Dec 2, 2000 - northwestern Florida (Panama City Beach, based on drag sampling, documented ... Florida. Rodent ectoparasites from two locations in northwestern Florida ..... J. Wrenn (University of North Dakota, Grand Forks) for help with ...
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Rodent ectoparasites from two locations in northwestern Florida Lance A. Durden, Renjie Hu1, James H. Oliver, Jr. and James E. Cilek2 Institute of Arthropodology and Parasitology Georgia Southern University, P. O. Box 8056, Statesboro, Georgia 30460 2 John A. Mulrennan, Sr., Public Health Entomology Research & Education Center Florida A & M University, 4000 Frankford Ave., Panama City, Florida 32405 1
Current address: California Department of Health Sciences Vector-Borne Diseases Section 2151 Convention Center Way, Suite 218B, Ontario, California 91764 Received 3 March 2000; Accepted 18 July 2000 ABSTRACT: From Feb.-Apr. 1999, 19 species of ectoparasitic arthropods (2 sucking lice, 4 fleas, 4 ticks, 2 mesostigmatid mites, 5 chiggers, 2 fur mites) were recovered from 106 rodents belonging to 5 species (cotton mouse, Peromyscus gossypinus, n=64; cotton rat, Sigmodon hispidus, n=23; eastern woodrat, Neotoma floridana, n=9; golden mouse, Ochrotomys nuttalli, n=9; eastern gray squirrel, Sciurus carolinensis, n=1) at Tall Timbers Research Station, Leon County, Florida. During the same period, 13 species of ectoparasites (2 sucking lice, 1 flea, 3 ticks, 3 mesostigmatid mites, 2 chiggers, 2 fur mites) were recovered from 57 rodents belonging to 3 species (S. hispidus, n=40; black rat, Rattus rattus, n=16; S. carolinensis, n=1) from Panama City, Bay County, Florida. Noteworthy ectoparasite records include Ixodes minor from both sites, which extends the known geographical range of this tick, and Stenoponia americana from Tall Timbers that represents the second documented Florida record of this flea. Potential tick vectors (Dermacentor variabilis and Ixodes scapularis) of zoonotic pathogens (Rickettsia rickettsii and Borrelia burgdorferi) were collected at both sites. On S. hispidus, fleas were more prevalent at Tall Timbers but sucking lice, chiggers, ticks, mesostigmatid and listrophorid mites all were more prevalent at the Panama City site. Arthropods recovered from arboreal nests (n=3) of O. nuttalli at Tall Timbers included 3 species of ectoparasites (1 tick, 2 laelapid mites). Journal of Vector Ecology 25(2): 222-228. 2000. Keyword Index: Ectoparasites, rodents, Florida INTRODUCTION Few studies have documented the ectoparasites of rodents in northwestern Florida. Most rodentectoparasite surveys within this state have been completed in peninsular Florida (Worth 1950a,b, 1951; Forrester 1992; Durden et al. 1993). Nevertheless, one wide-ranging survey of ectoparasites associated with commensal rats and mice (Pratt and Good 1954) included data for northwestern Florida. Also, because of their proximity to this part of Florida, some rodentectoparasite surveys from adjacent locations such as southwest Georgia (Morlan 1952, Smith and Love 1958), Jacksonville, Florida (Rumreich and Wynn 1945, Wilson et al. 1991), and Mobile, Alabama (Cole and Koepke 1946) have relevance to this region. Some surveys for the entire state of Florida (Layne 1971, Forrester 1992) have additionally included rodent-
ectoparasite data for this part of northern Florida. A recent two-year tick survey at a recreational area in northwestern Florida (Panama City Beach, based on drag sampling, documented the presence of four medically important tick species (Cilek and Olson 2000). Because of the role of some rodent ectoparasites, such as certain ticks and fleas, as vectors of zoonotic pathogens, and the reservoir potential of certain rodents for these pathogens, it is important to document hostparasite associations and infestation parameters for parasitic arthropods infesting rodents. In connection with a survey in the southeastern United States to identify potential vectors and reservoirs of Borrelia burgdorferi sensu lato, the etiologic agent of Lyme borreliosis in North America, we conducted a rodentectoparasite study at two locations in northwestern Florida.
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MATERIALS AND METHODS The study was completed at two sites. One site, the Tall Timbers Research Station, near Tallahassee (Leon County), is about 35 km from the Gulf of Mexico, and characterized by secondary growth fields, scrub, and woodland. The other site, Panama City (Bay County), is situated along the Gulf Coast and characterized by the same type of vegetation as the previous site but in tracts adjacent to residential neighborhoods. The distance between the two sites is approximately 160 km. Rodents were captured using two sizes (23 cm and 30 cm) of ShermanR traps (H. B. Sherman Traps, Tallahassee, Florida) during Feb.- Apr., 1999. These months were targeted because a previous study in southern Georgia (Durden and Oliver 1999) showed that immature stages of Ixodes scapularis, an important vector of B. burgdorferi sensu lato (see Oliver 1996), began attaching to rodents during early spring. Traps were baited with bird seed and arranged along random transects in areas of suspected rodent activity. Traps were set overnight and checked the following morning. The number of trap-nights was 1,023 at the Tall Timbers site and 456 at the Panama City site. Captured rodents were taken to a laboratory and anesthetized with 1:10 ketamine hydrochloride: xylazine sulphate injected intramuscularly. Each rodent was then visually checked for ectoparasites by carefully examining the ears, systematically parting the fur over the entire body surface, and then pulling a flea comb through the pelage. Ticks recovered were kept alive in humidified vials for spirochete screening in connection with a separate study. Other ectoparasites were stored in appropriately labeled vials containing 70% ethanol. All ectoparasites were subsequently identified using standard taxonomic keys and identification guides. Voucher specimens are deposited in the Institute of Arthropodology and Parasitology (IAP) at Georgia Southern University, Statesboro (accession numbers: L2300-L2327, L2378-L2412, L2420-2451, L24592498, L2500-L2527). Using Berlese funnel apparatus, arthropods were extracted and identified from three arboreal nests of the golden mouse, Ochrotomys nuttalli, collected at the Tall Timbers site. This was done to determine the prevalence and intensity of ectoparasitic arthropods, particularly potential tick vectors, in these nests. Voucher specimens of arthropods recovered from these nests are also deposited in the IAP (accession numbers: L2413, L2455, L2456).
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RESULTS AND DISCUSSION At the Tall Timbers site, 106 rodents belonging to 5 species were live trapped (trap success rate = 10.4%) and examined for ectoparasites. Table 1 lists infestation prevalence (percent of hosts infested), mean intensity (mean per infested host) following Bush et al. (1997), and numbers and stages recovered for ectoparasites collected from each rodent species at this site. Nineteen species of ectoparasites (2 sucking lice, 4 fleas, 4 ticks, 2 mesostigmatid mites, 5 chiggers, 2 fur mites) were recovered from rodents trapped at Tall Timbers. At the Panama City site, 57 rodents belonging to three species were live trapped (trap success rate = 12.5%) and examined for ectoparasites. Table 2 lists infestation data for ectoparasite species collected from each rodent species at this site. Thirteen species of ectoparasites (2 sucking lice, 1 flea, 3 ticks, 3 mesostigmatid mites, 2 chiggers, 2 fur mites) were recovered at this site. Representatives of a wide variety of arthropod higher taxa, most of them non-parasitic, were extracted from the three arboreal nests of O. nuttalli recovered at the Tall Timbers site. These included arachnids (spiders, ticks, and astigmatid, cheyletid, oribatid, and mesostigmatid mites) and immature and adult stages of representatives from six orders of insects (Collembola, Coleoptera, Hymenoptera, Lepidoptera, Psocoptera, and Thysanoptera). Of relevance to this study, three species of ectoparasites were recovered from the nests. These were the American dog tick, Dermacentor variabilis (5 nymphs, 1 larva - 2 nests infested), and the facultatively hematophagous laelapid mites Androlaelaps casalis (6 males, 50 females, 10 nymphs - 1 nest infested), and Androlaelaps fahrenholzi (3 males, 28 females, 6 nymphs - 3 nests infested). We recorded different communities of ectoparasites associated with rodents at two geographically different (i.e., coastal and suburban vs. inland and relatively undisturbed) sites in northwestern Florida. Reasons for the ectoparasite faunal differences between the two sites are not known. However, habitat differences, contrasting rodent species and abundances, and different numbers of trap nights between the two sites, presumably all contributed to these differences. Only two rodent species were trapped at both sites, the cotton rat, Sigmodon hispidus, and the eastern gray squirrel, Sciurus carolinensis. The latter species was inadequately sampled (one squirrel from each site) but there were noteworthy differences in ectoparasite infestation parameters for S. hispidus between the two
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Table 1. Ectoparasites recovered from 106 rodents at Tall Timbers Research Station, Leon Co., Florida, 1999 Host species
Ectoparasites*
Cotton mouse, Peromyscus gossypinus n=64 (38M,26F**)
Sucking louse: Hoplopleura hesperomydis (22%; 12.5; 37M,134F,4N**) Fleas: Polygenis gwyni (5%; 1.0; 2M,1F) Orchopeas leucopus (2%; 1.0; 1F) Stenoponia americana (2%; 1.0; 1F) Ticks: Dermacentor variabilis (17%; 2.2; 5N, 19L**) Ixodes minor (6%; 1.3; 5N) Ixodes scapularis (2%; 1.0; 1L) Mesostigmatid mites: Androlaelaps fahrenholzi (5%; 1.0; 3F) Ornithonyssus bacoti (2%; 1.0; 1F) Chiggers: Leptotrombidium peromysci (14%; 7.4; 67L) Eutrombicula cinnabaris (5%; 15.3; 46L) Euschoengastia peromysci (2%; 1.0; 1L) Sucking louse: Hoplopleura hirsuta (30%; 8.3; 10M,33F,16N) Fleas: Polygenis gwyni (83%; 4.4; 34M,50F) Orchopeas leucopus (4%; 1.0; 1M) Ticks: Amblyomma maculatum (4%; 1.0; 1N) Ixodes minor (4%; 1.0; 1N) Mesostigmatid mites: Androlaelaps fahrenholzi (17%; 5.8; 22F,1N) Ornithonyssus bacoti (4%; 1.0; 1F) Fur mites: Radfordia sigmodontis (9%; 1.5; 3F) Prolistrophorus bakeri*** (4%; ca. 600; not determined***) Chigger: Leptotrombidium peromysci (4%; 1.0; 1L) Flea: Orchopeas leucopus (11%; 1.0; 1M) Ticks: Ixodes minor (22%; 1.0; 1N,1L) Amblyomma maculatum (11%; 1.0; 1N) Dermacentor variabilis (11%; 1.0; 1L) Chiggers: Cheladonta ouachitensis (33%; 4.7; 14L) Euschoengastia peromysci (22%; 8.5; 17L) Neotrombicula whartoni (11%; 14.0; 14L) Eutrombicula cinnabaris (11%; 4.0; 4L) Leptotrombidium peromysci (11%; 3.0; 3L)
Cotton rat, Sigmodon hispidus n=23 (10M,13F)
Eastern woodrat, Neotoma floridana n=9 (3M,6F)
Continued
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Table 1. Continued. Golden mouse, Ochrotomys nuttalli n=9 (4M,5F)
Eastern gray squirrel, Sciurus carolinensis n=1 (1F)
Sucking louse: Hoplopleura hesperomydis (11%; 1.0; 1F) Ticks: Ixodes scapularis (22%; 1.0; 2L) Dermacentor variabilis (11%; 1.0; 1L) Ixodes minor (11%; 1.0; 1L) Chigger: Euschoengastia peromysci (11%; 2.0; 2L) Flea: Orchopeas howardi (100%; 1.0; 1F)
*For each ectoparasite species, infestation parameters listed are prevalence (% infested) and mean intensity (mean per infested host), respectively, followed by the numbers of the different life stages that were recovered. **M, Male(s); F, Female(s); N, Nymph(s); L, Larva(e). ***Infestation density was estimated for P. bakeri; different life stages of this mite were not counted. sites (Tables 1 and 2). The hoplopleurid sucking louse Hoplopleura hirsuta, the mesostigmatid mites A. fahrenholzi and Ornithonyssus bacoti, the chigger Eutrombicula cinnabaris, and the listrophorid fur mite Prolistrophorus bakeri, all were more common on S. hispidus in Panama City, whereas the rhopalopsyllid flea Polygenis gwyni was more common on this host at Tall Timbers. Other differences included the absence of Dermacentor variabilis on S. hispidus at Tall Timbers and the frequent infestation (prevalence, 35%) of this host by this tick in Panama City (Tables 1 and 2). Although other differences in ectoparasite species infesting S. hispidus at the two sites are evident, most were infrequently collected at either site and could reflect sampling biases. Differences in overall ectoparasite faunas were also evident between the two sites. For example, five species of chiggers (169 specimens) were recovered from 106 rodents at Tall Timbers but two species of chiggers (345 specimens) were recovered from 57 rodents in Panama City (Tables 1 and 2). The pest chigger E. cinnabaris predominated at the Panama City site but was relatively uncommon at the Tall Timbers site. However, much of the chigger diversity at Tall Timbers was associated with eastern woodrats, Neotoma floridana, which were not collected in Panama City. Also, just one species of flea was recorded in Panama City compared to four species at Tall Timbers. In part, this presumably reflected the larger number of rodents and rodent species sampled from the latter site. We recorded 12 species of ectoparasites from the cotton mouse, Peromyscus gossypinus, which was
trapped only at Tall Timbers (Table 1). The hoplopleurid sucking louse Hoplopleura hesperomydis, the ceratophyllid flea Orchopeas leucopus, and the large ctenophthalmid flea Stenoponia americana, are well documented ectoparasites of this mouse (Durden and Musser 1994, Durden et al. 1999). Similarly, the tick Dermacentor variabilis, and the chiggers Euschoengastia peromysci, and Leptotrombidium peromysci are frequently recorded ectoparasites of Peromyscus spp. mice (Whitaker 1968). The ectoparasites we recorded from S. hispidus (Tables 1 and 2) are mostly species that are welldocumented parasites of this rodent (Worth 1950a,b, 1951; Morlan 1952; Smith and Love 1958; Pfaffenberger and de Bruin 1988; Forrester 1992; Durden et al. 1993). Some of these ectoparasites such as H. hirsuta, P. gwyni, P. bakeri, and the myobiid fur mite Radfordia sigmodontis are associated exclusively or principally with this rodent (Whitaker and Wilson 1974; Durden et al. 1993, 1999; Durden and Musser 1994). The macronyssid mite Ornithonyssus bacoti is a vector of the filarial nematode Litomosoides carinii to cotton rats (Forrester 1992, Durden et al. 1993). We recorded relatively few ectoparasites from the black rat, Rattus rattus, which was trapped only in Panama City (Table 2). In previous studies, populations of black rats in port cities along the Gulf Coast and in Jacksonville, Florida have yielded larger numbers and more species of ectoparasites than we recorded (Rumreich and Wynn 1945, Cole and Koepke 1946, Pratt and Good 1954). Characteristic ectoparasites of black rats such as the spined rat louse Polyplax spinulosa, the Oriental rat flea Xenopsylla cheopis,
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Table 2. Ectoparasites recovered from 57 rodents in Panama City, Bay Co., Florida, 1999. Host species
Ectoparasites*
Cotton rat, Sigmodon hispidus n=40 (18M,22F**)
Sucking louse: Hoplopleura hirsuta (65%; 11.0; 25M,88F,172N**) Flea: Polygenis gwyni (28%; 2.2; 6M,18F) Ticks: Dermacentor variabilis (35%; 3.4; 47N,1L**) Ixodes scapularis (5%; 1.5; 3L) Ixodes minor (3%; 1.0; 1N) Mesostigmatid mites: Androlaelaps fahrenholzi (55%; 6.0; 1M,128F,2N) Ornithonyssus bacoti (13%; 2.6; 11F,2N) Haemogamasus liponyssoides (3%; 1.0; 1F) Fur mites: Prolistropohorus bakeri*** (38%; ca. 127; not determined***) Radfordia sigmodontis (3%; 4.0; 3F,1N) Chigger: Eutrombicula cinnabaris (35%; 23.2; 325L) Sucking louse: Polyplax spinulosa (13%; 3.0; 1F,5N) Flea: Polygenis gwyni (6%; 1.0; 1F) Ticks: Dermacentor variabilis (19%; 1.3; 2N,2L). Ixodes scapularis (13%; 2.0; 3N,1L) Chiggers: Eutrombicula cinnabaris (13%; 8.5; 17L) Euschoengastia peromysci (6%; 3.0; 3L) No ectoparasites recovered.
Black rat, Rattus rattus n=16 (9M,7F)
Eastern gray squirrel, Sciurus carolinesis n=1 (1M) *,**,***Footnotes as in Table 1.
the spiny rat mite Laelaps echidnina, and the tropical rat mite, O. bacoti, were all well documented in these earlier ectoparasite studies but were rare (P. spinulosa) or absent (X. cheopis, L. echidnina, O. bacoti) on black rats in our study. A wide variety of ectoparasites, most with little apparent host specificity, parasitize the eastern woodrat (Durden et al. 1997a) which we trapped only at the Tall Timbers site (Table 1). We collected few ectoparasites from the nine woodrats sampled, but relatively diverse tick (three species) and chigger (five species) faunas were associated with these animals (Table 1). Although the ceratophyllid flea Orchopeas sexdentatus pennsylvanicus parasitizes the eastern woodrat across much of the southeastern United States (Durden et al.
1997a), this ectoparasite does not appear to have been recorded in Florida (Layne 1971) and the only flea we recovered from a woodrat during this study was the congeneric O. leucopus (Table 1). Few ectoparasite records from the golden mouse are available. In a previous survey in Appalachian Tennessee, Linzey (1968) reported one species of sucking louse, four species of fleas, one species of tick, three species of mesostigmatids, one species of glycyphagid, and one species of myocoptid from this host. Like Linzey (1968), we recorded both the louse H. hesperomydis and the tick D. variabilis from O. nuttalli. However, we recorded two additional species of ticks (Ixodes minor and Ixodes scapularis) that would not be expected to parasitize this rodent in
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Appalachia and one species of chigger (E. peromysci) that would parasitize it there (Table 1). In addition to containing abundant arthropods assignable to various taxonomic groups, the three golden mouse nests that we sampled included three species of ectoparasites. Although two of these, A. casalis and A. fahrenholzi, are facultative hematophages with no known medical importance, the tick D. variabilis is a vector of the rickettsial agent of Rocky Mountain spotted fever and can cause tick paralysis in humans (Felz and Durden 1998). We trapped just one eastern gray squirrel at each site; the squirrel from Tall Timbers was parasitized by the ceratophyllid flea Orchopeas howardi. This is a common ectoparasite of this rodent in North America including northern Florida (Wilson et al. 1991). Some of our tick records are noteworthy. The medical importance of D. variabilis has already been underscored. Ixodes scapularis is not only a vector of B. burgdorferi sensu lato, but also of the causative agents of human granulocytic ehrlichiosis and human babesiosis in some regions (Magnarelli et al. 1995). In addition, I. minor appears to be involved in enzootic transmission cyles of B. burgdorferi or closely related spirochetes between rodents and birds (Oliver 1996, Durden et al. 1997b). Our records of I. minor from both sites (Tables 1 and 2) extend the known geographical distribution of this tick. Previously, it had been documented in the United States from coastal South Carolina, coastal Georgia, southwest Georgia and peninsular Florida (Morlan 1952, Tedders et al. 1992). We also collected immatures of the Gulf Coast tick, Amblyomma maculatum, at the Tall Timbers site; this tick can be a pest of livestock, pets, deer, and humans (Felz and Durden 1998). Cilek and Olson (2000) collected four species of ticks during a two-year dragbased survey in Panama City, Florida: Amblyomma americanum, A. maculatum, D. variabilis and I. scapularis. In the same general area, we recovered immature stages of two of these species (D. variabilis and I. scapularis) plus a nymph of a third species (I. minor) from rodents. Although we did not record important flea vectors such as X. cheopis, which can transmit murine typhus rickettsiae and plague bacilli, one notable flea record is evident. The S. americana specimen we collected at Tall Timbers represents the second documented record of this flea in Florida (Layne 1971). Coincidentally, the previous record in 1938, was from “Tallahassee” on a cotton mouse (Fox 1940) and was therefore from the same host species and possibly from the same locality as our second record more than 60 years later. Clearly, a distinct and varied community of
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ectoparasites is associated with rodents in the Florida panhandle and this appears to differ with respect to host diversity, proximity to the coast, and probably other factors also. Some of these ectoparasites have zoonotic importance through their role as vectors of pathogens. Acknowledgments We are indebted to A. A. Kinsey and M. A. Olson for field assistance and to C. W. Banks and A. R. Banks for laboratory assistance. Thanks are extended to L. A. Brennan, Director of the Tall Timbers Research Station, for allowing us to collect at this site. We also thank W. J. Wrenn (University of North Dakota, Grand Forks) for help with identifying some of the chiggers. This work was supported in part by National Institutes of Health grants AI 24899 and AI 40729. REFERENCES CITED Bush, A. O., K. D. Lafferty, J. M. Lotz and A. W. Shostak. 1997. Parasitology meets ecology on its own terms: Margolis et al. revisited. J. Parasitol. 83: 575-583. Cilek, J. E. and M. A. Olson. 2000. Seasonal distribution and abundance of ticks (Acari: Ixodidae) in northwestern Florida. J. Med. Entomol. 37: 439-444. Cole, L. C. and J. A. Koepke. 1946. A study of rodent ectoparasites in Mobile, Ala. Publ. Hlth. Rep. 41: 1469-1487. Durden, L. A., J. S. H. Klompen and J. E. Keirans. 1993. Parasitic arthropods of sympatric opossums, cotton rats, and cotton mice from Merritt Island, Florida. J. Parasitol. 79: 283-286. Durden, L. A., C. W. Banks, K. L. Clark, B. V. Belbey and J. H. Oliver, Jr. 1997a. Ectoparasite fauna of the eastern woodrat, Neotoma floridana: composition, origin, and comparison with ectoparasite faunas of western woodrat species. J. Parasitol. 83: 374-381. Durden, L. A., R. G. McLean, J. H. Oliver, Jr., S. R. Ubico and A. M. James. 1997b. Ticks, Lyme disease spirochetes, trypanosomes, and antibody to encephalitis viruses in wild birds from coastal Georgia and South Carolina. J. Parasitol. 83: 11781182. Durden, L. A. and G. G. Musser. 1994. The sucking lice (Insecta, Anoplura) of the world: a taxonomic checklist with records of mammalian hosts and geographical distributions. Bull. Am. Mus. Nat. Hist. 218: 1-90. Durden, L. A. and J. H. Oliver, Jr. 1999. Ecology of
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Ixodes scapularis and Lyme disease in coastal Georgia, USA, Pp. 379-383, in Acarology IX: symposia (G. R. Needham, R. Mitchell, D. J. Horn and W. C. Welbourn, eds.). Ohio Biological Survey, Columbus. 507 p. Durden, L. A., W. Wills and K. C. Clark. 1999. The fleas (Siphonaptera) of South Carolina with an assessment of their vectorial importance. J. Vector Ecol. 24: 171-181. Felz, M. W. and L. A. Durden. 1998. Identifying ticks: a pictorial guide. Patient Care 32(13): 172-187. Forrester, D. J. 1992. Parasites and diseases of wild mammals in Florida. University Press of Florida, Gainesville. 459 pp. Fox, I. 1940. Fleas of eastern United States. Iowa State College Press, Ames. 191 pp. Layne, J. N. 1971. Fleas (Siphonaptera) of Florida. Florida Entomol. 54: 35-51. Linzey, D. W. 1968. An ecological study of the golden mouse, Ochrotomys nuttalli, in the Great Smoky Mountains National Park. Am. Midl. Nat. 79: 320345. Magnarelli, L. A., J. S. Dumler, J. F. Anderson, R. C. Johnson and E. Fikrig. 1995. Coexistence of antibodies to tick-borne pathogens of babesiosis, ehrlichiosis, and Lyme borreliosis in human sera. J. Clin. Microbiol. 33: 3054-3057. Morlan, H. B. 1952. Host relationships and seasonal abundance of some southwest Georgia ectoparasites. Am. Midl. Nat. 48: 74-93. Oliver, J. H., Jr. 1996. Lyme borreliosis in the southern United States: a review. J. Parasitol. 82: 926-935. Pfaffenberger, G. S. and D. de Bruin. 1988. Parasites of the hispid cotton rat, Sigmodon hispidus (Cricetidae), and population biology of the cotton rat louse, Hoplopleura hirsuta (Hoplopleuridae: Anoplura), in eastern New Mexico, including an annotated host-parasite bibliography. Texas J. Sci. 40: 369-399.
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Pratt, H. D. and N. E. Good. 1954. Distribution of some common domestic rat ectoparasites in the United States. J. Parasitol. 40: 113-129. Rumreich, A. S. and R. S. Wynn. 1945. A study of the rodent-ectoparasite population of Jacksonville, Fla. Publ. Hlth. Rep. 60: 885-905. Smith, W. W. and G. J. Love. 1958. Ectoparasite populations and prevalence of murine typhus antibodies in cotton rats of southwest Georgia during wet and dry periods. Trans. Am. Microscopical Soc. 77: 48-56. Tedders, S. H., J. E. Keirans, D. C. Williams and T. A. Gwinn. 1992. First report of Ixodes (Ixodes) minor Neumann (Acari: Ixodidae) from South Carolina. J. Med. Entomol. 29: 282-283. Whitaker, J. O., Jr. 1968. Parasites, pp. 254-311, In: J. A. King (ed.), Biology of Peromyscus (Rodentia). Special Publication No. 2, American Society of Mammalogists. 593 pp. Whitaker, J. O., Jr. and N. Wilson. 1974. Host and distribution lists of mites (Acari), parasitic and phoretic, in the hair of wild mammals of North America, north of Mexico. Am. Midl. Nat. 91: 167. Wilson, N. A., S. R. Telford, Jr. and D. J. Forrester. 1991. Ectoparasites of a population of urban gray squirrels in northern Florida. J. Med. Entomol. 28: 461-464. Worth, C. B. 1950a. Observations on ectoparasites of some small mammals in Everglades National Park and Hillsborough County, Florida. J. Parasitol. 36: 326-335. Worth, C. B. 1950b. A preliminary host-ectoparasite register for some small mammals of Florida. J. Parasitol. 36: 497-498. Worth, C. B. 1951. Indirect evidence supporting observations on the range of wild rodents. J. Mammal. 32: 76-79.
