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During 1998-2000, at least 14 species (n = 309) of small mammals were live-trapped and examined for ectoparasites in moist forests of the Taita and Shimba Hills and drier savannah habitats of Nguruman, southeastern Kenya. Ectoparasites were recorded from 11 species of mammals. Five species of sucking lice [Hoplopleura inexpectans Johnson, H. intermedia Kellogg & Ferris, Polyplax reclinata (Nitzsch), P. waterstoni Bedford and Schizophthirus graphiuri Ferris], six species of fleas (Ctenophthalmus leptodactylous Hubbard, Dinopsyllus grypurus Jordan & Rothschild, D. lypusus Jordan & Rothschild, Hypsophthalmus campestris Jordan & Rothschild, Listropsylla basilewskyi Smit and Xiphiopsylla lippa Jordan) and at least six species of ticks (Amblyomma sp., Haemaphysalis sp., Ixodes sp., I. alluaudi Neumann, I. cumulatimpunctatus Schulze, I. muniensis Arthur & Burrow and Rhipicephalus sp.) were recorded from these hosts. Four of the five species of sucking lice were host specific whereas P. reclinata was recorded from two different species of white-toothed shrews, Crocidura spp. Although fleas and ticks were less host specific, C. leptodactylous, D. grypurus and I. cumulatimpunctatus were only recorded from the murid rodent Praomys delectorum (Thomas), Amblyomma sp. was only recorded from the nesomyid rodent Beamys hindei Thomas, Rhipicephalus sp. was only recorded from the murid Lemniscomys striatus (L.) and I. muniensis was only recorded from the dormouse Graphiurus microtis (Noack). More species of ectoparasites and significantly greater infestation prevalences were recorded from small mammals in moist habitats compared with those from the savannah habitat. At least one of the fleas recorded, D. lypusus, is a known vector of Yersinia pestis Lehmann & Neumann, the causative agent of plague, which is present in the region.

Specimens of Amblyomma oudemansi (Neumann, 1910) were collected in Papua New Guinea from an endangered monotreme, Zaglossus bruijni (Peters & Doria), the western long-beaked echidna. These ticks were compared morphologically and molecularly with species formerly assigned to Aponomma Neumann, 1899 (now included in Bothriocroton Keirans, King, & Sharrad, 1994 or Amblyomma Koch, 1844), and a phylogeny was generated. Based on our results, we reassign this tick to Bothriocroton, as B. oudemansi (Neumann, 1910) n. comb. Original descriptions are provided for the female and the nymph of this species and the male is redescribed. A revised list of all Bothriocroton records and holdings in the US National Tick Collection is also provided.

Ixodes (Pholeoixodes) gregsoni Lindquist, Wu, and Redner, a species of hard tick described in 1999 in Canada, was recovered from a harvested fisher (Martes pennanti Erxleben) and a domestic cat (Felis silvestris catus Ragni and Randi) in Vermont in 2001 and from harvested mink (Mustela vison Schreber) in Maine in 2003. These samples are the first records of this species within the United States. Although knowledge of this tick's natural history and distribution are still preliminary, these records indicate a possible greater distribution for I. gregsoni than initially perceived. Although its status as a disease vector is presently unknown, natural resource professionals should be aware of the possibility of this tick's occurrence in the northeastern United States.

Anaplasma marginale Theiler is a tick-borne pathogen that causes anaplasmosis in cattle. There are approximately 20 tick species worldwide that are implicated as vectors of this pathogen. In the United States, Dermacentor andersoni Stiles and Dermacentor variabilis (Say) are the principal vectors. The risk of transmission of anaplasmosis to cattle has been largely based on the distribution of D. andersoni in the United States. We developed a centralized geographic database that incorporates collection records for D. andersoni from two large national databases. We reviewed the geographic records in each database and postings from MEDLINE and AGRICOLA to produce a national county-level distribution map based on a total of 5,898 records. The records spanned the period from 1903 through 2001 with the majority between 1921 to 1940. Populations of D. andersoni were recorded from 267 counties in 14 states and were distinguished as either established or reported. We found 180 counties with established populations of D. andersoni and 87 counties with reported occurrences in 14 states with the majority of established populations reported from Montana, Idaho, and Oregon. D. andersoni populations in the United States currently extend from the western portions of Nebraska and the Dakotas westward to the Cascade Mountains and from the northern counties of Arizona and New Mexico northward to the Canadian border. The data will be useful for identifying regions at increased risk of acquiring anaplasmosis in the United States. Based upon the database collection records, we also present a summary of recorded hosts for D. andersoni and comments on its seasonal occurrence.

Calreticulin (CRT) is a unique eukaryotic gene. The CRT gene product, calreticulin, was first identified as a calcium binding protein in 1974, but further investigations have indicated that CRT protein performs many functions in cells, including involvement in evading the host's immune system by parasites. Many studies of CRT have been published since the molecule was first discovered; however, the CRT gene exon-intron structure is only known for a limited number of ectoparasite species. In this study, we compared tick CRT genomic sequences to the corresponding cDNA from 28 species and found that 2 exons and 1 intron are present in the tick CRT gene. The intron position is conserved in 28 hard ticks, but intron size and nucleotide sequences vary. Three tick introns possess duplicated fragments and are twice as long as other introns. All tick CRT introns obey the GT-AG rule in the splice-site junctions and are phase 1 introns. By comparing tick CRT introns to those of fruit fly, mouse, and human, we conclude that tick CRT introns belong to the intron-late type. The number and size of CRT introns have increased through the evolution of eukaryotes.

In 2000, we initiated an investigation on the tick fauna of Rondônia State, where we collected many specimens of Amblyomma scalpturatum Neumann, 1906 and Amblyomma incisum Neumann, 1906. In addition, we also collected a third group of ticks that were morphologically closely related to those 2 species, but sufficiently different to be considered a distinct species; members of this group were subsequently identified as Amblyomma latepunctatum Tonelli-Rondelli, 1939, through comparison with the type specimens of this taxon. Herein, we redescribe both sexes of A. scalpturatum and A. incisum, the female of A. latepunctatum, and provide the first description of the male of this latter species. Molecular analysis of the second internal transcribed spacer (ITS2) from rDNA of specimens of the 3 species supports morphological results. Examination of both A. scalpturatum and A. incisum deposited in different tick collections revealed that A. latepunctatum appeared relatively frequently in the vials believed to contain specimens of A. incisum or A. scalpturatum. Before this study, A. latepunctatum was considered a synonym of A. scalpturatum. Herein, we provide morphological and molecular evidence to validate the species A. latepunctatum. The South American tapir (Tapirus terrestris) seems to be the primary host for the adult stage of A. latepunctatum, A. scalpturatum, and A. incisum.

Ticks exotic for the Neotropical region were found on Python regius imported into Argentina and Chile. All ticks (7 males and 3 females) were classified as Amblyomma latum Koch, 1844 ( = Aponomma latum). Additionally, four lots comprising 18 males of the Argentinean tortoise tick, Amblyomma argentinae Neumann, 1904, were found on a terrestrial tortoise, Chelonoidis chilensis, and on three terrestrial tortoises (probably C. chilensis) imported to Uruguay presumably from Argentina). These findings alert us to the risk of expanding the distribution of reptile parasites and their diseases into regions previously free of these parasites.

New Neotropical records are presented for ticks belonging to the Ixodes auritulus Neumann, 1904, species group, together with a review of hosts and localities from which members of this complex have previously been collected. The range of the I. auritulus species group is now understood to include Colombia, and 15 bird species are listed as new hosts. From Guatemala to southern Argentina and Chile, specimens of the I. auritulus group have been found on birds belonging to the orders Ciconiiformes, Columbiformes, Falconiformes, Galliformes, Passeriformes, Piciformes, Procellariiformes and Tinamiformes. Passeriform birds are probably the principal hosts, sustaining tick populations throughout the Neotropics. Collection data have yielded four areas--southern South America (from 56 degrees S to 51 degrees S), southern Brazil (25 degrees S-22 degrees S), south-central Peru (14 degrees S-10 degrees S) and Central America (10 degrees N-15 degrees N)--where the I. auritulus group appears to commonly parasitize birds, but additional collections may show that the range of this complex is less discontinuous than currently perceived. Several morphological differences are described for ticks within and among these areas, but it is still unclear whether the I. auritulus group comprises more than one species.

Ticks of the genus Ixodes are poorly known in the Neotropical zoogeographical region, from where only 45 species have been recorded. In Brazil, the genus is currently represented by eight species, four of which are known only from this country. This paper presents a redescription of all active stages in the life-cycle of I. loricatus , based on optical and scanning electron microscopy. In addition, the relationship of I. loricatus to other Neotropical Ixodes is presented and discussed.

The female of Ixodes neuquenensis Ringuelet, 1947 (Acari: Ixodidae) is redescribed and the nymph and larva are described from specimens collected from the endangered marsupial Dromiciops gliroides Thomas (Microbiotheria: Microbiotheriidae) in Argentina. At first sight the female of I. neuquenensis resembles a member of the subgenus Ixodes Latreille, 1795. However, the female of I. neuquenensis is peculiar in having the combination of two spurs on coxae II-IV and a pair of chitinous plaques internal to coxa I. Both the nymph and larva have an anterior and posterior process on palpal article I, characteristics of the subgenus Ixodiopsis Filippova, 1957 and some representatives of the subgenus Pholeoixodes Schulze, 1942. Analysis of 16S mitochondrial rDNA sequences showed no strong relationship with any known Ixodes subgenus.

Calreticulin (CRT) is a calcium-binding protein and has many functions in eukaryotic cells. CRT is possibly involved in parasite host immune system evasion. To better understand the molecular basis of CRT in ticks, we cloned and sequenced 4 full-length complementary DNAs (cDNAs) from the hard tick species, Dermacentor variabilis, Haemaphysalis longicornis, Ixodes scapularis, and Rhipicephalus sanguineus, using the technique of rapid amplification of cDNA ends. The deduced amino acid sequences share high identities (between 77 and 98%) with 3 known tick CRT sequences. The major characteristics of known CRTs are observed in all 4 of our deduced tick CRTs. These include 3 major domains, a signal peptide sequence at the beginning of the coding region, 2 triplets of conserved regions, cysteine sites providing disulfide bridges for N-terminal folding, and a nuclear localization signal. Remarkably, the replacement of the endoplasmic reticulum retention signal KDEL by HEEL, which is believed to be associated with secretion of CRT into the host during feeding and was previously recorded only in 2 ticks and a hookworm, is also present in all 4 of our tick putative CRTs. In addition, the CRT gene is potentially useful for tick phylogenetic reconstruction.

Two putative cDNAs of acetylcholinesterase (AChE), one from Dermacentor variabilis, and the other from Rhipicephalus sanguineus, were amplified and sequenced. The deduced amino acid sequences have high amino acid identities (between 70 and 94%) to known tick AChE sequences deposited in GenBank. Furthermore, these two AChEs also possess common features in their primary AChE structure such as catalytic active sites. A 2,220-bp contiguous sequence, containing a 1,791-bp open reading frame encoding an AChE precursor with 596 amino acid residues, was obtained from D. variabilis. The deduced proteins of R. sanguineus are different in size by 6 amino acids because of alternative splicing at the 5' end. A gene tree deduced from phylogenetic analysis indicates that there are at least three lineages of AChE in arthropods.

Five persons contracted tick-borne relapsing fever after staying in a cabin in western Montana. Borrelia hermsii was isolated from the blood of two patients, and Ornithodoros hermsi ticks were collected from the cabin, the first demonstration of this bacterium and tick in Montana. Relapsing fever should be considered when patients who reside or have vacationed in western Montana exhibit a recurring febrile illness.

The Ixodes ricinus species complex is a group of ticks distributed in almost all geographic regions of the world. Lyme borreliosis spirochetes are primarily transmitted by tick species within this complex. It has been hypothesized that the Lyme vector ticks around the world are closely related and represent a monophyletic group. This implies that vector competence in ixodid ticks for Lyme agents might have evolved only once. To test this hypothesis, we used a molecular phylogenetic approach. Two fragments of mitochondrial 16S ribosomal deoxyribonucleic acid were sequenced from 11 species in the I. ricinus complex and from 16 other species of Ixodes. Phylogenetic analysis using Bayesian methodology indicated that the I. ricinus complex is not a monophyletic group unless 3 additional Ixodes species are included in it. The known major vectors of Lyme disease agents in different areas of the world are not sister taxa. This suggests that acquisition of the ability to transmit borreliosis agents in species of Ixodes may have multiple origins.

DNA sequences of Amblyomma aureolatum (Pallas, 1772) and Amblyomma ovale Koch, 1844 were obtained to determine genetic differences between these tick species. Collections of these species are discussed in relation to distribution and hosts. Seven ticks collections (four from Brazil, one from Argentina, one from Uruguay and one from USA) house a total of 1272 A. aureolatum (224 males, 251 females, 223 nymphs and 574 larvae) and 1164 A. ovale (535 males, 556 females, 66 nymphs and 7 larvae). The length of the sequenced mitochondrial 16S rRNA gene fragment for A. aureolatum was 370bp and for A. ovale was 373bp. The DNA sequence analysis showed a 13.1% difference between the two species. Apart from one male A. ovale found on a toad, all adult ticks were found on mammals. The majority of adult specimens of both tick species were removed from Carnivora (96.1 and 84.3% of A. aureolatum and A. ovale, respectively), especially from dogs (53.1% of A. aureolatum, and 46.4% of A. ovale). Collections on wild Canidae were higher for A. aureolatum (23.3%) than for A. ovale (7.1%). On the other hand, collections of A. ovale adults on wild Felidae were higher (18.3%) than findings of A. aureolatum (9.2%). The contribution of other mammalian orders as hosts for adults of A. aureolatum and A. ovale was irrelevant, with the exception of Perissodactyla because Tapiridae contributed with 13.0% of the total number of A. ovale adults. Adults of both tick species have been found occasionally on domestic hosts (apart of the dog) and humans. Most immature stages of A. aureolatum were found on Passeriformes birds, while rodents and carnivores were the most common hosts for nymphs and larvae of A. ovale. A. aureolatum has been found restricted to the Neotropical region, covering the eastern area of South America from Uruguay to Surinam, including northeastern Argentina, eastern Paraguay, southeastern Brazil and French Guiana. A. ovale showed a distribution that covers the Neotropical region from central-northern Argentina throughout the Neotropics into the Nearctic region of Mexico with a few records from the USA, also with collection sites in Paraguay, Bolivia, most Brazilian states, Peru, Ecuador, French Guiana, Surinam, Guyana, Trinidad & Tobago, Venezuela, Colombia, Panama, Costa Rica, Nicaragua, Belize, Guatemala and several states of Mexico.

Partial mitochondrial 16S ribosomal ribonucleic acid gene sequences in the ticks Carios capensis collected from black-footed albatross. Diomedea nigripes, colonies on Torishima Island, Japan (30 degrees 28'N, 140 degrees 18'E), were examined. The sequence was compared with those of C. capensis from Hawaii, South Carolina, and Texas. The sequences were all identical in ticks from Torishima and 2 from Hawaii. There were 2-3 transitions between the other Hawaiian and Texas ticks and Torishima specimens. Two transitions were also observed when compared with the ticks from South Carolina. The results suggest the possibility of gene flow between tick populations at each of the 2 geographic sites, which probably was accomplished by tick-infested migratory seabirds at their breeding sites. Sequence comparison analysis indicated that the C. capensis ticks are on the branch with C. marginatus and C. mexicanus ticks and not with Ornithodoros. This supports the revision suggested by Klompen and Oliver (1993).

Collections of Amblyomma auricularium (Conil 1878) and A. pseudoconcolor Aragão, 1908 are discussed in relation to distribution and hosts. Three tick collections (two from Argentina and a third from the USA) house a total of 574 A. auricularium (307 males, 162 females, 73 nymphs and 32 larvae) and 179 A. pseudoconcolor (96 males, 74 females, 4 nymphs and 5 larvae). Apart from an adult A. pseudoconcolor found on a bird, Nothura maculosa Temminck, 1815, all ticks were found on mammals. The great majority of specimens of both ticks species were removed from the family Dasypodidae Gray, 1821 (84.9% and 93.8% of A. auricularium and A. pseudoconcolor, respectively). Amblyomma auricularium has also been found on wild hosts of the families Myrmecophagidae and occasionally Didelphidae, Caviidae, Chinchillidae, Hydrochaeridae, Muridae, Canidae, Mustelidae, Procyonidae and domestic animals (cattle, dogs, horses), while A. pseudoconcolor has also been found occasionally on wild hosts of the family Didelphidae and on domestic animals (cattle, dogs). Amblyomma pseudoconcolor appears to be restricted to the Neotropical region, covering northern Argentina and the eastern region of South America from Uruguay to Surinam, including south-eastern Paraguay, eastern Brazil and French Guiana. Amblyomma auricularium is distributed from northern Patagonia in Argentina throughout the Neotropics into the Nearctic region up to the southern USA (Texas, Florida), with collection localities also in Belize, Bolivia, Brazil, Colombia, Costa Rica, French Guiana, Honduras, Mexico, Nicaragua, Panama, Paraguay, Trinidad & Tobago, Uruguay and Venezuela. It is not known whether A. auricularium is an established resident of the USA.

Ixodes schulzei Aragão and Fonseca was described from Brazil in 1951 based on female ticks collected on wild rats from the states of Rio de Janeiro and Santa Catarina. Since that time, there have been no additional reports of I. schulzei in the literature. We report two new records of I. schulzei: a female collected on the water rat Nectomys squamipes (Brants) from Minas Gerais State, and another female from this same host species from São Paulo State. This last specimen was engorged and oviposited fertile eggs in the laboratory. Larvae hatched from these eggs were used for subsequent infestations under laboratory conditions, as were nymphs obtained from the engorged larvae. Naive laboratory rats (Rattus norvegicus Berkenhout) and wild mice (Calomys callosus Rengger) were used to feed ticks. C. callosus was a more suitable host than R. norvegicus, as significantly more ticks (P < 0.05) were recovered and successfully molted after feeding on the former host species. A significantly (P < 0.05) greater proportion of larvae detached from C. callosus during daylight (71.3%) when compared with those that detached from R. norvegicus in the same period (54.8%). A total of nine engorged nymphs (one from R. norvegicus, and eight from C. callosus) were recovered in the infestations. All of them successfully molted to adults, which were all females. The male of I. schulzei remains unknown.

During an ecological study, carried out between 1994 and 1996 with Streptoprocne biscutata (Sclater) (Apodiformes: Apodidae) birds, that inhabit caves in the Quatro Barras County, State of Paran , Southern Brazil, a new tick species of the subgenus Multidentatus was observed. The female, male, nymph, and larva of Ixodes (Multidentatus) paranaensis n. sp., are described. Of the 12 known species of the subgenus Multidentatus, only I. (M.) auritulus Neumann, 1904 and I. (M.) murreleti Cooley and Kohls, 1945 occur in the Neartic region and only I. (M.) auritulus occurs in the Neotropical region. As such, I. (M.) paranaensis n. sp. increases the number of species and the distribution area of the subgenus Multidentatus in the Americas.

The male and female of Amblyomma geochelone n. sp. are described and illustrated by both scanning electron micrographs and line drawings. Specimens of this new tick species were recovered from the endangered ploughshare tortoise, Geochelone yniphora (Vaillant), in northwestern Madagascar. This relatively large tick is morphologically most similar to Amblyomma nuttalli Dönitz, which occurs in mainland sub-Saharan Africa where it mainly parasitizes other species of tortoises. However, several characters distinguish the new species from A. nuttalli including the scutal ornamentation in both sexes and the characteristic patterns of shallow grooves on the alloscutum of the female of A. geochelone. Because the adult stages of A. geochelone are almost certainly host specific ectoparasites of the ploughsh are tortoise, this new tick species is also probably endangered.

The world's argasid tick fauna comprises 183 species in four genera, namely Argas, Carios, Ornithodoros and Otobius in the family Argasidae. The ixodid tick fauna consists of 241 species in the genus Ixodes and 442 species in the genera Amblyomma, Anomalohimalaya, Bothriocroton, Cosmiomma, Dermacentor, Haemaphysalis, Hyalomma, Margaropus, Nosomma, Rhipicentor and Rhipicephalus in the family Ixodidae, with the genus Boophilus becoming a subgenus of the genus Rhipicephalus. The family Nuttalliellidae is represented by the monospecific genus Nuttalliella. The species names of these ticks, based on seven previous complete or partial listings, as well as those of recently described new species, are presented in tabular format.

Phylogenetic studies of ticks have been increasing in recent years, particularly in the use of molecular data. However, all of the studies to date are either limited to the mitochondrial genome or to a few nuclear ribosomal genes. There is a need to explore the use of nuclear protein-encoding genes because these genes direct most aspects of the phenotypic traits in the development of an organism. We report here the test of a nuclear protein-encoding gene, RNA polymerase II, for the phylogenetic study of ticks. Thirty-eight ticks representing 26 species of hard and soft ticks were chosen for the study. The pairwise divergences among sampled species are ranged from 0.3 to 15.2% and most of the substitutions are transitions. In addition, the nucleotide composition is not obviously biased in POL II gene. The trees inferred from the POL II sequences using maximum parsimony (MP), neighbor joining (NJ), and maximum likelihood (ML) by PAUP* and MrBayes are largely concordant with the existing phylogenies. Our study demonstrated that POL II gene sequences contain strong phylogenetic signals in ticks at the generic and higher levels. POL II has proven to be a useful gene for resolving tick phylogeny.

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.

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.

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.

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.

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.

Systematic relationships among the basal Ixodidae are examined using one morphological and three molecular data sets, 18S and 28S nuclear and 16S mitochondrial rDNA. Although different combinations of partitions are incompatible in a partition homogeneity test, combining them produces similar or better support for most major lineages through both additive and complementary effects. The different data sets are not complete for all taxa, but inclusion or exclusion of taxa with missing data for one or more data sets (8 of 29 ingroup taxa) does not influence overall tree topology and only weakly affects support levels. The only notable effect was based on gap treatment in the 28S data set. Gap treatment completely changes the arrangement and support levels for one basal node. The combined analyses show strong support for the Metastriata, a lineage including most endemic Australian Ixodes, and a lineage including the remaining Ixodes, but not for the Prostriata (=Ixodes s.l.). The distribution pattern of endemic Australian taxa (nearly all included in three exclusively Australian basal lineages) suggests that these lineages, and by extension the Ixodidae, originated after the isolation of Australia in the late Cretaceous, much more recently than previously indicated.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

The authors present an introductory overview of the principal groups of ectoparasites (flukes, leeches, crustaceans, insects, arachnids, lampreys and vampire bats) associated with domestic animals. Currently-accepted higher-level classifications are outlined for these parasites. Almost all significant ectoparasites of domestic animals are invertebrates, the majority being arthropods (crustaceans, insects and arachnids). Some of these ectoparasites are of particular importance as vectors of pathogens. Many ectoparasite species are host-specific, and vector species typically transmit characteristic pathogens.

The larva, nymph, and female of Ixodes (Ixodes) venezuelensis are diagnosed using scanning electron microscopy, and the larva is described for the first time. This ixodid tick is recorded from the northern neotropics as a parasite of five species of didelphid marsupials and eighteen species of rodents (one sciurid, two heteromyids, eleven murids, two dasyproctids, and two echimyids). Collection records of I. venezuelensis are presented for Costa Rica, Colombia, Panama and Venezuela, with the Costa Rican records being new and extending the known range of this tick by at least 300 km. Altitudinally, I. venezuelensis has been collected between elevations of 24 and 2410 m with most collections between 100 and 800 m. The possibility that I. venezuelensis may be parthenogenetic is discussed.

Aponomma (Bothriocroton) glebopalma, n. subgen., n. sp., and Amblyomma glauerti, n. sp. (Acari: Ixodidae), are described from the monitor lizards Varanus glebopalma Mitchell and Varanus glauerti Mertens in Western Australia and the Northern Territory, Australia. The new subgenus is erected to accomodate A. (Bothriocroton) glebopalma, a new species with a deeply pitted and pilose scutum in both the adult and immature stages, unlike all other described species of Aponomma. This new species is found in an area where Aponomma fimbriatum Koch, a parasite of varanid lizards and various species of snakes, was the only known member of the genus found in both Western Australia and the Northern Territory. Amblyomma glauerti n. sp., an ornate Australian reptile tick found on the same two hosts in Western Australia and the Northern Territory, is also described. These two new species were recovered from formalin-preserved lizard specimens; therefore, nothing is known of their life cycles, except that all stages parasitize varanid lizards.

The male of Amblyomma rotundatum is described for the first time, and the nymph and larva are redescribed. This tick is a parthenogenetically reproducing species parasitic on amphibians and reptiles in the warmer areas of the New World. Attempts to mate male and female A. rotundatum under laboratory conditions were unsuccessful.

Haemaphysalis juxtakochi Cooley, 1946, an ectoparasite of deer and other large mammals from Mexico southward through Central and South America, was recovered from the ear of a white-tailed deer, Odocoileus virginanus Zimmermann, in Ohio. This represents the first record of H. juxtakochi from the United States.

The parthenogenetic tick Amblyomma rotundatum, a Central and South American species, has become established in southern Florida. The date of introduction is unknown, but it is suspected to be either during the 1930s, when 1 of its natural hosts, the giant or marine toad, Bufo marinus, was introduced to southern Florida as a potential biological control of pest beetles in sugar cane fields, or between 1955 and 1964 when specimens of B. marinus were accidentally or deliberately released in the greater Miami area. Several museum specimens of this toad collected in the Miami area 25 April 1979 had nymphal and adult A. rotundatum attached. Subsequent examination of living giant toads collected at another Miami area site from 1983 through 1985 revealed larval, nymphal, and adult A. rotundatum and confirmed colonization of this tick. Under laboratory conditions, another neotropical amphibian and reptile tick, Amblyomma dissimile, is capable of transmitting Cowdria ruminantium, the causative agent of heartwater, a disease present in the Caribbean area. Therefore, we suggest that A. rotundatum should also be tested for vectorial competence.

Rhipicephalus exophthalmos sp. nov., a species which has in the past been confused with Rhipicephalus oculatus Neumann, 1901, is described and illustrated from laboratory-reared specimens. Preferred hosts for the adults of this tick are various domestic and wild ungulates and hares. It is widely distributed in Namibia and in the southeastern Cape Province, South Africa, with only scattered records from elsewhere. The adults of R. oculatus are redescribed and illustrated, and the immature stages are described and illustrated for the first time. All stages feed virtually exclusively on leporids. It is common in parts of the eastern and southern Cape Province, with few records as yet from other parts of South Africa and Namibia.

This tick, originally referred to as Rhipicephalus species III by Yeoman & Walker (1967), was first collected from a sitatunga, Tragelaphus spekei, and then occasionally from cattle, in Tanzania. Further collections, mostly from sitatunga, have since been seen from Uganda and Zambia. Descriptions of the adults of this species are given, together with information on their hosts and their distribution.