOVA, E., HAILS, R. S. & NUTTALL, P. A. ( 1997). Tick-borne encephalitis virus transmission between ticks cofeeding on specific immune natural rodent hosts. Virology 235, 138143.
LABUDA, M., NUTTALL, P. A., KOZUCH, O., ELECKOVA, E., WILLIAMS, T., ZUFFOVA, E. & SABO, A. ( 1993 b). Non-viraemic transmission of tick-borne encephalitis virus: a mechanism for arbovirus survival in nature. Experientia 49, 802805.
LANE, R. S. & QUISTAD, G. B. ( 1998). Borreliacidal factor in the blood of the Western Fence Lizard (Sceloporus occidentalis). Journal of Parasitology 84, 2934.
LEES, A. D. & MILNE, A. ( 1951). The seasonal and diurnal activities of individual sheep ticks (Ixodes ricinus). Parasitology 41, 180209.
LIGHTON, J. R. B. & FIELDEN, L. J. ( 1995). Mass scaling of standard metabolism in ticks – a valid case of low metabolic rates in sit-and-wait strategists. Physiological Zoology 68, 4362.
LOGIUDICE, K., OSTFELD, R. S., SCHMIDT, K. A. & KEESING, F. ( 2003). The ecology of infectious disease: effects of host diversity and community composition on Lyme disease risk. Proceedings of the National Academy of Sciences, USA 100, 567571.
LOYE, J. E. & LANE, R. S. ( 1988). Questing behavior of Ixodes pacificus (Acari: Ixodidae) in relation to meteorological and seasonal factors. Journal of Medical Entomology 25, 391398.
MACLEOD, J. ( 1935). Ixodes ricinus in relation to its physical environment. II. The factors governing survival and activity. Parasitology 27, 123144.
MACLEOD, J. ( 1936). Ixodes ricinus in relation to its physical environment. IV. An analysis of the ecological complexes controlling distribution and activities. Parasitology 28, 295319.
MACLEOD, J. ( 1970). Tick infestation patterns in the Southern Province of Zambia. Bulletin of Entomological Research 60, 253274.
MADDER, M., SPEYBROECK, N., BRANDT, J. & BERKVENS, D. ( 1999). Diapause induction in adults of three Rhipicephalus appendiculatus stocks. Experimental and Applied Acarology 23, 961968.
MATUSCHKA, F.-R., FISCHER, P., HEILER, M., RICHTER, D. & SPIELMAN, A. ( 1992). Capacity of European animals as reservoir hosts for the Lyme disease spirochete. Journal of Infectious Diseases 165, 479483.
MATUSCHKA, F.-R., SCHINKEL, T. W., KLUG, B., SPIELMAN, A. & RICHTER, D. ( 2000). Relative importance of European rabbits for Lyme disease spirochaetes. Parasitology 121, 297302.
MILNE, A. ( 1949). The ecology of the sheep tick, Ixodes ricinus L. Host relationships of the tick. Part 2. Observations on hill and moorland grazings in northern England. Parasitology 39, 173194.
MOORE, S. L. & WILSON, K. ( 2002). Parasites as a viability cost of sexual selection in natural populations of mammals. Science 297, 20152018.
MORAN, M. C., NIGARURA, G. & PEGRAM, R. G. ( 1996). An assessment of host resistance to ticks on cross-bred cattle in Burundi. Medical and Veterinary Entomology 10, 1218.
NEEDHAM, G. R. & TEEL, P. D. ( 1991). Off-host physiological ecology of ixodid ticks. Annual Review of Entomology 36, 659681.
NEWSON, R. M. ( 1978). The life cycle of Rhipicephalus appendiculatus on the Kenyan coast. In Tick-borne Diseases and their Vectors ( ed. Wilde, J. K. H.), pp. 4650. Edinburgh, CTVM, University of Edinburgh.
NICHOLSON, M. C. & MATHER, T. N. ( 1996). Methods for evaluating Lyme disease risks using geographical information systems and geospatial analysis. Journal of Medical Entomology 33, 711720.
NORVAL, R. A. I., LAWRENCE, J. A., YOUNG, A. S., PERRY, B. D., DOLAN, T. T. & SCOTT, J. ( 1991). Theileria parva: influence of vector, parasite and host relationships on the epidemiology of theileriosis in southern Africa. Parasitology 102, 347356.
NUTTALL, P. A. & LABUDA, M. ( 1994). Tick-borne encephalitides. In Ecological Dynamics of Tick-borne Zoonoses ( ed. Sonenshine, D. E. & Mather, T. N.), pp. 351391. New York, Oxford, Oxford University Press.
OCHANDA, H., YOUNG, A. S., WELLS, C., MEDLEY, G. F. & PERRY, B. D. ( 1996). Comparison of the transmission of Theileria parva between different instars of Rhipicephalus appendiculatus. Parasitology 113, 243253.
OGDEN, N. H., HAILS, R. S. & NUTTALL, P. A. ( 1998). Interstadial variation in the attachment sites of Ixodes ricinus ticks on sheep. Experimental and Applied Acarology 22, 227232.
OGDEN, N. H., RANDOLPH, S. E. & NUTTALL, P. A. ( 1997). Natural Lyme disease cycle maintained via sheep by co-feeding ticks. Parasitology 115, 591599.
PACALA, S. W. & DOBSON, A. P. ( 1988). The relation between the number of parasites/host and host age: population dynamics causes and maximum likelihood estimation. Parasitology 96, 197210.
PADGETT, K. A. & LANE, R. S. ( 2001). Life cycle of Ixodes pacificus (Acari: Ixodidae): timing of development processes under field and laboratory conditions. Journal of Medical Entomology 38, 684693.
PEGRAM, R. G. & BANDA, D. S. ( 1990). Ecology and phenology of cattle ticks in Zambia: development and survival of free-living stages. Experimental and Applied Acarology 8, 291293.
PERRET, J.-L., GUERIN, P. M., DIEHL, P.-A., VLIMANT, M. & GERN, L. ( 2003). Darkness favours mobility and saturation deficit limits questing duration in Ixodes ricinus, the tick vector of Lyme disease in Europe. Journal of Experimental Biology 206, 18091815.
PERRET, J.-L., GUIGOZ, E., RAIS, O. & GERN, L. ( 2000). Influence of saturation deficit and temperature on Ixodes ricinus tick questing activity in a Lyme borrelosis-endemic area (Switzerland). Parasitology Research 86, 554557.
PETERS, A. ( 2000). Testosterone treatment is immunosuppressive in superb fairy-wrens, yet free-living males with high testosterone are more immunocompetent. Proceedings of the Royal Society of London, B 267, 883889.
PIESMAN, J. ( 2002). Ecology of Borrelia burgdorferi sensu lato in North America. In Lyme Borreliosis Biology, Epidemiology and Control ( ed. Gray, J. S., Kahl, O., Lane, R. S. & Stanek, G.), pp. 223250. Wallingford, CABI.
RAND, P. W., LUBELCZYK, C., LAVIGNE, G. R., ELIAS, S., HOLMAN, M. S., LACOMBE, E. H. & SMITH, R. P. J. ( 2003). Deer density and the abundance of Ixodes scapularis (Acari: Ixodidae). Journal of Medical Entomology 40, 179184.
RANDOLPH, S. E. ( 1975). Patterns of distribution of the tick Ixodes trianguliceps Birula on its hosts. Journal of Animal Ecology 44, 451474.
RANDOLPH, S. E. ( 1977). Changing spatial relationships in a population of Apodemus sylvaticus with the onset of breeding. Journal of Animal Ecology 46, 653676.
RANDOLPH, S. E. ( 1979). Population regulation in ticks: the role of acquired resistance in natural and unnatural hosts. Parasitology 79, 141156.
RANDOLPH, S. E. ( 1993). Climate, satellite imagery and the seasonal abundance of the tick Rhipicephalus appendiculatus in southern Africa: a new perspective. Medical and Veterinary Entomologyy 7, 243258.
RANDOLPH, S. E. ( 1994 a). Population dynamics and density-dependent seasonal mortality indices of the tick Rhipicephalus appendiculatus in east and southern Africa. Medical and Veterinary Entomology 8, 351368.
RANDOLPH, S. E. ( 1994 b). Density-dependent acquired resistance to ticks in natural hosts, independent of concurrent infection with Babesia microti. Parasitology 108, 413419.
RANDOLPH, S. E. ( 1995). Quantifying parameters in the transmission of Babesia microti by the tick Ixodes trianguliceps amongst voles (Clethrionomys glareolus). Parasitology 110, 287295.
RANDOLPH, S. E. ( 1997). Abiotic and biotic determinants of the seasonal dynamics of the tick Rhipicephalus appendiculatus in South Africa. Medical and Veterinary Entomology 11, 2537.
RANDOLPH, S. E. ( 1998). Ticks are not insects: consequences of contrasting vector biology for transmission potential. Parasitology Today 14, 186192.
RANDOLPH, S. E. ( 2000). Ticks and tick-borne disease systems in space and from space. Advances in Parasitology 47, 217243.
RANDOLPH, S. E. ( 2001). The shifting landscape of tick-borne zoonoses: tick-borne encephalitis and Lyme borreliosis in Europe. Philosophical Transactions of the Royal Society, B 356, 10451056.
RANDOLPH, S. E. & STEELE, G. M. ( 1985). An experimental evaluation of conventional control measures against the sheep tick Ixodes ricinus (L) (Acari: Ixodidae). II. The dynamics of the tick–host interaction. Bulletin of Entomological Research 75, 501518.
RANDOLPH, S. E., GREEN, R. M., PEACEY, M. F. & ROGERS, D. J. ( 2000). Seasonal synchrony: the key to tick-borne encephalitis foci identified by satellite data. Parasitology 121, 1523.
RANDOLPH, S. E., GREEN, R. M., HOODLESS, A. N. & PEACEY, M. F. ( 2002). An empirical quantitative framework for the seasonal population dynamics of the tick Ixodes ricinus. Internatioal Journal for Parasitology 32, 979989.
RANDOLPH, S. E., MIKLISOVÁ, D., LYSY, J., ROGERS, D. J. & LABUDA, M. ( 1999). Incidence from coincidence: patterns of tick infestations on rodents facilitate transmission of tick-borne encephalitis virus. Parasitology 118, 177186.
RANDOLPH, S. E. & ROGERS, D. J. ( 1997). A generic population model for the African tick Rhipicephalus appendiculatus. Parasitology 115, 265279.
RANDOLPH, S. E. & STOREY, K. ( 1999). Impact of microclimate on immature tick-rodent interactions (Acari: Ixodidae): implications for parasite transmission. Journal of Medical Entomology 36, 741748.
RECHAV, Y. ( 1979). Migration and dispersal patterns of three African ticks (Acari: Ixodidae) under field conditions. Journal of Medical Entomology 16, 150163.
RECHAV, Y. ( 1981). Ecological factors affecting the seasonal activity of the brown ear tick Rhipicephalus appendiculatus. In Tick Biology and Control ( ed. Whitehead, G. B. & Gibson, J. D.), pp. 187191. South Africa, Grahamstown University.
RECHAV, Y. ( 1982). Dynamics of tick populations (Acari: Ixodidae) in the Eastern Cape Province of South Africa. Journal of Medical Entomology 19, 679700.
RECHAV, Y. ( 1992). Naturally acquired resistance to ticks – a global view. Insect Science and its Application 13, 495504.
ROGERS, D. J. & RANDOLPH, S. E. ( 1993). Distribution of tsetse and ticks in Africa: past, present and future. Parasitology Today 9, 266271.
ROWSEWITT, C. N. ( 1986). Seasonal variation in activity rhythms of male voles: mediation by gonadal hormones. Physiology and Behaviour 37, 797803.
SHAW, D. J., GRENFELL, B. T. & DOBSON, A. P. ( 1998). Patterns of macroparasite aggregation in wildlife host populations. Parasitology 117, 597610.
SHORT, N. J., FLOYD, R. B., NORVAL, R. A. I. & SUTHERST, R. W. ( 1989). Survival and behaviour of unfed stages of the ticks Rhipicephalus appendiculatus, Boophilus decoloratus and B. microplus under field conditions in Zimbabwe. Experimental and Applied Acarology 6, 215236.
SHORT, N. J. & NORVAL, R. A. I. ( 1981). The seasonal activity of Rhipicephalus appendiculatus Neumann 1901 (Acarina: Ixodidae) in the high veld of Zimbabwe Rhodesia. Journal of Parasitology 67, 7784.
SONENSHINE, D. E. ( 1991). Biology of Ticks, vol. 1. Oxford, Oxford University Press.
SPIELMAN, A., WILSON, M. L., LEVINE, J. F. & PIESMAN, J. ( 1985). Ecology of Ixodes dammini-borne human babesiosis and Lyme disease. Annual Review of Entomology 30, 439460.
STEELE, G. M. & RANDOLPH, S. E. ( 1985). An experimental evaluation of conventional control measures against the sheep tick Ixodes ricinus (L) (Acari: Ixodidae). I. A unimodal seasonal activity pattern. Bulletin of Entomological Research 75, 489499.
STUBBS, M. ( 1977). Density dependence in the life-cycles of animals and its importance in K- and r-strategies. Journal of Animal Ecology 46, 677688.
SUTHERST, R. W., UTECH, K. B. W., KERR, J. D. & WHARTON, R. H. ( 1979). Density-dependent mortality of the tick Boophilus microplus, on cattle – further observations. Journal of Applied Ecology 16, 397403.
TALLEKLINT, L. & JAENSON, T. G. T. ( 1994). Transmission of Borrelia burgdorferi s.l. from mammal reservoirs to the primary vector of Lyme borreliosis, Ixodes ricinus (Acari: Ixodidae), in Sweden. Journal of Medical Entomology 31, 880886.
TALLEKLINT, L. & JAENSON, T. G. T. ( 1996). Relationship between Ixodes ricinus density and prevalence of infection with Borrelia-like spirochetes and density of infected ticks. Journal of Medical Entomology 33, 805811.
TALLEKLINT-EISEN, L. & LANE, R. S. ( 2000). Efficiency of drag sampling for estimating population sizes of Ixodes pacificus (Acari: Ixodidae) nymphs in leaf litter. Journal of Medical Entomology 37, 484487.
TATCHELL, R. J. & EASTON, E. ( 1986). Tick (Acari: Ixodidae) ecological studies in Tanzania. Bulletin of Entomological Research 76, 229246.
USPENSKY, I. ( 1995). Physiological age of Ixodid ticks: aspects of its determination and application. Journal of Medical Entomology 32, 751764.
UTECH, K. B. W., SEIFERT, G. W. & WHARTON, R. H. ( 1978). Breeding Australian Illawara shorthorn cattle for resistance to Boophilus microplus. I. Factors affecting resistance. Australian Journal of Agricultural Research 29, 411422.
VAN ES, R. P., HILLERTON, J. E. & GETTINBY, G. ( 1998). Lipid consumption in Ixodes ricinus (Acari: Ixodidae): temperature and potential longevity. Bulletin of Entomological Research 88, 567573.
VANDYK, J. K., BARTHOLOMEW, D. M., ROWLEY, W. A. & PLATT, K. B. ( 1996). Survival of Ixodes scapularis (Acari: Ixodidae) exposed to cold. Journal of Medical Entomology 33, 610.
VARLEY, G. C., GRADWELL, G. R. & HASSELL, M. P. ( 1973). Insect Population Ecology: An Analytical Approach. Oxford, Blackwell Scientific Publications.
WALKER, A. R. ( 2001). Age structure of a population of Ixodes ricinus (Acari: Ixodidae) in relation to its seasonal questing. Bulletin of Entomological Research 91, 6978.
WALKER, A. R., FLETCHER, J. D. & TODD, L. ( 1990). Resistance between stages of the tick Rhipicephalus appendiculatus (Acari: Ixodidae). Journal of Medical Entomology 27, 955961.
WILSON, M. L., ADLER, G. H. & SPEILMAN, A. ( 1985). Correlation between abundance of deer and that of the deer tick, Ixodes dammini (Acari: Ixodidae). Annals of the Entomological Society of America 78, 172176.
WOOLHOUSE, M. E. J., DYE, C., ETARD, J. F., SMITH, T., CHARLWOOD, J. D., GARNETT, G. P., HAGAN, P., HII, J. L. K., NDHLOVU, P. D., QUINNELL, R. J., WATTS, C. H., CHANDIWANA, S. K. & ANDERSON, R. M. ( 1997). Heterogeneities in the transmission of infectious agents: implications for the design of control programs. Proceedings of the National Academy of Sciences, USA 94, 338342.
WOOLHOUSE, M. E. J., ETARD, J.-F., DIETZ, K., NDHLOVU, P. D. & CHANDIWANA, S. K. ( 1998). Heterogeneities in schistosome transmission dynamics and control. Parasitology 117, 475482.
ZABICKA, J. ( 1994). [The tick-borne encephalitis in Poland.] (In Polish.) Przegl Epidemiology 48, 197203.
ZEMAN, P. & JANUSKA, J. ( 1999). Epizootiologic background of dissimilar distribution of human cases of Lyme borreliosis and tick-borne encephalitis in a joint endenic area. Comparative Immunology, Microbiology and Infectious Diseases 22, 247260.


Tick ecology: processes and patterns behind the epidemiological risk posed by ixodid ticks as vectors

  • S. E. RANDOLPH (a1)


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