Clarification of the Rhipicephalus sanguineus group (Acari,Ixodoidea, Ixodidae). II. R. sanguineus (Latreille, 1806) and related species

In this, our second study on the biosystematics of the Rhipicephalus sanguineus group, we consider an additional five species found in the Afrotropical region, namely R. sanguineus, R. camicasi, R. guilhoni, R. moucheti and R. bergeoni.Cross-breeding experiments have shown that R. sanguineus (Latreille, 1806) sensu stricto is genetically distinct from R. camicasi Morel, Mouchet & Rodhain, 1976. Further cross-breeding trials confirmed that R. camicasi also occurs in Saudi Arabia: a field strain from this country was more productive than a laboratory-reared strain from Egypt. R. sanguineus, which parasitises mainly dogs, probably occurs throughout the Afrotropical region. R. camicasi parasitises both large and small domestic ruminants and is restricted to the arid and semi-arid lowlands of northeastern Africa and the Arabian peninsula. The larva, nymph, male and female of these two last named species are described with the aid of scanning electron micrographs (SEMs).The two western African species, R. guilhoni Morel & Vassiliades, 1963 and R. moucheti Morel, 1964, are accepted as valid members of the R. sanguineus group, but we have little material available for critical study. The male and female of R. guilhoni are briefly redescribed with the aid of SEMs. R. bergeoni Morel & Balis, 1976 occurs predominantly on cattle in the wetter highlands and subhighlands of Ethiopia and Sudan. The male and female are briefly described. Current morphological and ecological data suggest that the inclusion of this species in the R. sanguineus group is questionable.     

Ticks (Acari:Ixodidae) of the Blue and White Nile ecosystems in the Sudan with particular reference to theRhipicephalus sanguineus group

Twenty-four adult ixodid tick species, infesting livestock and some wildlife hosts along the Blue and White Nile in the Sudan, were identified. Three species,Boophilus geigyi, Rhipicephalus camicasi andR. bergeoni, were recorded for the first time from the Sudan. Tick numbers on indigenous breeds of cattle (Bos indicus) were relatively low, ranging between 17.1 and 40.5 per animal. Young cattle grazing with the herd carried significantly fewer ticks than older animals. With the exception ofB. annulatus andR. simus, which have extended their distribution further north into Blue Nile, Gezira and Khartoum Provinces, the distribution patterns of the most important cattle ticks have been relatively unchanged over the past 30 years.TheRhicicephalus sanguineus group was represented by six species.R. camicasi was the only species present on cattle, sheep and goats in the north in Kassala and Khartoum Provinces, whereas this species occurred sympatrically withR. guilhoni andR. turanicus further south in Gezira and Blue Nile Provinces. In the Southern Region of the Sudan onlyR. turanicus andR. guilhoni were present, the latter being by far the predominant species, with peak activity towards the end of the rains in the Jonglei Canal Area.R. bergeoni was collected once, from cattle near the Ethiopian border in Blue Nile Province, whereasR. sanguineus sensu stricto was collected throughout the study area, from domestic dogs only. Finally,R. sulcatus was found once on a hare.The distributions of the common tick species are correlated with the occurrence of tick-borne diseases of domestic animals and recommendations for the control of tick-borne diseases and their vectors in the Sudan are given.     

Ticks Infesting Livestock on Farms in Western Sicily, Italy

The results of a 2-year survey on ticks infesting livestock on farms in Western Sicily, Italy, are reported. Several dogs living in the same area were also examined for ticks. Moreover, free-living ticks from one cattle farm were caught by CO₂ traps and flagging. A total of 6208 specimens was collected belonging to 9 species: Rhipicephalus bursa (32.4%), Rhipicephalus turanicus (22.7%), Rhipicephalus sanguineus (19.3%), Hyalomma lusitanicum (12.0%), Haemaphysalis punctata (7.8%), Hyalomma marginatum (3.5%), Dermacentor marginatus (1.4%), Ixodes ricinus (0.8%) and Ixodes hexagonus (0.1%). The species belonging to the genus Rhipicephalus were present on all hosts and habitats monitored, R. bursa was found to be abundant on cattle (46.3%) and sheep (62.0%), R. sanguineus was prevalent on dogs (50.4%), and R. turanicus on donkeys (41.0%) and as free-living species (48.5%).     

Molecular dissimilarities of <Emphasis Type="Italic">Rhipicephalus sanguineus</Emphasis> (Acari: Ixodidae) in Brazil and its relation with samples throughout the world: is there a geographical pattern?

In this study the genetic variability of Rhipicephalus sanguineus within Brazil and its relation with ticks of the same group from different continents was evaluated. Mitochondrial 12S and 16S rDNA fragments of R. sanguineus from seven Brazilian States were sequenced and compared to GenBank sequences of R. sanguineus and R. turanicus ticks from Africa, Asia, Europe, South America and USA. Results indicate a relatively high intra-specific variability between Brazilian samples but also a global latitude linked distribution pattern of at least two major R. sanguineus groups; one group distributed from latitude 25°N to 22°S including R. sanguineus from Brazil, Taiwan and Thailand and R. turanicus from Zambia and Zimbabwe, and the other group found closer to the poles, roughly above 29°N and below 30°S with ticks from Argentina, Uruguay, France, Oklahoma (USA), Israel and Egypt.     

Observations on the ixodid ticks associated with domestic ungulate animals in Jordan

Abstract

Fourteen species and subspecies of ixodid ticks, representing four genera, were collected from native sheep, goats, camels and cattle in Jordan. Haemaphysalis otophila was the most common species (48.1%) with Rhipicephalus sanguineus (15.3%), Boophilus kohlsi (14.4%) and Haemaphysalis sulcatus (11.6%) representing other common species among the 9,545 specimens examined.     

Rhipicephalus rossicus,a neglected tick at the margin of Europe: a review of its distribution,ecology and medical importance

Rhipicephalus rossicus (Ixodida: Ixodidae) is a three‐host tick with a broad host spectrum that includes wild animals, pets, livestock and humans. Despite its local abundance in certain areas, most of the available information on R. rossicus was published decades ago, mainly by former soviet authors. Its distribution largely overlaps the Eurasian steppe. However, its range may be more extensive than is currently known because this species may have been misidentified as Rhipicephalus sanguineus, principally in areas where the latter species is present. Although R. rossicus has been occasionally reported to feed on people, little attention has been given to its medical importance. It has been shown to have a vectorial role in the transmission of Francisella tularensis, Crimean–Congo haemorrhagic fever virus and West Nile virus. However, the vectorial importance of R. rossicus may be significantly greater, mainly as the closely related species R. sanguineus s.l. is known to transmit a very wide spectrum of pathogens. The probably underestimated vectorial role of R. rossicus may represent a hidden public health threat.     

Ticks Parasitizing Wild Birds in Portugal: Detection of Rickettsia aeschlimannii, R. helvetica and R. massiliae

From January 2002 to December 2004, 152 ticks were collected from 40 wild birds recovered in Santo André Natural Reserve and Monsanto Forestal Park, Portugal mainland. Five ticks species were identified from 22 species of birds, and new host record were provided for some species. In addition, 32 (21%) ticks were screened by PCR to detect infections with agents belonging to order Rickettsiales: Anaplasma phagocytophilum, Ehrlichia chaffeensis, and Rickettsia spp. PCR amplicons were obtained in 5 (15.6%) tick samples. Rickettsia DNA exhibiting gltA sequences similar to those of Rickettsia aeschilimannii, R. helvetica and R. massiliae were identified in Hyalomma marginatum, Ixodes ventalloi and in Rhipicephalus turanicus, respectively. This is the first report of rickettsiae infections in ticks collected from wild birds in Portugal. Giving the results presented above wild birds play an important role in the maintenance and dissemination of several tick species and associated rickettsiae.     

Clarification of the status of Rhipicephalus tricuspis Dönitz, 1906 and Rhipicephalus lunulatus Neumann, 1907 (Ixodoidea,Ixodidae)

The systematic status of Rhipicephalus species whose males have tricuspid adanal plates has been confused for many years. Some authors have regarded Rhipicephalus tricuspis Dönitz, 1906 as the only valid entity with this morphological character and synonymized both Rhipicephalus lunulatus Neumann, 1907 and Rhipicephalus glyphis Dönitz, 1910 with it. Others, however, have always maintained that R. tricuspis and R. lunulatus (syn. R. glyphis) are separate species. Detailed comparative studies, including scanning electron microscopy, of laboratory-reared series as well as numerous field collections of these ticks have now confirmed that the latter view is correct. In addition, a third species, designated here as a Rhipicephalus sp. near tricuspis, has been identified as a member of this group.All stages of R. tricuspis and R. lunulatus are herein redescribed and illustrated by means of scanning electron micrographs. Their life cycles in the laboratory, hosts, distribution and disease relationships are discussed and their differentiation is described with the aid of line drawings. Rhipicephalus tricuspis has been recorded primarily in southern Africa, but also in Zambia and western Zaire, in various types of dry woodland. Its adults are most commonly parasitic on relatively small mammals such as hares, spring hares, jackals and small antelopes. R. lunulatus is much more widespread in the Afrotropical region, most commonly in different types of woodland but also in a variety of other habitats. Its adults parasitize a very wide range of hosts including domestic animals (especially cattle and dogs), the African buffalo, many different antelopes (especially the larger species) and wild pigs. The Rhipicephalus sp. near tricuspis occurs in eastern and parts of central Africa, where its distribution often overlaps with that of R. lunulatus.     

Synergism of thymol,carvacrol and eugenol in larvae of the cattle tick,Rhipicephalus microplus,and brown dog tick,Rhipicephalus sanguineus

The effects of combinations of the monoterpenes thymol and carvacrol and the phenylpropanoid eugenol in larvae of Rhipicephalus microplus (Canestrini, 1888) (Acari: Ixodidae) and Rhipicephalus sanguineus sensu lato (s.l.) (Acari: Ixodidae) were assessed by the larval packet test. The CompuSyn program was used to make qualitative assessments of the effects (synergistic, additive and antagonistic) of the associations. The effects of all combinations tested against R. microplus larvae were synergistic, with combination indices (CIs) <0.70. When tested against R. sanguineus, eight of the mixtures showed a synergistic effect (CI < 0.70); only the carvacrol + thymol mixture at LC₅₀ presented a moderate synergistic effect, with CIs between 0.70–0.90. This study is the first to determine the effects of the interactions of these substances in the control of these two tick species. The combinations of carvacrol + thymol, carvacrol + eugenol and thymol + eugenol have synergistic effects in R. microplus and R. sanguineus s.l. larvae.     

Morphological identification of ticks (Acari: Ixodidae) infesting donkeys (Equus asinus) in Maputo Province,Mozambique

Donkeys (Equus asinus) are rustic animals, but in Africa’s poorest regions, they can present multiple health problems, including tick infestation. The study’s objective was to determine the species composition of ticks that infest donkeys in Maputo Province (Mozambique). Ticks were collected in five conveniently selected southern districts of Maputo Province (Moamba, Matutuíne, Marracuene, Boane, and Matola) and were identified to species level using a stereoscopic microscope with the aid of dichotomous identification keys. In total, 500 ticks were collected from all 88 selected donkeys. Three genera of ticks were identified, namely Rhipicephalus (97.2%; 486/500), Amblyomma (2.2%; 11/500), and Hyalomma (0.6%; 3/500). Seven species were identified, of which Rhipicephalus evertsi evertsi with 50.4% (252/500) was the most prevalent, followed by Rhipicephalus appendiculatus (27.4%; 137/500), Rhipicephalus turanicus (11.6; 10/500), Rhipicephalus (boophilus) microplus (6.8; 20/500), Amblyomma hebraeum (2.2%; 11/500), Rhipicephalus sanguineus (1%; 5/500) and Hyalomma truncatum (0.6%; 3/500). Rhipicephalus evertsi evertsi occurred in all locations, whereas Hyalomma truncatum occurred only in the Boane district. Males were the most prevalent (67.2%; 336/500). The study revealed that donkeys in Maputo Province were infested with seven tick species of which R. evertsi evertsi was the main species.

     

Mitochondrial DNA analysis of Rhipicephalus sanguineus s.l. from the western Iberian peninsula

Rhipicephalus sanguineus Latreille (1806) (Ixodida: Ixodidae) is considered to be the most widely distributed tick and to have a vast range of habitats and hosts, including livestock, pets and wildlife. In addition to morphological differences, recent investigations using approaches based on molecular genetic markers have revealed the existence of different R. sanguineus lineages in different geographic regions. In this study, 475 ticks collected from dogs in the western Iberian peninsula were studied both morphologically and genetically, using 12S and 16S rDNA and COI gene markers in order to clarify the controversy over the systematic status of R. sanguineus sensu lato in Western Europe, and to compare the present data with those sourced from studies conducted in other regions of the world. Despite the high morphometric variability, particularly on spiracles in both genders and in female genitalia, data obtained with different genetic molecular markers show very low variability, suggesting the existence of a unique species. In addition, the phylogenetic analysis showed genetic uniformity, supporting the existence of a well‐defined clade consisting of R. sanguineus s.l. specimens from Western Europe that are distinct from R. sanguineus s.l. from Africa. Furthermore, these data corroborate the existence of a polymorphic species in Western Europe, which requires to be consensually redescribed in view of its medical and veterinary importance in pathogen transmission.     

Genetic Diversity of Hard Ticks (Acari: Ixodidae) in the South and East Regions of Kazakhstan and Northwestern China

To date, there is no report on the genetic diversity of ticks in these regions. A total of 370 representative ticks from the south and east regions of Kazakhstan (SERK) and Xinjiang Uygur Autonomous Region (XUAR) were selected for molecular comparison. A fragment of the mitochondrial cytochrome c oxidase subunit I (cox1) gene, ranging from 631 bp to 889 bp, was used to analyze genetic diversity among these ticks. Phylogenetic analyses indicated 7 tick species including Hyalomma asiaticum, Hyalomma detritum, Hyalomma anatolicum, Dermacentor marginatus, Rhipicephalus sanguineus, Rhipicephalus turanicus and Haemaphysalis erinacei from the SERK clustered together with conspecific ticks from the XUAR. The network diagram of haplotypes showed that i) Hy. asiaticum from Almaty and Kyzylorda Oblasts together with that from Yuli County of XUAR constituted haplogroup H-2, and the lineage from Chimkent City of South Kazakhstan was newly evolved; and ii) the R. turanicus ticks sampled in Israel, Almaty, South Kazakhstan, Usu City, Ulugqat and Baicheng Counties of XUAR were derivated from an old lineage in Alataw City of XUAR. These findings indicate that: i) Hy. asiaticum, R. turanicus and Ha. erinacei shared genetic similarities between the SERK and XUAR; and ii) Hy. marginatum and D. reticulatus show differences in their evolution.     

Spotted Fever Group <Emphasis Type="Italic">Rickettsiae</Emphasis> in Ticks in Cyprus

In two surveys conducted from March 1999 to March 2001 and from January 2004 to December 2006, a total of 3,950 ticks (belonging to ten different species) were collected from seven domestic and wild animals (goat, sheep, cattle, dog, fox, hare, and mouflon) from different localities throughout Cyprus. In order to establish their infection rate with Spotted Fever Rickettsiae (SFG), ticks were pooled and tested by polymerase chain reaction targeting gltA and ompA genes, followed by sequencing analysis. When tick pools tested positive, individual ticks were then tested one by one, and of the 3,950 ticks screened, rickettsial DNA was identified in 315 ticks (infection rate, 8%). Five SFG Rickettsiae were identified: Rickettsia aeschlimannii in Hyalomma marginatum marginatum, Rickettsia massiliae in Rhipicephalus turanicus and Rhipicephalus sanguineus, Rickettsia sibirica mongolotimonae in Hyalomma anatolicum excavatum, and a Rickettsia endosymbiont of Haemaphysalis sulcata (later described as Rickettsia hoogstraalii) in Haemaphysalis punctata. Two additional genes, 17 kDa and ompB, were targeted to characterize a new genotype of “Candidatus Rickettsia barbariae” genotype in R. turanicus, designated here as “Candidatus Rickettsia barbariae” Cretocypriensis. These results confirm the presence of a spectrum of SFG Rickettsiae on the island. Further studies are necessary to gain better knowledge on the epidemiology of SFG Rickettsiae in Cyprus.     

Comparison of closely related,uncultivated Coxiella tick endosymbiont population genomes reveals clues about the mechanisms of symbiosis

Understanding the symbiotic interaction between Coxiella‐like endosymbionts (CLE) and their tick hosts is challenging due to lack of isolates and difficulties in tick functional assays. Here we sequenced the metagenome of a CLE population from wild Rhipicephalus sanguineus ticks (CRs) and compared it to the previously published genome of its close relative, CLE of R. turanicus (CRt). The tick hosts are closely related sympatric species, and their two endosymbiont genomes are highly similar with only minor differences in gene content. Both genomes encode numerous pseudogenes, consistent with an ongoing genome reduction process. In silico flux balance metabolic analysis (FBA) revealed the excess production of L‐proline for both genomes, indicating a possible proline transport from Coxiella to the tick. Additionally, both CR genomes encode multiple copies of the proline/betaine transporter, proP gene. Modelling additional Coxiellaceae members including other tick CLE, did not identify proline as an excreted metabolite. Although both CRs and CRt genomes encode intact B vitamin synthesis pathway genes, which are presumed to underlay the mechanism of CLE‐tick symbiosis, the FBA analysis indicated no changes for their products. Therefore, this study provides new testable hypotheses for the symbiosis mechanism and a better understanding of CLE genome evolution and diversity.     

Species distribution and detection of Crimean Congo Hemorrhagic Fever Virus (CCHFV) in field-collected ticks in Ankara Province, Central Anatolia, Turkey

Ticks may act as vectors for a number of infectious diseases including Crimean Congo Hemorrhagic Fever (CCHF). The causative agent is Crimean Congo Hemorrhagic Fever Virus (CCHFV), a member of Bunyaviridae, causing extensive ecchymosis, visceral bleeding and hepatic dysfunction with a high fatality rate in the affected individuals. CCHF was initially recognized in Turkey in 2002 and the current number of reported cases exceeds 4,400. This study was conducted to confirm the presence of tick species established as potential CCHFV vectors and investigate CCHFV activity in ticks at Ankara province, Turkey’s second most-densely populated province, where CCHF cases were demonstrated. A total of 1,196 adult ticks, collected from various animals and vegetation in 12 sites located in 5 counties of Ankara during April–July 2010 were identified to species level. Twenty-two tick pools from county K2 were also evaluated for the presence of CCHFV RNA via a one-step real-time RT-PCR assay and reactive results were further confirmed by an in house nested RT-PCR assay. Nine tick species were identified: Rhipicephalus bursa (44.9%), R. sanguineus (18.9%), R. turanicus (18.1%), Haemaphysalis parva (8.3%), Hyalomma marginatum marginatum (5.4%), H. aegyptium (1.4%), H. anatolicum excavatum (1.3%), Hae. punctata (0.3%) and Dermacentor marginatus (0.2%). A total of five tick pools (22.7%) were reactive in real-time and nested RT-PCR assays. The pools included R. bursa, H. m. marginatum and Hae. parva ticks, collected from mammal hosts from two villages in one county. This is the first documentation of CCHFV activity in ticks from Ankara province, which indicates requirement for detailed surveillance to predict high risk zones in the region.     

Rickettsial infections of dogs, horses and ticks in Juiz de Fora, southeastern Brazil, and isolation of Rickettsia rickettsii from Rhipicephalus sanguineus ticks

The present study was performed in an area endemic for Brazilian spotted fever (BSF) in Juiz de Fora, state of Minas Gerais, Brazil, during the years 2007 and 2008, when fatal cases of BSF (caused by Rickettsia rickettsii) were reported. Adult ticks (Acari: Ixodidae) identified as Rhipicephalus sanguineus (Latreille) and Amblyomma cajennense (Fabricius) were collected from dogs and horses, respectively, and tested by polymerase chain reaction (PCR). Overall, 13.1% of the Rh. sanguineus ticks and none of the A. cajennense were found to be infected with R. rickettsii. Two isolates of R. rickettsii were successfully established in Vero cell culture from two Rh. sanguineus ticks. An indirect immunofluorescence assay (IFA) using R. rickettsii antigens detected blood serological reaction to R. rickettsii in 67.9% (53/78) of dogs and 41.0% (16/39) of horses living in the study area. Larval offspring from two Rh. sanguineus engorged females, naturally infected by R. rickettsii, were reared to adult stage in the laboratory. All active stages (larvae, nymphs, adults) remained 100% infected by R. rickettsii, which was efficiently transmitted to naïve rabbits. Overall, the results of the present study indicate a potential risk for transmission of R. rickettsii to humans by Rh. sanguineus, an occurrence yet to be documented in Brazil.     

Seasonal dynamics of the brown dog tick,Rhipicephalus sanguineus,on a confined dog population in Italy

This study evaluated the seasonal dynamics of Rhipicephalus sanguineus (Latreille) (Acari: Ixodidae) on naturally infested dogs in a private shelter in southern Italy. From March to May 2008, 39 autochthonous mixed‐breed young dogs and 10 beagles were enrolled in the study. From March 2008 until March 2009, every 21 ± 2 days, 11 body sites of each dog were checked for ticks. At each follow‐up, the number of ticks, their developmental stage, sex and location on the dog's body were recorded. Adult ticks were found throughout the year, but immatures were absent in January and February. The adult tick population increased from July to August, whereas the load of immatures increased in early July and peaked in September, which suggests that R. sanguineus develops one generation per year in this area. The mean number of immature ticks per infested dog was higher than that of adults from March to October 2008. Ears, interdigital areas and armpits were the most frequent attachment sites of adult ticks. At the last follow‐up, a total of 2266 ticks were collected and identified as R. sanguineus. The results suggest that R. sanguineus develops one generation per year in the study area, but that it infests dogs in all seasons. This information should be taken into account when planning control programmes against this tick species and the pathogens it transmits.     

The seasonal occurrence of ticks (Acari: Ixodidae) on sheep and in the field in the Judean area of Israel

A 2 year survey of ixodid ticks in the Judean area of Israel between 1983 and 1985 showed that sheep were parasitized by the following species:Rhipicephalus sanguineus group (30.6%),Rhipicephalus bursa (25.3%),Haemaphysalis cretica (20.5%),Haemaphysalis otophila (20.1%) andHyalomma anatolicum excavatum (3.1%). Three other species,Hyalomma marginatum rufipes, Boophilus annulatus andAmblyomma lepidum were also present but in very low numbers. A total of 50 816 specimens (78.1% adult ticks) were collected from the sheep. In the field, a total of 4780 ticks (8.7% adults) were collected and 64.3% consisted ofR. bursa. Other species found wereH. cretica (14.5%),H. a. excavatum (13.7%),H. otophila (3.4%),R. sanguineus group (1.9%) and 1.7% wasIxodes eldaricus. The number of ticks collected varied from year to year and seasonal activity was observed in all species except forH. a. excavatum. In all species, males outnumbered females, with the exception ofH. cretica. The seasonal occurrence, tick ecology and disease relationship is discussed for each species.     

Incongruent effects of two isolates of <Emphasis Type="Italic">Rickettsia conorii</Emphasis> on the survival of <Emphasis Type="Italic">Rhipicephalus sanguineus</Emphasis> ticks

Rickettsia conorii, the etiologic agent of Mediterranean spotted fever is widely distributed in Southern Europe, the Middle East, Africa, India and the Caspian region. In the Mediterranean region, the brown dog tick, Rhipicephalus sanguineus, is the recognized vector of R. conorii. To study tick-pathogen relationships and pathogenesis of infection caused in model animals by the bite of an infected tick, we attempted to establish a laboratory colony of Rh. sanguineus persistently infected with R. conorii. Rhipicephalus sanguineus ticks of North American and Mediterranean origin were exposed to R. conorii isolates of African (R. conorii conorii strain Malish) and Mediterranean (R. conorii israelensis strain ISTT) origin. Feeding of ticks upon infected mice and dogs, intra-hemocoel inoculation, and submersion in suspensions of purified rickettsiae were used to introduce the pathogen into uninfected ticks. Feeding success, molting success and the longevity of molted ticks were measured to assess the effects of R. conorii on the survival of Rh. sanguineus. In concordance with previously published results, Rh. sanguineus larvae and nymphs from both North American and Mediterranean colonies exposed to R. conorii conorii Malish experienced high mortality during feeding and molting or immediately after. The prevalence of infection in surviving ticks did not exceed 5%. On the other hand, exposure to ISTT strain had lesser effect on tick survival and resulted in 35–66% prevalence of infection. Rh. sanguineus of Mediterranean origin were more susceptible to infection with either strain of R. conorii than those from North America. Previous experimental studies had demonstrated transovarial and transstadial transmission of R. conorii in Rh. sanguineus; however, our data suggest that different strains of R. conorii may employ different means of maintenance in nature. The vertebrate host may be a more important reservoir than previously thought, or co-feeding transmission between different generations of ticks may obviate or lessen the requirement for transovarial maintenance of R. conorii.     

Off‐host aggregation in the non‐fed,female brown dog tick,Rhipicephalus sanguineus (Latreille), is induced by tick excreta and enhanced by low relative humidity

We report that Rhipicephalus sanguineus (Ixodida: Ixodidae) faeces and its main component, guanine, act as assembly pheromones in short‐range Petri plate bioassays. Arrestment activity in response to guanine was lower than that in response to natural excreta, indicating the presence of other active ingredients in natural excreta. The selective removal of appendages was used to establish the important roles played by the palps and the front pair of legs in the detection of the pheromone. Reaction to chemically pure guanine at varying concentrations occurred without a dose response; thus only the presence of guanine, not a critical amount, is required to induce assembly. Higher speed and intensity of clustering occurred at 33% relative humidity (RH). We conclude that female adults of R. sanguineus are more prone to assemble under dry conditions that match the arid microhabitats preferred by this species and that this tendency allows this tick to reside in human dwellings and dog kennels that maintain standards of comfort at 30–50% RH. Cleaning or removing tick excreta‐covered surfaces on which ticks aggregate from within and around human dwellings may prove useful as a means of interfering with the establishment of off‐host clusters of R. sanguineus.