Comments on controversial tick (Acari: Ixodida) species names and species described or resurrected from 2003 to 2008

There are numerous discrepancies in recent published lists of the ticks of the world. Here we review the controversial names, presenting evidence for or against their validity and excluding some altogether. We also address spelling errors and present a list of 17 species described or resurrected during the years 2003–2008. We consider the following 35 tick species names to be invalid: Argas fischeri Audouin, 1826, Ornithodoros boliviensis Kohls and Clifford, 1964, Ornithodoros steini (Schulze, 1935), Amblyomma acutangulatum Neumann, 1899, Amblyomma arianae Keirans and Garris, 1986, Amblyomma bibroni (Gervais, 1842), Amblyomma colasbelcouri (Santos Dias, 1958), Amblyomma concolor Neumann, 1899, Amblyomma cooperi Nuttall and Warburton, 1908, Amblyomma curruca Schulze, 1936, Amblyomma cyprium Neumann, 1899, Amblyomma decorosum (Koch, 1867), Amblyomma nocens Robinson, 1912, Amblyomma perpunctatum (Packard, 1869), Amblyomma striatum Koch, 1844, Amblyomma superbum Santos Dias, 1953, Amblyomma testudinis (Conil, 1877), Amblyomma trinitatis Turk, 1948, Dermacentor confractus (Schulze 1933), Dermacentor daghestanicus Olenev, 1928, Haemaphysalis himalaya Hoogstraal, 1966, Haemaphysalis vietnamensis Hoogstraal and Wilson, 1966, Hyalomma detritum Schulze, 1919, Ixodes apteridis Maskell, 1897, Ixodes donarthuri Santos Dias, 1980, Ixodes kempi Nuttall, 1913, Ixodes neotomae Cooley, 1944, Ixodes rangtangensis Teng, 1973, Ixodes robertsi Camicas, Hervy, Adam and Morel, 1998, Ixodes serrafreirei Amorim, Gazetta, Bossi and Linhares, 2003, Ixodes tertiarius Scudder, 1885, Ixodes uruguayensis Kohls and Clifford, 1967, Ixodes zealandicus Dumbleton, 1961, Ixodes zumpti Arthur, 1960 and Rhipicephalus camelopardalis Walker and Wiley, 1959. We consider the following 40 names valid: Argas delicatus Neumann, 1910, Argas vulgaris Filippova, 1961, Ornithodoros aragaoi Fonseca, 1960, Ornithodoros dugesi Mazzoti, 1943, Ornithodoros knoxjonesi Jones and Clifford, 1972, Ornithodoros marocanus Velu, 1919, Ornithodoros nattereri Warburton, 1927, Amblyomma beaurepairei Vogelsang and Santos Dias, 1953, Amblyomma crassipes (Neumann, 1901), Amblyomma echidnae Roberts, 1953, Amblyomma fuscum Neumann, 1907, Amblyomma orlovi (Kolonin, 1995), Amblyomma parkeri Fonseca and Arag?o, 1952, Amblyomma pseudoconcolor Arag?o, 1908, Bothriocroton oudemansi (Neumann, 1910), Bothriocroton tachyglossi (Roberts, 1953), Dermacentor abaensis Teng, 1963, Dermacentor confragus (Schulze 1933), Dermacentor ushakovae Filippova and Panova, 1987, Haemaphysalis anomaloceraea Teng, 1984, Haemaphysalis filippovae Bolotin, 1979, Haemaphysalis pavlovskyi Pospelova-Shtrom, 1935, Hyalomma excavatum Koch, 1844, Hyalomma isaaci Sharif, 1928, Hyalomma rufipes Koch, 1844, Hyalomma turanicum Pomerantzev, 1946, Ixodes arabukiensis Arthur, 1959, Ixodes boliviensis Neumann, 1904, Ixodes columnae Takada and Fujita, 1992, Ixodes maslovi Emel′yanova and Kozlovskaya, 1967, Ixodes sachalinensis Filippova, 1971, Ixodes siamensis Kitaoka and Suzuki, 1983, Ixodes sigelos Keirans, Clifford and Corwin, 1976, Ixodes succineus Weidner, 1964, Rhipicephalus aurantiacus Neumann, 1907, Rhipicephalus cliffordi Morel, 1965, Rhipicephalus pilans Schulze, 1935, Rhipicephalus pseudolongus Santos Dias, 1953, Rhipicephalus serranoi Santos Dias, 1950 and Rhipicephalus tetracornus Kitaoka and Suzuki, 1983.     

Ticks (Acari: Ixodidae) parasitizing wild carnivores in Phu Khieo Wildlife Sanctuary, Thailand

Ixodid ticks were collected and identified from 8 wild carnivore species in Phu Khieo Wildlife Sanctuary, northeastern Thailand. Six tick species belonging to 4 genera were recovered and identified from 132 individuals. These included Amblyomma testudinarium (n = 36), Haemaphysalis asiatica (n = 58), H. hystricis (n = 31), H. semermis (n = 3), Rhipicephalus haemaphysaloides (n = 3), and Ixodes granulatus (n = 1). Leopard cats (Prionailurus bengalensis) (n = 19) were infested with 4 tick species, whereas yellow-throated marten (Martes flavigula) (n = 4), clouded leopard (Neofelis nebulosa) (n = 2), and dhole (Cuon alpinus) (n = 1) were infested with 3 tick species, Asiatic golden cat (Catopuma temmincki) (n = 2) with 2 species, and marbled cat (Pardofelis marmorata), binturong (Arctictis binturong), and large Indian civet (Viverra zibetha) each infested with 1 species. This information contributes to the knowledge available on the ectoparasites of wild carnivores in Southeast Asia.     

Ticks (Acari: Ixodidae) on wild animals from the Porto-Primavera Hydroelectric power station area,Brazil

From June 2000 to June 2001, a total of 741 ticks were collected from 51 free-living wild animals captured at the Porto-Primavera Hydroelectric power station area, located alongside an approximately 180 km course of the Paran river, between the states of S o Paulo and Mato Grosso do Sul, comprising 9 species of 3 genera: Ambly-omma (7 species), Boophilus (1) and Anocentor (1). A total of 421 immature Amblyomma ticks were reared in laboratory until the adult stage, allowing identification of the species. A. cajennense was the most frequent tick species (mostly immature stages) collected on 9 host species: Myrmecophaga tridactyla, Tamandua tetradactyla,Cerdocyon thous, Puma concolor,Tayassu tajacu, Mazama gouazoubira,Hydrochaeris hydrochaeris,Alouatta caraya, Cebus apella. Other tick species were less common, generally restricted to certain host taxa.     

Using habitat models to map diversity: pan-African species richness of ticks (Acari: Ixodida)

Aims To show how logistic regression models for individual species can be used to produce improved estimates of species richness at a continental scale; to present these data for African ticks (Acari: Ixodida); and to address the question of whether there is a latitudinal gradient in tick species richness. Location Africa. Methods A database of 34,060 collection records for African ticks is used to produce a pan‐African map of known tick species richness at 0.25 × 0.25‐degree resolution. The likely distributions of seventy‐three species are then estimated from environmental factors using logistic regression, and localities where there is a suitably high probability of occurrence for a given species are added to the original data for that species. These augmented data are combined to produce a map of the predicted pan‐African distribution of tick species richness. The relationship of species richness to latitude is considered along a transect placed across some of the more extensively collected areas. Results Maps of known and predicted pan‐African tick species richness are presented, and deficiencies in the available data are highlighted. Correlations using both known and predicted estimates of tick species richness suggest that ticks follow similar species richness patterns to those described for African mammals and birds, with a latitudinal gradient and highest species richness in east equatorial Africa. Tick species ranges are log‐normally distributed. Main conclusions Carefully constructed probability surfaces offer a more powerful approach to mapping species ranges than simple presence‐absence maps. Such models are a useful extension to current biogeographical methods and have a wide range of potential applications in ecology, epidemiology and conservation. Tick species richness at a continental scale follows similar trends to those reported for mammals and birds.     

Ticks (Ixodidae) on humans in South America

Twenty eight species of Ixodidae have been found on man in South America (21 Amblyomma, 1 Boophilus, 2 Dermacentor, 2 Haemaphysalis, 1 Ixodes and 1 Rhipicephalus species). Most of them are rarely found on man. However, three species frequently parasitize humans in restricted areas of Argentina (A. neumanni reported from 46 localities), Uruguay (A. triste from 21 sites) and Argentina–Brazil (A. parvum from 27 localities). The most widespread ticks are A. cajennense (134 localities in Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Paraguay, Suriname and Venezuela), A. ovale (37 localities in Argentina, Brazil, Ecuador, French Guiana, Guyana, Paraguay, Suriname and Venezuela) and A. oblongoguttatum (28 sites in Brazil, Colombia, French Guiana, Guyana, Suriname and Venezuela). Amblyomma aureolatum (18 localities in Argentina, Brazil, French Guiana and Paraguay), A. cajennense, and A. triste are vectors of rickettsioses to man in South America. A better understanding of the respective roles of these and other tick species in transmitting pathogens to humans will require further local investigations. Amblyomma ticks should be the main subjects of these studies followed by species of Boophilus, Dermacentor, Haemaphysalis and Rhipicephalus species. In contrast with North America, Europe and Asia, ticks of the genus Ixodes do not appear to be major players in transmitting diseases to human. Indeed, there is only one record of an Ixodes collected while feeding on man for all South America.     

Ticks (Acari: Ixodidae) of the state of Amazonas,Brazil

The tick fauna of Brazil is currently composed by 72 species. The state of Amazonas is the largest of Brazil, with an area of ≈ 19% of the Brazilian land. Besides its vast geographic area, only 19 tick species have been reported for Amazonas. Herein, lots containing ticks from the state of Amazonas were examined in three major tick collections from Brazil. A total of 5933 tick specimens were examined and recorded, comprising 2693 males, 1247 females, 1509 nymphs, and 484 larvae. These ticks were identified into the following 22 species: Amblyomma cajennense sensu lato, Amblyomma calcaratum, Amblyomma coelebs, Amblyomma dissimile, Amblyomma dubitatum, Amblyomma geayi, Amblyomma goeldii, Amblyomma humerale, Amblyomma latepunctatun, Amblyomma longirostre, Amblyomma naponense, Amblyomma oblongoguttatum, Amblyomma ovale, Amblyomma rotundatum, Amblyomma scalpturatum, Amblyomma varium, Dermacentor nitens, Haemaphysalis juxtakochi, Ixodes cf. Ixodes fuscipes, Ixodes luciae, Rhipicephalus microplus, Rhipicephalus sanguineus sensu lato. Ticks were collected from 17 (27.4%) out of the 62 municipalities that currently compose the state of Amazonas. The following four species are reported for the first time in the state of Amazonas: A. coelebs, A. dubitatum, H. juxtakochi, and Ixodes cf. I. fuscipes. The only tick species previously reported for Amazonas and not found in the present study is Amblyomma parvum. This study provides a great expansion of geographical and host records of ticks for the state of Amazonas, which is now considered to have a tick fauna composed by 23 species. It is noteworthy that we report 1391 Amblyomma nymphs that were identified to 13 different species.     

Ticks (Acari: Ixodidae) parasitizing humans in an Atlantic rainforest reserve of Southeastern Brazil with notes on host suitability

While conducting projects on ticks from deer and on tick ecology in animal trails in an Atlantic rainforest reserve in Southeastern Brazil, researchers of our group were bitten by ticks several times. Some of these episodes were recorded. Three species of adult ticks attached to humans: Amblyomma brasiliense Aragão, Amblyomma incisum Neumann, and Amblyomma ovale Koch. Eight nymphal attachments with engorgement on humans were recorded. From these, six molted to adults of A. incisum, one to an adult of A. brasiliense, and one had an anomalous molting, therefore the adult tick could not be properly identified. Local reactions to tick attachment varied among individual hosts from almost imperceptible to intense. Especially itching, but hyperemia and swelling as well, were prominent features of the reaction. Overall it can be affirmed that human beings can be a physiologically suitable host species for ticks in the Atlantic rainforest and that itching was an important if not the major component of the resistance to tick bite.     

Parthenogenesis in Mites and Ticks (Arachnida: Acari)

SYNOPSIS. Parthenogenesis is discussed according to type (arrhenotoky,thelytoky, deuteroLoky, artificial parthenogenesis, gynogenesis)and presence in various acarine taxa. Evidence for parthenogenesisis based on rearing and/or cytological studies. Arrhenotokyoccurs in Mesostigmata, Prostigmata and Astigmata and usuallydoes not occur sporadically throughout these taxa; when presentit operates in many closely related species as the major typeof reproduction in certain genera, subfamilies or families.Thelytoky is present in Mesostigmata, Metastigmata, Prostigmata,Astigmata and Cryptostigmata, but in no case is the major modeof reproduction in higher taxa, i.e. subfamilies, families.Its sporadic occurrence may or may not be accompanied by polyploidy.Deuterotoky occurs in one acarine group (Listrophoridae) andartificial parthenogenesis is possible in ticks (Metastigmata).Gynogenesis is present in several species of haplo-diploid Mesostigmata(some Dermanyssidae, Phytoseiidae), although it is usually consideredin connection with thelytoky. Current studies of the effectsof ploidy level on morphological variation indicate greatercharacter variability in diploid females than in haploid males.In some species of ticks thelytoky is expressed differentiallyeven on an intraspecific level. One geographic strain of a speciesmay have the ability to reproduce thelytokously, but if matedproduces males and females.     

Ticks (Acari: Ixodida) infesting humans in the provinces of Kelkit Valley, a Crimean-congo hemorrhagic fever endemic region in Turkey

Ticks are mandatory blood feeding ectoparasites leading transmission of various tick-borne pathogens to human and animals. Since 2002, thousands of human tick bites and numerous Crimean-Congo Hemorrhagic Fever cases have been reported in several provinces in the Kelkit Valley region in Turkey. Despite increased cases of tick bites and tick-borne diseases, no taxonomic information is available about the tick species infesting humans in the region. In the present study, a tick survey on humans was performed to determine the species composition of ticks infesting humans in several provinces of Kelkit Valley. In the survey, 1,460 ticks (721 males, 516 females and 223 nymphs) were collected from tick-infested humans. A total of 19 tick species have been found on humans in the region, including 7 Hyalomma, 2 Argas, 2 Haemaphysalis, 2 Ixodes, Dermacentor and 3 Rhipicephalus species. Infestation of Dermacentor reticulatus on humans was documented for the first time in Turkey.     

Ticks on captive and free-living wild animals in northeastern Brazil

From 2005 to 2009, 147 ticks found on 32 wild animals from or referred to two zoobotanical parks (Parque Zoobotânico Arruda Câmara and Parque Estadual Dois Irmãos) located in northeastern Brazil were identified. Ticks found on two veterinarians working in one of the parks (i.e., Parque Estadual Dois Irmãos), after return from forested areas within the park’s territory, were also identified. The following tick-host associations were recorded: Amblyomma fuscum Neumann on Boa constrictor L.; Amblyomma longirostre Koch on Ramphastos vitellinus ariel Vigors and Coendou prehensilis (L.); Amblyomma varium Koch on Bradypus variegates Schinz; Amblyomma rotundatum Koch on Chelonoidis carbonaria (Spix), Chelonoidis denticulata (L.), Micrurus ibiboboca (Merrem), Python molurus bivittatus Kuhl, Iguana iguana (L.) and B. variegatus; Amblyomma nodosum Neumann on Myrmecophaga tridactyla L. and Tamandua tetradactyla (L.); and Rhipicephalus sanguineus (Latreille) on Nasua nasua (L.). The ticks found on the veterinarians were identified as nine Amblyomma larvae. The presence of Am. nodosum in Pernambuco and Am. rotundatum and Am. varium in Paraíba is recorded for the first time and the occurrence of Am. longirostre in Pernambuco is confirmed. Ramphastos vitellinus ariel is a new host record for Am. longirostre whereas M. ibiboboca and B. variegatus are new host records for Am. rotundatum. Finally, the human parasitism by Amblyomma ticks is reported for the first time in Pernambuco, highlighting the potential of tick-borne pathogen transmission in this state.     

Fruit consumption by carnivores in Mediterranean Europe

• 1 We reviewed patterns of fruit consumption amongst 10 species of mesocarnivores: red fox Vulpes vulpes, weasel Mustela nivalis, stoat Mustela erminea, polecat Mustela putorius, stone marten Martes foina, pine marten Martes martes, Eurasian badger Meles meles, common genet Genetta genetta, Egyptian mongoose Herpestes ichneumon and wildcat Felis silvestris in Mediterranean Europe.
• 2 The 65 reviewed studies recorded 79 different fruits eaten by carnivores, 58 of which were identified to species. Most records (63%) were of fleshy fruits with high pulp content. The frequency of occurrence of fruit items varied widely amongst species and regions. Four of the carnivore species (red fox, stone marten, badger and common genet) included more than 30 fruit species in their diet.
• 3 A longitudinal pattern was detected in the consumption of fruit in the Mediterranean region, with the frequency of occurrence of fruit consumption increasing towards the east.

     

Evolution of skull shape in carnivores 1. Representative modern carnivores

Fifteen variables, selected primarily to reflect functionally significant aspects of cranial morphology, were measured on one skull each of 62 species of modern carnivores, including viverrids, canids, mustelids and felids. To allow comparisons between species of different sizes without the potentially confounding effects of allometric shape changes, the measurements were transformed to dimensionless variables, based on the residuals from allometric equations. Fourteen out of 15 of the transformed variables distinguish one or more of the four family groups and the rotated first two axes of a principal components analysis distinguish all four families from each other. The following functional hypotheses are proposed: mustelids and felids have the most powerful bites and canids the weakest among the four family groups studied; mustelids and, to a lesser degree, felids have more powerful neck musculature than do canids and viverrids; and visual abilities are best developed among felids and least developed among mustelids. The first two functional hypotheses suggest possible differences in killing behaviour, which are supported by a preliminary survey of the literature on such behaviour. Allometric analysis of the 15 cranial measures shows that the neurocranial components scale with negative allometry, while most of the other measures scale approximately isometrically.     

Ticks (Acari: Ixodidae) within various phytophysiognomies of a Cerrado reserve in Uberlandia, Minas Gerais, Brazil

Cerrado biome, the South American savannah, covers about 2 million km² and is very rich in endemic species but threatened by agriculture. In this report free-living tick species are presented, and their seasonal and relative distribution within the various phytophysiognomies in a small Cerrado reserve in Minas Gerais State, Brazil. Overall 2,694 free-living ticks were found during a 2 years sampling period with CO₂ traps and cloth dragging. Of these, 73.5% were Amblyomma cajennense and 0.6% Amblyomma dubitatum. All other ticks (25.9%) were retained as Amblyomma spp. Adults of A. cajennense peaked in spring, the nymphs in winter of both years. Amblyomma larval clusters were found in autumn and winter. Adult ticks (46.7%) and nymphs (39.5%) were most often found in woodlands, whereas most larval clusters were found in valley-side marshes (39%). Amblyomma cajennense, Anocentor nitens, Rhipicephalus (Boophilus) microplusand Rhipicephalus sanguineus ticks were found on domestic animals from neighboring properties. Search for Rickettsia in the hemolymph of 497 A. cajennense and one A. dubitatum ticks yielded negative results. Results confirmed earlier reports on the overwhelming prevalence of A. cajennense ticks in the Cerrado biome of Brazil and added information to habitat preferences of this tick species, a major vector in Brazil of the Rocky Mountain spotted fever.     

Phylogeny, Evolution and Historical Zoogeography of Ticks: A Review of Recent Progress

There has been much progress in our understanding of the phylogeny and evolution of ticks, particularly hard ticks, in the past 5 years. Indeed, a consensus about the phylogeny of the hard ticks has emerged. Our current working hypothesis for the phylogeny of ticks is quite different to the working hypothesis of 5 years ago. So that the classification reflects our knowledge of ticks, several changes to the nomenclature of ticks are imminent. One subfamily, the Hyalomminae, will probably be sunk, yet another, the Bothriocrotoninae n. subfamily, will be created. Bothriocrotoninae n. subfamily, and Bothriocroton n. genus, are being created to house an early-diverging (‘basal’) lineage of endemic Australian ticks that used to be in the genus Aponomma (ticks of reptiles). There has been progress in our understanding of the subfamily Rhipicephalinae. The genus Rhipicephalus is almost certainly paraphyletic with respect to the genus Boophilus. Thus, the genus Boophilus will probably become a subgenus of Rhipicephalus. This change to the nomenclature, unlike other options, will keep the name Boophilus in common usage. Rhipicephalus (Boophilus) microplus may still called B. microplus, and Rhipicephalus (Boophilus) annulatus may still be called B. annulatus, but the nomenclature will have been changed to reflect our knowledge of the phylogeny and evolution of these ticks. New insights into the historical zoogeography of ticks will also be presented. This revised version was published online in July 2006 with corrections to the Cover Date.