Systematics and Biogeography of Hard Ticks, a Total Evidence Approach.

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.

Ontogeny of Rhyncoptes grabberi, n. sp. (Acari: Astigmata: Rhyncoptidae) associated with Macaca mulatta.

All ontogenetic stages of a new species of follicle mite, Rhyncoptes grabberi, n. sp., associated with rhesus monkeys, Macaca mulatta, are described and illustrated. The ontogeny of the new species is characterized by numerous retardations and losses of ancestral transformations, especially in the female developmental line. In the male developmental line, many transformations are delayed to the male itself.

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.

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.

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.

Haller's organ in the tick family Argasidae (Acari: Parasitiformes: Ixodida).

The structure of Haller's organ in all major groups of soft ticks, family Argasidae, is reexamined, and hypotheses of setal homology are proposed based on a comparative study of developmental patterns. The larvae of most taxa have 7 setae in the anterior pit (1 serrate, 1 conical, 2 fine, 2 porose, 1 grooved) and 4 porose setae in the capsule. The postlarval stages add up to 3 setae to the anterior pit (1 serrate, 1 porose, 1 grooved) and up to 3 porose setae to the capsule. Preliminary hypotheses of polarity, distinguishing ancestral from derived character states, are generated for most characters examined by comparisons with results obtained for the other major family of ticks, Ixodidae, and (where possible) other members of Parasitiformes. Most of the variability in the setal complement of the anterior pit and capsule results from paedomorphosis, and nearly all the observed changes in the setal complement (both modifications of addition patterns and changes in fundamental numbers) are restricted to single species or small species groups. A correlation between changes in Haller's organ structure and changes in life history patterns is suggested but cannot be confirmed based on presently available data.

Parasitic arthropods of sympatric opossums, cotton rats, and cotton mice from Merritt Island, Florida.

Six species of parasitic arthropods were collected from 12 opossums (Didelphis virginiana), 8 species were obtained from 28 cotton rats (Sigmodon hispidus), and 4 species were collected from 10 cotton mice (Peromyscus gossypinus) from Merritt Island, Brevard County, Florida. The flea Polygenis gwyni was the only parasite that infested all 3 host species, and it was present in very high densities (mean intensity = 73.7) on the opossums. The American dog tick, Dermacentor variabilis, and the tropical rat mite, Ornithonyssus bacoti, both infested 2 host species but were principally associated with opossums and cotton rats, respectively. All remaining arthropod species were confined to single host species, suggesting host specificity by these parasites. The atopomelid mite Didelphilichus serrifer and the myobiid mite Radfordia sigmodontis constitute new state records for Florida. The reptile tick Amblyomma dissimile is recorded from a cotton mouse for the first time; Merritt Island may represent the most northerly location for viable populations of this tick in the U.S.A.

Occurrence and treatment of the mange mite Notoedres muris in marsh rats from South America.

The mange mite Notoedres muris is reported from a new host, the marsh rat (Holochilus brasiliensis) from Argentina. The infection involved alopecia and encrustations on the ears and face, and was treated successfully by a subcutaneous injection of ivermectin. The new record suggests that Notoedres muris has become self-maintaining within this marsh rat population.

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.

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.