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Biomedical subjects

J E Keirans

Publications and source records attributed to J E Keirans.

At least 19 recordsLinked to original sources

Ixodes (Haemixodes) longiscutatum Boero (new status) and I. (H.) uruguayensis Kohls & Clifford, a new synonym of I. (H.) longiscutatum (Acari: Ixodidae).

Females of Ixodes (Haemixodes) uruguayensis Kohls & Clifford, 1967, a species whose adults were unknown until the present, were obtained in the laboratory from engorged nymphs collected on rodents (Scapteromys tumidus and Oxymycterus nasutus) in the counties of Maldonado and San José, Uruguay. Morphological characters of these females were identical to those given in the description of the female of Ixodes longiscutatum Boero, 1944. I. uruguayensis is, thus, relegated to a junior subjective synonym of I. longiscutatum. However, because of the unique morphological characters of the immature stages, the validity of the subgenus Haemixodes Kohls & Clifford, 1967 is not questioned. Therefore, the new status of Ixodes (Haemixodes) uruguayensis Kohls & Clifford, 1967 is Ixodes (Haemixodes) longiscutatum Boero, 1944.

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Invasion: exotic ticks (Acari: Argasidae, Ixodidae) imported into the United States. A review and new records.

A review of the literature and unpublished records from the U.S. National Tick Collection on the importation of ticks from foreign lands reveals that at least 99 exotic tick species assignable to 11 genera have been either detected and destroyed at ports of entry or inadvertently imported into the United States in the past half century. This number includes four argasid and 95 ixodid species, some of which are important vectors of agents that cause disease to both man and animals. If one includes Aponomma sp. and Hyalomma sp. and the subspecies of Rhipicephalus, the total exceeds 100 taxa. It is notable that the number of imported tick species recorded herein exceeds the total number of tick species native to the United States. It appears that the soft tick genera Argas, Antricola and Nothoaspis have not been imported, although at some point in time Argas persicus (Oken) was introduced because it is resident although not often collected. The hard tick genera Anomalohimalaya, Cosmiomma, Margaropus, Nosomma and Rhipicentor, and the nuttalliellid genus Nuttalliella have also not been imported.

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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.

A portion of mitochondrial 12S rDNA sequences (337-355 base pairs) and 63 morphological characters of 36 hard-tick species belonging to 7 genera were analyzed to determine the phylogenetic relationships among groups and species of Rhipicephalus and between the genera Rhipicephalus and Boophilus. Molecular and morphological data sets were first examined separately. The molecular data were analyzed by maximum parsimony (MP), maximum likelihood, and neighbor-joining distance methods; the morphological data were analyzed by MP After their level of congruence was evaluated by a partition homogeneity test, all characters were combined and analyzed by MP. The branches of the tree obtained by combining the data sets were better resolved than those of the trees inferred from the separate analyses. Boophilus is monophyletic and arose within Rhipicephalus. Boophilus species clustered with species of the Rhipicephalus evertsi group. Most of the clustering within Rhipicephalus was, however, consistent with previous classifications based on morphological data. Morphological characters were traced on the molecular reconstruction in order to identify characters diagnostic for monophyletic clades. Within the Rhipicephalus sanguineus complex, the sequences of specimens morphologically identified as Rhipicephalus turanicus were characterized by a high level of variability, indicating that R. turanicus-like morphology may cover a spectrum of distinct species.

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Ticks associated with wild animals in the Nhecolândia Pantanal, Brazil.

A study of ticks associated with wild animals was carried out from September 1996 to April 1998 at the Fazenda Alegria (21,000 ha), in the Nhecolândia Pantanal, State of Mato Grosso do Sul, Brazil, a sunken plain bordering the upper Paraguay river, located 19 x 08'S; 56 x 46'W. A total of 81 wild animals (13 species, 6 orders) were captured with the aid of nets, and ticks were found on 63 (78%). Tick species identified included Boophilus microplus (Canestrini), Amblyomma cajennense (F.), A. parvum (Aragão), A pseudoconcolor (Aragão), A. scalpturatum (Neumann), A. nodosum (Neumann), A. ovale (Koch), and A. tigrinum (Koch). Dragging from grasslands (campos) yielded negative results compared to the high concentration of ticks, mainly nymphs, that were collected from leaves in the forests (capão). Predominance of immature instars (Amblyomma genera) was observed in the end of winter (August-September). Ticks were associated mainly with coatis, deer (Mazama gouazoubira) and anteater, and these animals may play a role in the epidemiology of tick-transmitted pathogens in the Pantanal if one considers their co-existence with local domestic animals.

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Ticks (Acari: Ixodidae) associated with wild animals in the Pantanal region of Brazil.

This paper describes the identification of ticks from wild animals of the Pantanal region in Brazil as part of a comprehensive study about established and emerging tick-host relationships and related pathological aspects. Eighty-one animals were captured (representing 13 species, six orders), and ticks were found on 63 (78%). Tick species identified included Boophilus microplus (Canestrini), Amblyomma cajennense (F.), Amblyomma parvum Aragão, Amblyomma pseudoconcolor Aragão, Amblyomma scalpturatum Neumann, Amblyomma nodosum Neumann, Amblyomma ovale Koch, and Amblyomma tigrinum Koch. Dragging from grasslands yielded negative results compared with the high concentration of ticks that were collected from leaves in the forests.

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Evaluation of host preferences by helminths and ectoparasites among black-tailed jackrabbits in northern California.

Fifty-four black-tailed jackrabbits (Lepus californicus) (five juvenile males, 22 adult males, five juvenile females, and 22 adult females) from Humboldt County, California (USA) were evaluated for sex and age-specific differences in parasite prevalences and intensities, 26 February through 30 October 1996. Nematodes found included Biogastranema leporis in 42 hares (78% prevalence), Rauschia triangularis in 26 hares (48%), Trichostrongylus calcaratus in 14 hares (26%), and Trichuris sylvilagi in two hares (4%). Cestodes found included Taenia sp. cysticerci in five hares (9%) and Taenia sp. coenurus found in one hare (2%). Ectoparasites found included the ticks Dermacentor variabilis on 10 hares (19%) and Ixodes spinipalpis (= Ixodes neotomae) on nine hares (17%), as well as the anoplurid louse Haemodipsus setoni on 12 hares (22%). No significant differences in the parasite prevalences or intensities were found between male and female jackrabbits; this was for all males and females collectively, juvenile males and females only, as well as adult males and females only. Combining male and female hosts, adult jackrabbits had a significantly higher prevalence of B. leporis and R. triangularis compared to juveniles. This is the first known report of Trichostrongylus calcaratus, Rauschia triangularis, Trichuris sylvilagi, and Dermacentor variabilis among black-tailed jackrabbits and the first known report of T. calcaratus and T. sylvilagi in the western USA. This is the first published report of I. spinipalpis, the vector for Lyme disease in California, on black-tailed jackrabbits.

Age Distribution↗

Molecular genetic key for the identification of 17 Ixodes species of the United States (Acari:Ixodidae): a methods model.

A taxonomic key, based on restriction enzyme analysis of the second internal-transcribed spacer (ITS-2) in the nuclear ribosomal DNA gene, was developed for identification of 17 Ixodes tick species in the United States. This key includes: Ixodes affinis Neumann, Ixodes angustus Neumann, Ixodes baergi Cooley and Kohls, Ixodes brunneus Koch, Ixodes cookei Packard, Ixodes dentatus Marx, Ixodes jellisoni Cooley and Kohls, Ixodes kingi Bishopp, Ixodes minor Neumann, Ixodes muris Bishopp and Smith, Ixodes pacificus Cooley and Kohls, Ixodes scapularis Say, Ixodes sculpularis Neumann, I. spinipalpis Hadwen and Nuttall, Ixodes texanus Banks, Ixodes uriae White, and Ixodes woodi Bishopp. A 900-bp fragment of the ITS-2 was amplified using the polymerase chain reaction. This fragment was then digested with the restriction enzymes MspI and CfoI, and the digested fragments were size fractionated on a 2.5% high-resolution agarose gel. A dichotomous key was developed based on digested fragment sizes relative to a standard set of size markers. Little intraspecific variation in restriction fragment banding patterns was detected.

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A world checklist of genera, subgenera, and species of ticks (Acari: Ixodida) published from 1973-1997.

Researchers on ticks and tickborne diseases have been extremely fortunate in having at their fingertips the tick bibliographies produced by Harry Hoogstraal and his coworkers at the U.S. Naval facility at Cairo, Egypt, and by Mildred Doss and her colleagues at the U.S. Department of Agriculture laboratory at Beltsville, Maryland, USA. The Doss checklist of tick families, genera, species, and subspecies is now 25 years out of date, and the following checklist of one new genus, nine new subgenera, and 110 new species of Ixodida brings together the nomenclature on ticks produced during the last quarter century.

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Illustrated key to nymphs of the tick genus Amblyomma (Acari: Ixodidae) found in the United States.

A dichotomous identification key, with important characters illustrated by scanning electron micrographs, is provided for nymphs of the 9 species of the genus Amblyomma occurring in the United States. Species included are A. americanum (L.), A. cajennense (F.), A. dissimile Koch, A. imitator Kohls, A. inornatum (Banks), A. longirostre (Koch), A maculatum Koch, A. rotundatum Koch, and A. tuberculatum Marx. Notes on the geographical distributions, medical and veterinary importance, and nymphal stage hosts are included for each species.

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First records of Amblyomma americanum, Ixodes (Ixodes) dentatus, and Ixodes (Ceratixodes) uriae (Acari: Ixodidae) from Maine.

The first records of 3 ixodid tick species collected in the state of Maine are reported. A total of 23 records of the lone star tick, Amblyomma americanum (L., 1758), in 11 counties from hosts with no history of travel outside the state demonstrates that this tick is now a resident of Maine. Ixodes dentatus Marx, 1899 is recorded from Waldo and Lincoln counties, and Ixodes uriae White, 1852 is recorded from Matinicus Rock in Knox County. This is the first report of I. uriae from the eastern United States. Disease agents such as those causing human monocytic ehrlichiosis, Lyme disease, Rocky Mountain spotted fever, and several arboviruses have been recorded from 1 or more of these tick species.

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Phylogenetic relationships among tick subfamilies (Ixodida: Ixodidae: Argasidae) based on the 18S nuclear rDNA gene.

Phylogenetic relationships among tick subfamilies have been estimated using morphological and molecular characters. However, the phylogeny based on a portion of the 16S mitochondrial rDNA gene differed from the morphologically based phylogeny in a number of important respects. The entire 18S rDNA gene was examined in 18 taxa from all tick subfamilies to test the 16S rDNA based phylogeny. The 18S phylogeny supports the earlier 16S based phylogeny in placing members of Hyalomminae on a common branch with members of the Rhipicephalinae and in indicating long branch lengths among soft tick taxa. However, unlike the 16S phylogeny, Amblyomminae was monophyletic and members of Haemaphysalinae did not arise within Amblyomminae. Argasinae formed a monophyletic group within Argasidae and was not a sister taxon of the hard ticks. In most respects, the phylogeny based on the 18S rDNA gene resembles the morphologically based phylogeny.

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Taxonomic status of Ixodes neotomae and I. spinipalpis (Acari: Ixodidae) based on mitochondrial DNA evidence.

Ixodes spinipalpis Hadwen & Nuttall and I. neotomae Cooley are enzootic vectors of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner in western North America. The taxa overlap in host preference, habitat, and morphology. Mitochondrial DNA was compared between the taxa to test for reproductive isolation. A 300-bp region of the mitochondrial 16S ribosomal DNA gene was amplified in 28 specimens of I. neotomae and 149 specimens of I. spinipalpis. These products were screened for sequence variation using single-strand conformation polymorphism analysis, and 9 haplotypes were detected. Haplotype frequencies varied between taxa; however, Shannon diversity analysis indicated that most variation arose among collections within each taxon, and no unique haplotypes characterized either one. Phylogenetic analysis of 18 sequences, representing a replicate of each of the 9 haplotypes, was performed with Ixodes pacificus Cooley & Kohls and Ixodes jellisoni Cooley & Kohls as outgroups. Strong monophyletic support was found for a clade containing I. neotomae and I spinipalpis and within this clade no lineages comprised exclusively either taxon. These results argue against treatment of I. neotomae and I. spinipalpis as distinct species.

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Population genetics of Ixodes scapularis (Acari: Ixodidae) based on mitochondrial 16S and 12S genes.

Blacklegged tick, Ixodes scapularis Say, is the principal vector of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner in the eastern half of the United States. Populations exhibit extreme variation in morphology, host usage, development time, and behavior. We examined sequence variation in the 16S and 12S mitochondrial ribosomal DNA genes to determine genetic relationships among I. scapularis collections from throughout its range. Single strand conformation polymorphism analysis of 300 bp of the 16S molecule was used to identify different haplotypes and estimate their relative frequencies among 198 ticks. Eleven different haplotypes were detected. Haplotype diversity was least in northeastern collections and greatest in the southeast. The 11 haplotypes were sequenced in 24 specimens. In total, 462 bp in the 16S gene and 420 bp in the 12S gene were sequenced to reveal 66 informative sites. Phylogenetic analysis, using I. ricinus L. and I. pacificus Cooley & Kohls as outgroups, revealed 2 clades within I. scapularis. One clade was limited to the South and the other was distributed throughout the range of I. scapularis. Specimens from the Southern United States were basal in the broadly distributed clade. Random amplified polymorphic DNA by polymerase chain reaction patterns examined between members of the 2 clades provided no evidence for reproductive isolation. These patterns suggest that I. scapularis arose in the South but that a large geographic split gave rise to 2 distinct lineages. These lineages now interbreed and are partially sympatric.

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Ixodes (Ixodes) scapularis (Acari:Ixodidae): redescription of all active stages, distribution, hosts, geographical variation, and medical and veterinary importance.

The blacklegged tick, Ixodes (Ixodes) scapularis Say, 1821, is redescribed, based on laboratory reared specimens originating in Bulloch County, Georgia. Information on distribution, host associations, morphological variation, and medical/veterinary importance is also presented. A great deal of recent work has focused on this species because it is the principal vector of the agent of Lyme disease (Borrelia burgdorferi Johnson, Schmidt, Hyde, Steigerwaldt & Brenner) in eastern North America. Its distribution appears to be expanding, and includes the state of Florida in the southeastern United States north to the provinces of Nova Scotia and Prince Edward Island, Canada, west to North and South Dakota, United States, and south to the state of Coahuila, Mexico. Although I. scapularis feeds on at least 125 species of North American vertebrates (54 mammalian, 57 avian, and 14 lizard species), analysis of the U.S. National Tick Collection holdings show that white-tailed deer, Odocoileus virginianus (Zimmermann), cattle, Bos taurus L., dogs, Canis lupus L., and other medium-to-large sized mammals are important hosts for adults as are native mice and other small mammals, certain ground-frequenting birds, skinks, and glass lizards for nymphs and larvae. This tick is a polytypic species exhibiting north-south and east-west morphological clines. Analysis of variance and Student-Newman-Keuls multiple comparisons revealed significant interpopulational variation that is expressed most significantly in the nymphal stage. Nymphs from northern (Minnesota, Massachusetts, Maryland) populations had relatively larger basis capituli with shorter cornua (except Maryland) than southern (North Carolina, Georgia) populations. Midwestern populations (Minnesota, Missouri) differed from eastern populations (Massachusetts, Maryland, North Carolina, Georgia) in idiosomal characters (broader scuta, larger coxae III, and IV). In addition to Lyme disease, this tick is also a primary vector of the agent of human and rodent babesiosis, Babesia microti Franca. Under laboratory conditions it has transmitted the agents of deer babesiosis, Babesia odocoilei Emerson & Wright, tularemia, Francisella tularensis McCoy & Chapin, and anaplasmosis, Anaplasma marginale Theiler. Moreover, I. scapularis can reach pest proportions on livestock, and females can cause tick paralysis in dogs.

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Ixodes (Ixodes) jellisoni and I. (I.) neotomae (Acari:Ixodidae): descriptions of the immature stages from California.

Nymphal and larval stages of Ixodes (Ixodes) jellisoni Cooley & Kohls and I. (I.) neotomae Cooley are described for the first time. These 2 tick species occur only in the western United States, predominantly in California. The primary host for I. jellisoni is the California kangaroo rat, Dipodomys californicus (Merriam); that for I. neotomae is the dusky-footed woodrat, Neotoma fuscipes Baird. The etiologic agent of Lyme disease Borrelia burgdorferi Johnson, Schmidt, Hyde, Steigerwalt & Brenner has recently been isolated from both tick species, and I. neotomae was proven a competent enzootic vector of the Lyme disease spirochete.

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Evolution of ticks.

Evolutionary patterns in ticks have traditionally been cast in terms of host associations. Largely untested assumptions of cospeciation and observations of current host associations are used to estimate the age of different taxa. Several recent phylogenetic studies of supraspecific relationships in ticks, based on both morphological and DNA-sequence data, allow the first rigorous testing of these assumptions. Reanalysis of patterns of tick-host associations suggests that the perception of host specificity in ticks may be an artifact of incomplete sampling. An analysis of tick-host and -habitat associations and biogeographical patterns, in the context of the newly derived phylogenies, suggests that much of the existing host-association patterns may be explained as artifacts of biogeography and ecological specificity.

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Rhipicephalus interventus sp. nov. (Acari: Ixodidae), a new tick species closely related to Rhipicephalus tricuspis Dönitz, 1906 and Rhipicephalus Iunulatus Neumann, 1907, from east and central Africa.

Descriptions of the adults of this new species are given, together with information on its hosts and distribution. Previously it was referred to briefly by Walker, Keirans, pegram & Clifford (1988), who noted that in many respects it is intermediate in appearance between R. tricuspis and R. Iunulatus.

Africa, Central↗