Amblyomma americanum (Acari: Ixodidae) ticks are not vectors of the lyme disease agent, Borrelia burgdorferi (Spirocheatales: Spirochaetaceae): A review of the evidence

Journal of Medical Entomology

Volumn 55, Issue 3, 2018, Pages 501-514

  Stromdahl, Ellen Y ^a   Nadolny, Robyn M ^a   Hickling, Graham J ^b   Hamer, Sarah A ^c   Ogden, Nicholas H ^d   Casal, Cory ^a   Heck, Garrett A ^a,e   Gibbons, Jennifer A ^f   Cremeans, Taylor F ^a   Pilgard, Mark A ^g  

^a Public Health Command (Prov)   (United States)

^b UNIVERSITY OF TENNESSEE INSTITUTE OF AGRICULTURE   (United States)

^c TEXAS A AND M UNIVERSITY   (United States)

^d PUBLIC HEALTH AGENCY OF CANADA   (Canada)

^e Department of Environmental and Radiological Health Sciences   (United States)

^f AMERICAN TYPE CULTURE COLLECTION   (United States)

^g NATIONAL CENTER FOR EMERGING AND ZOONOTIC INFECTIOUS DISEASES   (United States)

Author keywords

Amblyomma americanum; Borrelia burgdorferi; Prevalence; Vector competency

Indexed keywords

ANIMAL; ARACHNID VECTOR; BORRELIA BURGDORFERI; IXODIDAE; LYME DISEASE; MICROBIOLOGY; PHYSIOLOGY; TRANSMISSION;

ANIMALS; ARACHNID VECTORS; BORRELIA BURGDORFERI; IXODIDAE; LYME DISEASE;

EID: 85049747066     PISSN: 00222585     EISSN: 19382928     Source Type: Journal    
DOI: 10.1093/jme/tjx250     Document Type: Review

References (108)

1. Allan, B. F., L. S. Goessling, G. A. Storch, and R. E. Thach. 2010. Blood meal analysis to identify reservoir hosts for Amblyomma americanum ticks. Emerg. Infect. Dis. 16: 433-440.
2. Armstrong, P. M., S. M. Rich, R. D. Smith, D. L. Hartl, A. Spielman, and S. R. Telford, 3rd. 1996. A new Borrelia infecting Lone Star ticks. Lancet. 347: 67-68.
3. Armstrong, P. M., L. R. Brunet, A. Spielman, and S. R. Telford, 3rd. 2001. Risk of Lyme disease: perceptions of residents of a Lone Star tick-infested community. Bull. World Health Organ. 79: 916-925.
4. Bacon, R. M., R. D. Gilmore, Jr, M. Quintana, J. Piesman, and B. J. Johnson. 2003. DNA evidence of Borrelia lonestari in Amblyomma americanum (Acari: Ixodidae) in southeast Missouri. J. Med. Entomol. 40: 590-592.
5. Bacon, R. M., M. A. Pilgard, B. J. Johnson, S. J. Raffel, and T. G. Schwan. 2004. Glycerophosphodiester phosphodiesterase gene (glpQ) of Borrelia lonestari identified as a target for differentiating Borrelia species associated with hard ticks (Acari:Ixodidae). J. Clin. Microbiol. 42: 2326-2328.
6. Bacon, R. M., M. A. Pilgard, B. J. Johnson, J. Piesman, B. J. Biggerstaff, and M. Quintana. 2005. Rapid detection methods and prevalence estimation for Borrelia lonestari glpQ in Amblyomma americanum (Acari: Ixodidae) pools of unequal size. Vector Borne Zoonotic Dis. 5: 146-156.
7. Barbour, A. G., W. Burgdorfer, S. F. Hayes, O. Péter, and A. Aeschlimann. 1983a. Isolation of a cultivable spirochete from Ixodes ricinus ticks of Switzerland. Curr. Microbiol. 8:123-126.
8. Barbour, A. G., S. L. Tessier, and W. J. Todd. 1983b. Lyme disease spirochetes and ixodid tick spirochetes share a common surface antigenic determinant defined by a monoclonal antibody. Infect. Immun. 41: 795-804.
9. Barbour, A. G., G. O. Maupin, G. J. Teltow, C. J. Carter, and J. Piesman. 1996. Identification of an uncultivable Borrelia species in the hard tick Amblyomma americanum: possible agent of a Lyme disease-like illness. J. Infect. Dis. 173: 403-409.
10. Bockenstedt, L. K., D. G. Gonzalez, A. M. Haberman, and A. A. Belperron. 2012. Spirochete antigens persist near cartilage after murine Lyme borreliosis therapy. J. Clin. Invest. 122: 2652-2660.
11. Bouchard, C., P. A. Leighton, G. Beauchamp, S. Nguon, L. Trudel, F. Milord, L. R. Lindsay, D. Bélanger, and N. H. Ogden. 2013. Harvested white-tailed deer as sentinel hosts for early establishing Ixodes scapularis populations and risk from vector-borne zoonoses in southeastern Canada. J. Med. Entomol. 50: 384-393.
12. Bunikis, J., J. Tsao, U. Garpmo, J. Berglund, D. Fish, and A. G. Barbour. 2004. Typing of Borrelia relapsing fever group strains. J. Emerg. Infect. Dis. 10: 1161-1164.
13. Burgdorfer, W., A. G. Barbour, S. F. Hayes, J. L. Benach, E. Grunwaldt, and J. P. Davis. 1982. Lyme disease-a tick-borne spirochetosis? Science. 216: 1317-1319.
14. Burkot, T. R., G. R. Mullen, R. Anderson, B. S. Schneider, C. M. Happ, and N. S. Zeidner. 2001. Borrelia lonestari DNA in adult Amblyomma americanum ticks, Alabama. Emerg. Infect. Dis. 7: 471-473.
15. Carroll, J. F., and T. L. Cyr. 2005. A note on the densities of Ixodes scapularis (Acari:Ixodidae) and white-tailed deer on the campus of the National Institutes of Standards and Technology, Maryland, USA. Proc. Entomol. Soc. Wash. 107: 973-976.
16. Castellaw, A. H., J. Showers, J. Goddard, E. F. Chenney, and A. S. Varela-Stokes. 2010. Detection of vector-borne agents in lone star ticks, Amblyomma americanum (Acari: Ixodidae), from Mississippi. J. Med. Entomol. 47: 473-476.
17. Clark, K. 2004. Borrelia species in host-seeking ticks and small mammals in northern Florida. J. Clin. Microbiol. 42: 5076-5086.
18. Clark, K. L., J. H. Oliver, Jr, A. M. James, L. A. Durden, and C. W. Banks. 2002. Prevalence of Borrelia burgdorferi sensu lato infection among rodents and host-seeking ticks in South Carolina. J. Med. Entomol. 39: 198-206.
19. Clark, K., A. Hendricks, and D. Burge. 2005. Molecular identification and analysis of Borrelia burgdorferi sensu lato in lizards in the southeastern United States. Appl. Environ. Microbiol. 71: 2616-2625.
20. Clark, K. L., B. Leydet, and S. Hartman. 2013. Lyme borreliosis in human patients in Florida and Georgia, USA. Int. J. Med. Sci. 10: 915-931.
21. Crowder, C. D., H. E. Matthews, S. Schutzer, M. A. Rounds, B. J. Luft, O. Nolte, S. R. Campbell, C. A. Phillipson, F. Li, R. Sampath, et al. 2010. Genotypic variation and mixtures of Lyme Borrelia in Ixodes ticks from North America and Europe. PLos One. 5: e10650.
22. Cyr, T. L., M. C. Jenkins, R. D. Hall, E. J. Masters, and G. A. McDonald. 2005. Improving the specificity of 16S rDNA-based polymerase chain reaction for detecting Borrelia burgdorferi sensu lato-causative agents of human Lyme disease. J. Appl. Microbiol. 98: 962-970.
23. Demaerschalck, I., A. Ben Messaoud, M. De Kesel, B. Hoyois, Y. Lobet, P. Hoet, G. Bigaignon, A. Bollen, and E. Godfroid. 1995. Simultaneous presence of different Borrelia burgdorferi genospecies in biological fluids of Lyme disease patients. J. Clin. Microbiol. 33: 602-608.
24. Feir, D., C. R. Santanello, B. W. Li, C. S. Xie, E. Masters, R. Marconi, and G. Weil. 1994. Evidence supporting the presence of Borrelia burgdorferi in Missouri. Am. J. Trop. Med. Hyg. 51: 475-482.
25. Fritzen, C. M., J. Huang, K. Westby, J. D. Freye, B. Dunlap, M. J. Yabsley, M. Schardein, J. R. Dunn, T. F. Jones, and A. C. Moncayo. 2011. Infection prevalences of common tick-borne pathogens in adult lone star ticks (Amblyomma americanum) and American dog ticks (Dermacentor variabilis) in Kentucky. Am. J. Trop. Med. Hyg. 85: 718-723.
26. Fryxell, R. T., C. D. Steelman, A. L. Szalanski, K. L. Kvamme, P. M. Billingsley, and P. C. Williamson. 2012. Survey of Borreliae in ticks, canines, and white-tailed deer from Arkansas, U.S.A. Parasit. Vectors. 5: 139.
27. Gleim E. R. 2013. The effects of long-term prescribed burning on ticks and tick-borne pathogen prevalence. Ph.D. dissertation, The University of Georgia, Athens, GA.
28. Gleim, E. R., L. E. Garrison, M. S. Vello, M. Y. Savage, G. Lopez, R. D. Berghaus, and M. J. Yabsley. 2016. Factors associated with tick bites and pathogen prevalence in ticks parasitizing humans in Georgia, USA. Parasit. Vectors. 9: 125.
29. Goddard, J., J. W. Sumner, W. L. Nicholson, C. D. Paddock, J. Shen, and J. Piesman. 2003. Survey of ticks collected in Mississippi for Rickettsia, Ehrlichia, and Borrelia species. J. Vector Ecol. 28: 184-189.
30. Guttman, D. S., P. W. Wang, I. N. Wang, E. M. Bosler, B. J. Luft, and D. E. Dykhuizen. 1996. Multiple infections of Ixodes scapularis ticks by Borrelia burgdorferi as revealed by single-strand conformation polymorphism analysis. J. Clin. Microbiol. 34: 652-656.
31. Guy, E. C., and G. Stanek. 1991. Detection of Borrelia burgdorferi in patients with Lyme disease by the polymerase chain reaction. J. Clin. Pathol. 44: 610-611.
32. Han, S., G. J. Hickling, and J. I. Tsao. 2016. High Prevalence of Borrelia miyamotoi among Adult Blacklegged Ticks from White-Tailed Deer. Emerg. Infect. Dis. 22: 316-318.
33. Harmon, J. R., M. C. Scott, E. M. Baker, C. J. Jones, and G. J. Hickling. 2015. Molecular identification of Ehrlichia species and host bloodmeal source in Amblyomma americanum L. from two locations in Tennessee, United States. Ticks Tick Borne Dis. 6: 246-252.
34. Haynes, J. M., L. P. Prescott, R. L. Seipelt, and S. M. Wright. 2005. Detection of Borrelia burgdorferi sequences in a biopsy from a Tennessee patient. J. TN Acad. Sci. 80: 57-59.
35. Hudman, D. A., and N. J. Sargentini. 2016. Detection of Borrelia, Ehrlichia, and Rickettsia spp. in ticks in northeast Missouri. Ticks Tick Borne Dis. 7: 915-921.
36. James, A. M., D. Liveris, G. P. Wormser, I. Schwartz, M. A. Montecalvo, and B. J. Johnson. 2001. Borrelia lonestari infection after a bite by an Amblyomma americanum tick. J. Infect. Dis. 183: 1810-1814.
37. Johnson, B. J., C. M. Happ, L. W. Mayer, and J. Piesman. 1992. Detection of Borrelia burgdorferi in ticks by species-specific amplification of the flagellin gene. Am. J. Trop. Med. Hyg. 47: 730-741.
38. Jordan, B. E., K. R. Onks, S. W. Hamilton, S. E. Hayslette, and S. M. Wright. 2009. Detection of Borrelia burgdorferi and Borrelia lonestari in birds in Tennessee. J. Med. Entomol. 46: 131-138.
39. Kardatzke, J. T., K. Neidhardt, D. P. Dzuban, J. L. Sanchez, and A. F. Azad. 1992. Cluster of tick-borne infections at Fort Chaffee, Arkansas: Rickettsiae and Borrelia burgdorferi in ixodid ticks. J. Med. Entomol. 29: 669-672.
40. Killmaster, L. F., A. D. Loftis, G. E. Zemtsova, and M. L. Levin. 2014. Detection of bacterial agents in Amblyomma americanum (Acari: Ixodidae) from Georgia, USA, and the use of a multiplex assay to differentiate Ehrlichia chaffeensis and Ehrlichia ewingii. J. Med. Entomol. 51: 868-872.
41. Kollars, T. M. Jr, J. H. Oliver, P. G. Kollars Jr, A. M. James, A. Richardson, L. A. Durden, D. O. Miles, and E. J. Masters. 2000. Phenotypic variation in Borrelia burgdorferi sensu lato in ticks (Acari: Ixodidae) and isolates from Missouri, USA. Int. J. Acarol. 26: 167-172.
42. Lane, R. S., J. Piesman, and W. Burgdorfer. 1991. Lyme borreliosis: relation of its causative agent to its vectors and hosts in North America and Europe. Annu. Rev. Entomol. 36: 587-609.
43. Ledin, K. E., N. S. Zeidner, J. M. Ribeiro, B. J. Biggerstaff, M. C. Dolan, G. Dietrich, L. Vredevoe, and J. Piesman. 2005. Borreliacidal activity of saliva of the tick Amblyomma americanum. Med. Vet. Entomol. 19: 90-95.
44. Lee, J. K., W. C. Smith, C. McIntosh, F. G. Ferrari, B. Moore-Henderson, and A. Varela-Stokes. 2014. Detection of a Borrelia species in questing Gulf Coast ticks, Amblyomma maculatum. Ticks Tick Borne Dis. 5: 449-452.
45. Levine, J. F., C. S. Apperson, W. L. Nicholson. 1989. The occurrence of spirochetes in ixodid ticks in North Carolina. J. Entomol. Sci. 24: 594-602.
46. Levine, J. F., D. E. Sonenshine, W. L. Nicholson, and R. T. Turner. 1991. Borrelia burgdorferi in ticks (Acari: Ixodidae) from coastal Virginia. J. Med. Entomol. 28: 668-674.
47. Lin, T., J., H. Oliver, Jr, and L. Gao. 2002. Genetic diversity of the outer surface protein C gene of southern Borrelia isolates and its possible epidemiological, clinical, and pathogenetic implications. J. Clin. Microbiol. 40: 2572-2583.
48. Lo, Y. M., and K. C. Chan. 2006. Setting up a polymerase chain reaction laboratory. Methods Mol. Biol. 336: 11-18.
49. Luckhart, S., G. R. Mullen, and J. C. Wright. 1991. Etiologic agent of Lyme disease, Borrelia burgdorferi, detected in ticks (Acari: Ixodidae) collected at a focus in Alabama. J. Med. Entomol. 28: 652-657.
50. Luckhart, S., G. R. Mullen, L. A. Durden, and J. C. Wright. 1992. Borrelia sp. in ticks recovered from white-tailed deer in Alabama. J. Wildl. Dis. 28: 449-452.
51. Maegli, A., J. D. Loy, and R. Cortinas. 2016. Note on Ehrlichia chaffeensis, Ehrlichia ewingii, and "Borrelia lonestari" infection in lone star ticks (Acari: Ixodidae), Nebraska, USA. Ticks Tick Borne Dis. 7: 154-158.
52. Magnarelli, L. A., J. F. Anderson, C. S. Apperson, D. Fish, R. C. Johnson, and W. A. Chappell. 1986. Spirochetes in ticks and antibodies to Borrelia burgdorferi in white-tailed deer from Connecticut, New York State, and North Carolina. J. Wildl. Dis. 22: 178-188.
53. Margos, G., A. G. Gatewood, D. M. Aanensen, K. Hanincová, D. Terekhova, S. A. Vollmer, M. Cornet, J. Piesman, M. Donaghy, A. Bormane, et al. 2008. MLST of housekeeping genes captures geographic population structure and suggests a European origin of Borrelia burgdorferi. Proc. Natl. Acad. Sci. USA 105: 8730-8735.
54. Marques, A., S. R. Telford, 3rd, S. P. Turk, E. Chung, C. Williams, K. Dardick, P. J. Krause, C. Brandeburg, C. D. Crowder, H. E. Carolan, et al. 2014. Xenodiagnosis to detect Borrelia burgdorferi infection: a first-in-human study. Clin. Infect. Dis. 58: 937-945.
55. Masters, E. J., C. N. Grigery, and R. W. Masters. 2008. STARI, or Masters disease: Lone Star tick-vectored Lyme-like illness. Infect. Dis. Clin. North Am. 22: 361-376, viii.
56. Mather, T. N., and M. E. Mather. 1990. Intrinsic competence of three ixodid ticks (Acari) as vectors of the Lyme disease spirochete. J. Med. Entomol. 27: 646-650.
57. Mitchell, E. A., P. C. Williamson, P. M. Billingsley, J. P. Seals, E. E. Ferguson, and M. S. Allen. 2016. Frequency and distribution of Rickettsiae, Borreliae, and Ehrlichiae detected in human-parasitizing ticks, Texas, USA. Emerg. Infect. Dis. 22: 312-315.
58. Mixson, T. R., S. R. Campbell, J. S. Gill, H. S. Ginsberg, M. V. Reichard, T. L. Schulze, and G. A. Dasch. 2006. Prevalence of Ehrlichia, Borrelia, and Rickettsial agents in Amblyomma americanum (Acari: Ixodidae) collected from nine states. J. Med. Entomol. 43: 1261-1268.
59. Moyer, P. L., A. S. Varela, M. P. Luttrell, V. A. Moore, 4th, D. E. Stallknecht, and S. E. Little. 2006. White-tailed deer (Odocoileus virginianus) develop spirochetemia following experimental infection with Borrelia lonestari. Vet. Microbiol. 115: 229-236.
60. Mukolwe, S. W., A. A. Kocan, R. W. Barker, K. M. Kocan, and G. L. Murphy. 1992. Attempted transmission of Borrelia burgdorferi (Spirochaetales: Spirochaetaceae) (JDI strain) by Ixodes scapularis (Acari: Ixodidae), Dermacentor variabilis, and Amblyomma americanum. J. Med. Entomol. 29: 673-677.
61. Nelson, J. A. 1995. Evidence supporting the presence of Borrelia burgdorferi in Missouri. Infect. Dis. Clin. Prac. 4: 110-111
62. Oliver, J. H., Jr, F. W. Chandler, Jr, M. P. Luttrell, A. M. James, D. E. Stallknecht, B. S. McGuire, H. J. Hutcheson, G. A. Cummins, and R. S. Lane. 1993. Isolation and transmission of the Lyme disease spirochete from the southeastern United States. Proc. Natl. Acad. Sci. USA 90: 7371-7375.
63. Ouellette, J., C. S. Apperson, P. Howard, T. L. Evans, and J. F. Levine. 1997. Tick-raccoon associations and the potential for Lyme disease spirochete transmission in the coastal plain of North Carolina. J. Wildl. Dis. 33: 28-39.
64. Philipp, M. T., E. Masters, G. P. Wormser, W. Hogrefe, and D. Martin. 2006. Serologic evaluation of patients from Missouri with erythema migranslike skin lesions with the C6 Lyme test. Clin. Vaccine Immunol. 13: 1170-1171.
65. Pichon, B., D. Egan, M. Rogers, and J. Gray. 2003. Detection and identification of pathogens and host DNA in unfed host-seeking Ixodes ricinus L. (Acari: Ixodidae). J. Med. Entomol. 40: 723-731.
66. Piesman, J., and R. J. Sinsky. 1988. Ability to Ixodes scapularis, Dermacentor variabilis, and Amblyomma americanum (Acari: Ixodidae) to acquire, maintain, and transmit Lyme disease spirochetes (Borrelia burgdorferi). J. Med. Entomol. 25: 336-339.
67. Piesman, J., and C. M. Happ. 1997. Ability of the Lyme disease spirochete Borrelia burgdorferi to infect rodents and three species of human-biting ticks (blacklegged tick, American dog tick, lone star tick) (Acari:Ixodidae). J. Med. Entomol. 34: 451-456.
68. Rawlings, J. A., and G. J. Teltow. 1994. Prevalence of Borrelia (Spirochaetaceae) spirochetes in Texas ticks. J. Med. Entomol. 31: 297-301.
69. Rich, S. M., P. M. Armstrong, R. D. Smith, and S. R. Telford, 3rd. 2001. Lone star tick-infecting borreliae are most closely related to the agent of bovine borreliosis. J. Clin. Microbiol. 39: 494-497.
70. Rijpkema, S. G., M. J. Molekenboer, L. M. Schouls, F. Jongejan, and J. G. Schellekens. 1995. Simultaneous detection and genotyping of three genomic groups of Borrelia burgdorferi sensu lato in Dutch Ixodes ricinus ticks by characterization of the ampli? ed intergenic spacer region between 5S and 23S rRNA genes. J. Clin. Microbiol. 33: 3091-3095.
71. Rollend, L., D. Fish, and J. E. Childs. 2013. Transovarial transmission of Borrelia spirochetes by Ixodes scapularis: a summary of the literature and recent observations. Ticks Tick Borne Dis. 4: 46-51.
72. Roome, A., L. Hill, V. Al-Feghali, C. G. Murnock, J. A. Goodsell, R. Spathis, and R. M. Garruto. 2017. Impact of white-tailed deer on the spread of Borrelia burgdorferi. Med. Vet. Entomol. 31: 1-5.
73. Rosa, P. A., D. Hogan, and T. G. Schwan. 1991. Polymerase chain reaction analyses identify two distinct classes of Borrelia burgdorferi. J. Clin. Microbiol. 29: 524-532.
74. Rudenko, N., M. Golovchenko, L. Grubhoffer, and J. H. Oliver, Jr. 2011. Updates on Borrelia burgdorferi sensu lato complex with respect to public health. Ticks Tick Borne Dis. 2: 123-128.
75. Rudenko, N., M. Golovchenko, K. Clark, J. H. Oliver, and L. Grubhoffer. 2016. Detection of Borrelia burgdorferi sensu stricto in Amblyomma americanum ticks in the southeastern United States: the case of selective compatibility. Emerg. Microbes Infect. 5: e48.
76. Ryder, J. W., R. R. Pinger, and T. Glancy. 1992. Inability of Ixodes cookei and Amblyomma americanum nymphs (Acari: Ixodidae) to transmit Borrelia burgdorferi. J. Med. Entomol. 29: 525-530.
77. Sanders, F. H., Jr, and J. H. Oliver, Jr. 1995. Evaluation of Ixodes scapularis, Amblyomma americanum, and Dermacentor variabilis (Acari: Ixodidae) from Georgia as vectors of a Florida strain of the Lyme disease spirochete, Borrelia burgdorferi. j. Med. Entomol. 32: 402-406.
78. Sayler, K. A., A. D. Loftis, S. K. Beatty, C. L. Boyce, E. Garrison, B. Clemons, M. Cunningham, A. R. Alleman, and A. F. Barbet. 2016. Prevalence of tick-borne pathogens in host-seeking Amblyomma americanum (Acari: Ixodidae) and Odocoileus virginianus (Artiodactyla: Cervidae) in Florida. J. Med. Entomol. 53: 949-956.
79. Sayler, K., J. Rowland, C. Boyce, and E. Weeks. 2017. Borrelia burgdorferi DNA absent, multiple Rickettsia spp. DNA present in ticks collected from a teaching forest in North Central Florida. Ticks Tick Borne Dis. 8: 53-59.
80. Schulze, T. L., G. S. Bowen, E. M. Bosler, M. F. Lakat, W. E. Parkin, R. Altman, B. G. Ormiston, and J. K. Shisler. 1984. Amblyomma americanum: A potential vector of Lyme disease in New Jersey. Science 224: 601-603.
81. Schulze, T. L., M. F. Lakat, W. E. Parkin, J. K. Shisler, D. J. Charette, and E. M. Bosler. 1986. Comparison of rates of infection by the Lyme disease spirochete in selected populations of Ixodes dammini and Amblyomma americanum (Acari: Ixodidae). Zentralbl. Bakteriol. Mikrobiol. Hyg. A. 263: 72-78.
82. Schulze, T. L., R. A. Jordan, C. J. Schulze, T. Mixson, and M. Papero. 2005. Relative encounter frequencies and prevalence of selected Borrelia, Ehrlichia, and Anaplasma infections in Amblyomma americanum and Ixodes scapularis (Acari: Ixodidae) ticks from central New Jersey. J. Med. Entomol. 42: 450-456.
83. Schulze, T. L., R. A. Jordan, S. P. Healy, V. E. Roegner, M. Meddis, M. B. Jahn, and D. L. Guthrie Sr. 2006. Relative abundance and prevalence of selected Borrelia infections in Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae) from publicly owned lands in Monmouth County, New Jersey. J. Med. Entomol. 43: 269-275.
84. Schulze, T. L., R. A. Jordan, J. C. White, V. E. Roegner, and S. P. Healy. 2011. Geographical distribution and prevalence of selected Borrelia, Ehrlichia, and Rickettsia infections in Amblyomma americanum (Acari: Ixodidae) in New Jersey. J. Am. Mosq. Control Assoc. 27: 236-244.
85. Simpson, K. K., and L. W. Hinck. 1993. The prevalence of Borrelia burgdorferi, the Lyme disease spirochete, in ticks and rodents in Northeast Arkansas. Proc. Arkansas Acad. Sci. 47:110-114.
86. Soares, C. A., N. S. Zeidner, C. B. Beard, M. C. Dolan, G. Dietrich, and J. Piesman. 2006. Kinetics of Borrelia burgdorferi infection in larvae of refractory and competent tick vectors. J. Med. Entomol. 43: 61-67.
87. Sonenshine, D. E., R. E. Ratzlaff, J. Troyer, S. Demmerle, E. R. Demmerle, W. E. Austin, S. Tan, B. A. Annis, and S. Jenkins. 1995. Borrelia burgdorferi in eastern Virginia: comparison between a coastal and inland locality. Am. J. Trop. Med. Hyg. 53: 123-133.
88. Springer, Y. P., L. Eisen, L. Beati, A. M. James, and R. J. Eisen. 2014. Spatial distribution of counties in the continental United States with records of occurrence of Amblyomma americanum (Ixodida: Ixodidae). J. Med. Entomol. 51: 342-351.
89. Steere, A. C., S. E. Malawista, D. R. Snydman, R. E. Shope, W. A. Andiman, M. R. Ross, and F. M. Steele. 1977. Lyme arthritis: an epidemic of oligoarticular arthritis in children and adults in three connecticut communities. Arthritis Rheum. 20: 7-17.
90. Stegall-Faulk, T., D. C. Clark, and S. M. Wright. 2003. Detection of Borrelia lonestari in Amblyomma americanum (Acari: Ixodidae) from Tennessee. J. Med. Entomol. 40: 100-102.
91. Straubinger, R. K. 2000. PCR-Based quantification of Borrelia burgdorferi organisms in canine tissues over a 500-Day postinfection period. J. Clin. Microbiol. 38: 2191-2199.
92. Stromdahl, E. Y., and G. J. Hickling. 2012. Beyond Lyme: aetiology of tickborne human diseases with emphasis on the south-eastern United States. Zoonoses Public Health. 59 (Suppl 2): 48-64.
93. Stromdahl, E. Y., S. R. Evans, J. J. O'Brien, and A. G. Gutierrez. 2001. Prevalence of infection in ticks submitted to the human tick test kit program of the U.S. Army Center for Health Promotion and Preventive Medicine. J. Med. Entomol. 38: 67-74.
94. Stromdahl, E. Y., P. C. Williamson, T. M. Kollars, Jr, S. R. Evans, R. K. Barry, M. A. Vince, and N. A. Dobbs. 2003. Evidence of Borrelia lonestari DNA in Amblyomma americanum (Acari: Ixodidae) removed from humans. J. Clin. Microbiol. 41: 5557-5562.
95. Stromdahl, E. Y., R. M. Nadolny, J. A. Gibbons, L. D. Auckland, M. A. Vince, C. E. Elkins, M. P. Murphy, G. J. Hickling, M. W. Eshoo, H. E. Carolan, et al. 2015. Borrelia burgdorferi not confirmed in human-biting Amblyomma americanum ticks from the southeastern United States. J. Clin. Microbiol. 53: 1697-1704.
96. Taft, S. C., M. K. Miller, and S. M. Wright. 2005. Distribution of borreliae among ticks collected from eastern states. Vector Borne Zoonotic Dis. 5: 383-389.
97. Telford, S. R., 3rd, T. N. Mather, S. I. Moore, M. L. Wilson, and A. Spielman. 1988. Incompetence of deer as reservoirs of the Lyme disease spirochete. Am. J. Trop. Med. Hyg. 39: 105-109.
98. Teltow, G. J., P. V. Fournier, and J. A. Rawlings. 1991. Isolation of Borrelia burgdorferi from arthropods collected in Texas. Am. j. Trop. Med. Hyg. 44: 469-474.
99. Tsao, J. I., J. T. Wootton, J. Bunikis, M. G. Luna, D. Fish, and A. G. Barbour. 2004. An ecological approach to preventing human infection: vaccinating wild mouse reservoirs intervenes in the Lyme disease cycle. Proc. Natl. Acad. Sci. USA 101: 18159-18164.
100. Varela, A. S., M. P. Luttrell, E. W. Howerth, V. A. Moore, W. R. Davidson, D. E. Stallknecht, and S. E. Little. 2004a. First culture isolation of Borrelia lonestari, putative agent of southern tick-associated rash illness. J. Clin. Microbiol. 42: 1163-1169.
101. Varela, A. S., V. A. Moore, and S. E. Little. 2004b. Disease agents in Amblyomma americanum from northeastern Georgia. J. Med. Entomol. 41: 753-759.
102. Varela-Stokes, A. S. 2007. Transmission of bacterial agents from lone star ticks to white-tailed deer. J. Med. Entomol. 44: 478-483.
103. Williamson, P. C., P. M. Billingsley, G. J. Teltow, J. P. Seals, M. A. Turnbough, and S. F. Atkinson. 2010. Borrelia, Ehrlichia, and Rickettsia spp. in ticks removed from persons, Texas, USA. Emerg. Infect. Dis. 16: 441-446.
104. Wormser, G. P., E. Masters, D. Liveris, J. Nowakowski, R. B. Nadelman, D. Holmgren, S. Bittker, D. Cooper, G. Wang, and I. Schwartz. 2005a. Microbiologic evaluation of patients from Missouri with erythema migrans. Clin. Infect. Dis. 40: 423-428.
105. Wormser, G. P., E. Masters, J. Nowakowski, D. McKenna, D. Holmgren, K. Ma, L. Ihde, L. F. Cavaliere, and R. B. Nadelman. 2005b. Prospective clinical evaluation of patients from Missouri and New York with erythema migrans-like skin lesions. Clin. Infect. Dis. 41: 958-965.
106. Xu, G., T. N. Mather, C. S. Hollingsworth, and S. M. Rich. 2016. Passive surveillance of Ixodes scapularis (Say), their biting activity, and associated pathogens in Massachusetts. Vector Borne Zoonotic Dis. 16: 520-527.
107. Yuan, D. T., 2010: A metagenomic study of the tick midgut. UT GSBS Dissertations and Theses. Paper 85. University of Texas Graduate School of Biomedical Sciences, Houston, TX. http://digitalcommons.library.tmc.edu/utgsbs-dissertations8/85.
108. Zeidner, N., A. Ullmann, C. Sackal, M. Dolan, G. Dietrich, J. Piesman, and D. Champagne. 2009. A borreliacidal factor in Amblyomma americanum saliva is associated with phospholipase A2 activity. Exp. Parasitol. 121: 370-375.