Didelphimorphia

Guglielmone, Alberto A., Nava, Santiago & Díaz, Mónica, 2011, Relationships of South American marsupials (Didelphimorphia, Microbiotheria and Paucituberculata) and hard ticks (Acari: Ixodidae) with distribution of four species of Ixodes, Zootaxa 3086, pp. 1-30 : 3-14

Didelphimorphia : Didelphidae Gray (only family for this order)

1) Amblyomma aureolatum ( Pallas, 1772) is a South American tick whose adults are mostly found on Carnivora Bowdich , while most records for subadult ticks are from birds ( Guglielmone et al. 2003c).

Unknown South American country, M on Didelphis sp. ( Guglielmone et al. 2003c).

Brazil, Río Grande do Sul, Pelotas (31º46´S 52º20´W), M on Didelphis albiventris Lund ( Muller et al. 2005) .

Santa Catarina, Area de Proteção Ambiental do Anhatomirim (27º25´S 48º34´W), MF on Didelphis aurita Wied-Neuweid ; Ilha de Ratones Grande (27º29´S 48º 34´W), MF on D. aurita ; Ilha de Santa Catarina (27º43´S 48º31´W), MF on D. aurita ( Salvador et al. 2007).

2) Amblyomma auricularium ( Conil, 1878) is a Neotropical and Nearctic tick species prone to feed on Cingulata Illiger ( Guglielmone et al. 2003a).

Brazil. There is a Brazilian record for A. auricularium on Lutreolina crassicaudata Desmarest but tick stages found on hosts were not determined ( Linardi et al. 1991).

Panama, LU, MF on Didelphis marsupialis Linnaeus; LU, MFNL on Philander opossum (Linnaeus) ( Fairchild et al. 1966).

3) Amblyommma cajennense ( Fabricius, 1787) is a Neotropical and Nearctic tick species with an ample tetrapod host range and locality records (1435 records for the whole Neotropical Region) but the name is considered to represent a tick species group (Lorenza Beati personal communication).

Argentina, Chaco, Riacho El Correntino (26º41´S 59º14´W), N on D. albiventris .

Formosa, El Colorado INTA (26º19´S 59º21´W), N on L. crassicaudata ( Ivancovich & Luciani 1992) .

Brazil, Mato Grosso do Sul, Campo Grande (20º27´S 54º36´W), A on D. albiventris ( Costa et al. 2002).

Rio de Janeiro, Seropédica (22º44´S 43º42´W), N on D. aurita ( Santos Abel et al. 2000) .

Santa Catarina, Ilha de Ratones Grande (27º29´S 48º 34´W), MFN on D. aurita ( Salvador et al. 2007).

Sao Paulo, Caçapava (23º06´S 45º42´W), N on Didelphis sp.; Campinas (22º54´S 47º04´W), N on Didelphis sp. ( Souza et al. 2006), A on D. albiventris ( Souza & Souza, 2008) ; Jaguarina (22º40´S 46º59´W), N on Didelphis sp.; Monte Alegre do Sul (22º39´S 46º40´W), AN on Didelphis sp. ( Souza et al. 2006); Pedreira (22º44´S 46º57´W), AN on D. marsupialis ( Lemos et al. 1997, but the host is probably D. aurita ); Piracicaba (22º43´S 47º38´W), NL on D. albiventris ( Pérez et al. 2008) ; Porto Martin (22º39´S 48º22´W), N on D. aurita ( Aragão 1918) .

Colombia is included in the list of A. cajennense feeding on marsupials, but no tick stages found on hosts are provided by Wells et al. (1981).

Mexico, Tabasco, Frontera (18º32´N 92º38´W), N on Didelphis sp. ( Keirans 1982).

Panama, LU, A on Didelphis sp. (Fairchild et al. 1966) but the host should be D. marsupialis because it is the only species of this genus found in Panama (Wilson & Reeder 2005).

4) Amblyomma coelebs Neumann, 1899 is basically a Neotropical tick species with a few records in the southern Nearctic. Adult ticks are prone to feed on Perissodactyla Owen (Guglielmone et al. 2003b). Brazil, LU, N on D. albiventris (Labruna et al. 2005a).

5) Amblyomma dissimile Koch, 1844 is a Nearctic and Neotropical species usually feeding on Amphibia Linnaeus and Squamata Oppel with sporadic findings on Mammalia Linnaeus (Guglielmone & Nava 2010b).

Brazil, Pernambuco, Reserva Biológica (08º39´S 38º01´W), N on Monodelphis domestica Wagner, N on D. albiventris (Botelho et al. 2002) , but these records were found unconvincing by Guglielmone and Nava (2010b) because the amphibian-squamatan A. dissimile was the only tick found on rodents and marsupials after two months of work and capture of 83 mammals recognized as hosts for several species of ticks.

6) Amblyomma dubitatum Neumann, 1899 is an exclusive South American species. The usual host for all tick stages is the caviid rodent Hydrochoerus hydrochaeris (Linnaeus) (Nava et al. 2010) .

Brazil, São Paulo, Mogi das Cruzes (23º31´S 46º12´W), N on D. albiventris and-or D. aurita (Horta et al. 2007) ; Piracicaba (22º43´S 47º38´W), NL on D. albiventris (Pérez et al. 2008) .

7) Amblyomma fuscum Neumann, 1907 is an exclusive South American species whose adults feed on a variety of hosts while immature stages have been found on rodents and marsupials (Barros-Battesti et al. 2005; Martins et al. 2010).

Brazil, Pernambuco, Paudalho (07º53´S 35º10´W), N on D. aurita and D. albiventris ; São Lourenço da Mata (08º00´S 35º01´W), N on D. albiventris ; Tamandaré (08º45´S 35º06´W), N on D. albiventris (Martins et al. 2010) .

São Paulo, Guarujá (23º59´S 46º16´W), NL on D. aurita (Martins et al. 2010) ; Iguapé (24º42´S 47º33´W), N on D. aurita (Barros-Battesti et al. 2005) ; Serra do Mar State Park (23º52´S 46º26´W), N on D. aurita (Sabatini et al. 2010) .

8) Amblyomma geayi Neumann, 1899 is a Neotropical species non-exclusively South American known mainly from adult ticks feeding on Pilosa Flower : Bradypodidae Gray and Rodentia Bowdich : Erethizontidae Bonaparte (Guglielmone et al. 2003b).

Panama, LU, MF on D. marsupialis, N on Caluromys derbianus (Waterhouse) (Fairchild et al. 1966).

9) Amblyomma humerale Koch, 1844 is a species found only in South America because records for Central America and Nearctic Region are considered misidentifications as discussed in Labruna et al. (2002a), who also state that the usual hosts for adult ticks are Testudines Linnaeus : Testudinidae Gray , while subadult ticks show a more ample host range than adult ticks. Differential diagnoses with Amblyomma crassum Robinson, 1926 and Amblyomma sabanerae Stoll, 1894 is difficult (Guglielmone et al. 2003b).

Brazil, Rondônia, Jamari River (10º17´S 63º14´W), N on D. marsupialis (Labruna et al. 2002a).

10) Amblyomma incisum Neumann, 1906 is an exclusive South American tick whose adults are usually found on Perissodactyla : Tapiridae Gray , redescribed convincingly by Labruna et al. (2005b)

Brazil. There are records of A. incisum on South American marsupials for this country in Barros-Battesti (2008) but tick stages found on hosts are not provided.

11) Amblyomma maculatum Koch, 1844 is a Neotropical-Nearctic tick species very close morphologically to Amblyomma triste Koch, 1844 , that parasitizes an ample range of hosts (Barker et al. 2004; Estrada-Peña et al. 2005).

Venezuela is included in the range of A. maculatum by Díaz-Ungría (1957) who mentions its presence on Didelphimorphia in Venezuela, but the tick stages found on them were not provided.

12) Amblyomma oblongoguttatum Koch, 1844 feeds on different types of hosts from the southern Neartic into the Neotropical Region reaching Bolivia and Brazil (Guglielmone et al. 2003b).

Panama, LU, A on Chironectes minimus (Zimmermann) (Fairchild et al. 1966).

13) Amblyomma ovale Koch, 1844 is a Nearctic and Neotropical species whose adult ticks are usually found on Carnivora Bowdich ( Canidae Fischer and Felidae Fischer de Waldheim), and to a lesser extent on Perissodactyla : Tapiridae apart from several findings on a variety of hosts (Guglielmone et al. 2003c).

Central or South America country, LU, NL on Didelphis sp., N on D. albiventris and Marmosa robinsoni Bangs (Guglielmone et al. 2003c).

Brazil. Barros-Battesti (2008) registers the presence of marsupials infested with A. ovale , but the tick stages found on hosts are not provided.

Colombia. There is a vague record of A. ovale on “marsupials” by López and Parra (1985).

14) Amblyomma pacae Aragão, 1911 is a Neotropical species distributed beyond South America whose usual hosts for adult ticks are Rodentia : Cuniculidae Miller and Gildney , although records on other types of hosts are not unusual (Guzmán Cornejo et al. 2006a).

Brazil, Rondônia, LU, N on D. marsupialis (Labruna et al. 2005a).

15) Amblyomma parvum Aragão, 1908a is a Neotropical species. Adult ticks have an ample host range, but Rodentia : Caviidae Fischer de Waldheim are the usual hosts for their larvae and nymphs (Nava et al. 2008).

Argentina, Formosa, El Colorado INTA (26º19´S 59º21´W), NL on D. albiventris (Ivancovich & Luciani 1992) .

16) Amblyomma pseudoconcolor Aragão, 1908b is a South American species whose usual hosts are Cingulata : Dasypodidae Gray (Guglielmone et al. 2003a).

South America, Guglielmone et al. (2003a) register the parasitism of males of A. pseudoconcolor on Philan- der sp. without any additional data.

Argentina, Formosa, El Colorado (26º18´S 59º23´W), N on D. albiventris (Ivancovich 1973) .

17) Amblyomma romitii Tonelli-Rondelli, 1939 had been considered a junior synonym of A. extraoculatum Neumann, 1899 , but was definitively resurrected by Barros-Battesti et al. (2007). Santos Dias (1955) treat as erroneous the labeling of Singapore for the only known specimen of A. extraoculatum because he regarded A. romittii as a junior synonym of the former, but A. romitii is now considered an exclusive South American tick whereas A. extraoculatum is considered an Oriental species. Most records are from Rodentia : Caviidae .

Venezuela, Bolívar, South-East of Ciudad Bolívar (06º52´N 63º29´W), M on D. marsupialis (Jones et al. 1972, who originally named this tick as A. extraoculatum ).

18) Amblyomma sabanerae Stoll, 1894 is basically a parasite of Testudines with few records on other types of hosts with a Neotropical range from southern Mexico to Colombia. This species is very close morphologically to A. crassum and A. humerale (Fairchild et al. 1966; Guglielmone et al. 2003b).

Panama, Panamá, LU, M on M. robinsoni (Fairchild et al. 1966) .

19) Amblyomma scutatum Neumann, 1899 is a Neotropical species non-exclusively South American that usually feeds on reptiles (Guglielmone et al. 2003b).

Brazil, LU, N on Didelphys (sic) pusilla Desmarest (= Thylamys pusillus (Desmarest) (Neumann 1899). The host in Neumann (1899) is probably wrong because Brazil is out of the range of T. pusillus (Wilson & Reeder 2005). Most probably the host is Thylamys karimii (Petter) (Giarla et al. 2010).

20) Amblyomma tigrinum Koch, 1844 is a Neotropical species with adults feeding mostly on Carnivora : Canidae , nymphs on Rodentia : Caviidae and larvae with a more ample range of hosts (Nava et al. 2006).

Paraguay, Massi Pallarés and Benítez Usher (1982) report infestation of “marsupials” with sub-adults of A. tigrinum . We consider these records doubtful because they were done long before the description of these stages by Estrada-Peña et al. (1993), who stress the difficulties to separate morphologically the larva and nymph of A. tigrinum from the same stages of related species (Estrada-Peña et al. 1993, 2005).

21) Amblyomma triste Koch, 1844 , a Neotropical and Nearctic tick species (Guzmán-Cornejo et al. 2006b) whose larvae and nymphs are mainly collected from Cricetidae Fischer : Sigmodontinae Wagner and Caviidae , but to a much lesser extent on birds (Nava et al. 2011).

Brazil, São Paulo, Pedreira (22º44´S 46º57´W), A on D. marsupialis (Lemos et al. 1997, but the host is probably D. aurita ).

Uruguay, LU, NL on Monodelphis dimidiata (Wagner) (Venzal et al. 2008 a).

22) Amblyomma varium Koch, 1844 is a Neotropical species non-exclusively South American that usually feeds on Pilosa : Bradypodidae-Megalonchydae Ameghino (Onofrio et al. 2008).

Panama, Panamá, Canal Zone nearby Panamá City (08º57´N 79º32´W), F on D. marsupialis (Fairchild et al. 1966) .

23) Haemaphysalis leporispalustris (Packard, 1869) is established in the Nearctic and Neotropical Regions, parasitizing usually Lagomorpha Brandt : Leporidae Fischer , although it was also found on an ample range of hosts. Colombia, Bolívar, Socorro (09º14´N 74º25´W), N on Metachirus nudicaudatus (Geoffroy) (Kohls 1960) . Costa Rica. Campbell et al. (1979) register the presence of H. leporispalustris infesting Didelphimorphia in this country but tick stages found on hosts are not stated.

24) Ixodes affinis Neumann, 1899 is a Nearctic and Neotropical tick species, but there is confusion about the presence (and hosts) of I. affinis in the southern range of its distribution because they may represent Ixodes aragaoi Fonseca, 1935 a or Ixodes pararicinus Keirans and Clifford, 1985 as discussed in Guglielmone et al. (2003b). Therefore, data below should be considered tentative. See also I. aragaoi for unjustified synonym with I. affinis .

Panama, Colón, Piña (09º16´N 80º03´W), MF on D. marsupialis (Fairchild et al. 1966) .

25) Ixodes amarali Fonseca, 1935 b is a South American tick species whose adults are mostly found on Didelphimorphia and to a lesser extent on sigmodontin rodents, but most nymphs are found on Sigmodontinae (Barros-Battesti & Knysak 1999) .

Brazil, LU in the northeastern area of the country, F on Monodelphis domestica (Wagner) (Barros-Battesti & Knysak 1999) .

Ceará, São Benedito (04º02´S 40º45´W), F on D. albiventris (Barros-Battesti & Knysak 1999) .

Paraíba, Princesa Izabel (07º44´S 37º59´W), F on M. domestica (Fonseca 1958) .

Pernambuco, Pesqueira (08º21´S 36º43´W), F on M. domestica (Fonseca 1958) ; Triunfo (07º49´S 38º06´W), FNL on M. domestica (Barros-Battesti & Knysak 1999) .

Rio de Janeiro, Seropedica (22º42´S 43º42´W), F on Didelphis sp. (Faccini et al. 1999, but the host is probably D. aurita ).

Additional geographical information from parasitism on non-marsupial hosts

Brazil, Alagoas, Anadias (09º41´S 36º18´W), N on Rodentia ; Palmeira dos Indios (09º24´S 36º37´W), N on Rodentia (Barros-Battesti & Knysak 1999) .

Ceará, Viçosa (04º27´S 37º47´W), N on Rodentia (Barros-Battesti & Knysak 1999) .

Minas Gerais, Belo Horizonte (19º55´S 43º56´W), F on Bolomys lasiurus (Lund) (= Necromys lasiurus (Lund)) , F on Oryzomys subflavus (Wagner) (= Cerradomys subflavus (Wagner)) (Botelho and Linardi 1996) ; Guaraciaba (20º33´S 43º00´W), N on Rodentia (Barros-Battesti & Knysak 1999) ; Ouro Preto (20º23´S 43º30´W), N on Oligoryzomys microtis (Allen) (Barros-Battesti & Knysak 1999) . The last host is mainly established in the Amazonian (Wilson & Reeder 2005; Oliveira & Bonvicino 2006) and the actual host is most probably different to O. microtis .

Pernambuco, Bom Conselho (09º09´S 36º41´W), N on O. subflavus (Wagner) (= C. subflavus (Wagner)) ; Garanhuns (08º53´S 36º29´W), FNL on O. subflavus (= C. subflavus ) (Barros-Battesti & Knysak 1999).

26) Ixodes aragaoi Fonseca, 1935 a, is a South American species very close to Ixodes pararicinus that is found on a variety of hosts (Guglielmone et al. 2003b). Durden and Keirans (1996) and Guzmán-Cornejo and Robbins (2010), among others, consider I. aragaoi to be a synonym of I. affinis but this is unjustified as discussed in Onofrio et al. (2009).

Brazil. Pinter et al. (2005) register the presence of I. aragaoi sub-adults on “marsupials”.

27) Ixodes boliviensis Neumann, 1904 is very close to Ixodes diversifossus Neumann, 1899 (Nearctic distribution); this last name will prevail if they are demonstrated to be synonyms. Under the current conditions I. boliviensis is a Neotropical species found on a vast array of hosts but more often on carnivores from southern Mexico to Bolivia (Guglielmone et al. 2003b).

Panama, Chiriqui, LU, F on D. marsupialis (Fairchild et al. 1966) .

Colombia, Boyacá, Muzo (05º32´N 74º06´W), N on D. marsupialis (Osorno Mesa 1942, who states that the tick found is near to I. boliviensis ).

28) Ixodes fuscipes Koch, 1844 is a South America species found usually on rodents (Guglielmone et al. 2003b). Brazil, Santa Catarina, Florianápolis (27º36´S 48º33´W), N on L. crassicaudata (Arzua et al. 2005) .

29) Ixodes lasallei Méndez Arocha and Ortiz, 1958 is a South American tick species prone to feed on rodents (Guglielmone et al. 2003b).

Venezuela, Amazonas, Cerro La Neblina (00º48´N 66º00´W), MFNL on Marmosa demerarae Thomas (Guerrero 1996) ; Río Cunucunuma at North-North West of La Esmeralda (03º39´N 65º21´W), N on P. opossum (Jones et al. 1972) . The last host is probably Philander mondolfi Lew, Pérez-Hernández and Ventura , and surely is not P. opossum because there are no Venezuelan populations of this species (Patton & Silva 2007).

30) Ixodes longiscutatus Boero, 1944 is a species found only in South American whose ecology need of further studies, but Sigmodontinae appears to be the most relevant host for its larvae and nymphs (Venzal et al. 2008b).

Argentina, Misiones, Ruta 14 and Río Victoria (26º57´S 54º27´W), F on P. opossum (= Philander frenatus (Olfers) (Ivancovich & Luciani 1992) , but this record is considered doubtful by Venzal et al. (2008b).

31) Ixodes loricatus Neumann, 1899 is a Neotropical species with adult ticks usually found on South American didelphimorphs. Relevant hosts for larvae and nymphs are marsupials and sigmodontin rodents. All tick stages of I. loricatus were considered to be highly specific to Didelphidae (Hoogstraal & Aeschlimann 1982) , but Nava et al.

(2004) clearly show that rodents are important for its life cycle. This study revealed that in the southern range of the distribution of I. loricatus adults were only found on Didelphidae and larvae and nymphs on Sigmodontinae .

Argentina, LU, MF on Didelphis paraguayensis Allen (= D. albiventris) (Aragão 1935).

Buenos Aires, LU, AN on Didelphis azarae Temminck (= D. albiventris ) and L. crassicaudata (Mauri & Navone 1988) , F on Didelphys (sic) azarai (sic) Temminck (= D. albiventris ) (Neumann 1910, who previously classified this tick as I. angustus Neumann, 1889 , and this error has been repeated until recently by several authors but I. angustus is a Nearctic-Palearctic species); Delta del Paraná (33º43´S 59º15´W), MF on L. crassicaudata (Keirans 1982) ; Delta del Paraná INTA (34º25´S 58º35´W), MFN on D. albiventris , MFN on L. crassicaudata (Ivancovich & Luciani 1992) , which is a controversial record for ticks classified first as Ixodes brunneus Koch, 1844 , and then as I. luciae as discussed in the note below; La Balandra (34º56´S 57º42´W), F on L. crassicaudata (Nava et al. 2004) .

Chaco, Campo Bermejo (26º38´S 59º03´W), F on L. crassicaudata ; Campo Winter (26º44´S 59º17´W), MF on D. albiventris (Ivancovich & Luciani 1992) , MFN on L. crassicaudata (Ivancovich 1973) ; Colonia Benítez INTA (27º19´S 58º56´W), ML on D. albiventris ; Estancia La Aurora (26º44´S 58º59´W), F on D. albiventris (Ivancovich & Luciani 1992) ; Ruta 90 and Río de Oro (26º25´S 59º23´W), MFN on D. albiventris (Ivancovich 1973) ; General José de San Martín (26º32´S 59º21´W), F on L. crassicaudata (Ivancovich & Luciani 1992) .

Córdoba, Morteros (30°42´S 62º00´W), MF on D. albiventris (Guglielmone et al. 2007a).

Corrientes, Corrientes (27º28´S 58º50´W), FL on D. albiventris (Gómez et al. 2000) .

Formosa, Colonia El Colorado (26º18´S 59º24´W), MF on L. crassicaudata ; Colonia Villafañe (26º12´S 59º03´W), M on D. albiventris ; El Colorado (26º19´S 59º23´W), F on D. albiventris ; El Colorado INTA (26º19´S 59º21´W), MF on D. albiventris , MFNL on L. crassicaudata , F on P. opossum (= P. frenatus ); Laguna Blanca (25º08´S 58º15´W), F on P. opossum (= P. frenatus ) (Ivancovich & Luciani 1992).

Misiones, Cuartel Victoria (26º44´S 54º20´W), F on D. albiventris (Ivancovich & Luciani 1992) .

Salta, LU, M on D. paraguayensis (= D. albiventris) (Aragão 1935).

Note: the record in Aragão (1935) is considered doubtful for the reason explained for I. loricatus in Brazil under the heading “ Additional information from parasitism on non-marsupial hosts ”.

Santa Fe, Funes (32º55´S 60º48´W), M on D. albiventris (Nava et al. 2004) ; Malabrigo (29º20´S 59º59´W), MF on D. paraguayensis (= D. albiventris ), F on L. crassicaudata (Aragão 1935) ; Santa Clara de Saguier (31º20´S 61º49´W), F on D. albiventris (Nava et al. 2004) ; Sauce Viejo (31º46´S 60º49´W), F on D. albiventris ; Sauce Viejo Aeropuerto (31º43´S 60º48´W), F on L. crassicaudata ; North of Sauce Viejo (31º 42´S 60º48´W) (Faccioli 2011).

Note: Ivancovich and Luciani (1992) present records of I. luciae from man, D. albiventris and L. crassicaudata in Delta of Paraná River which corresponds to the southernmost record for this species of tick. The authors state that the ticks were classified as I. brunneus but this diagnosis was later changed to I. luciae These specimens (all collected on December 20, 1976) were kept in three vials at the Estación Experimental Agropecuaria Colonia Benítez, Chaco, Argentina; vial numbered 451 with a label of I. brunneus contained one nymph (host non-stated), vial 452 with I. brunneus label contained seven males, two females and three nymphs (host non-stated) and a vial with repeated number 452 and with an Ixodes label contained one male and four females (from L. crassicaudata ). However, the vial 451 contained a nymph of the Ixodes auritulus Neumann, 1904 tick group, and all the remaining 17 ticks were identified as I. loricatus . We are not absolutely certain that all ticks indicated in Ivancovich and Luciani (1992) were revised because in the article it is stated that they found one female of I. luciae on man, eight males, one female and four nymphs on D. albiventris and one male, one female and four nymphs on L. crassicaudata . However, what is certain is: 1) most of the alleged records of I. luciae are in fact I. loricatus , 2) I. auritulus were first confused with I. brunneus and later with I. luciae , 3) the alleged infestation of man with I. luciae is most probably the result of tick misidentification. Consequently, we regard records of I. luciae on D. albiventris and L. crassicaudata in Ivancovich and Luciani (1992) as I. loricatus .

Brazil, LU, MF on Didelphis quica Temminck (= P. frenatus) (Neumann 1899); ML on D. albiventris (Keirans 1982), F on C. minimus, ML on D. marsupialis (Keirans 1985).

Meridional Region, LU, N on D. quica (= P. frenatus) (Neumann 1899).

Northeast Region, LU, MF on D. albiventris (Barros-Battesti & Knysak 1999).

Alagoas, Palmeira dos Indios (09º24´S 36º37´W), F on D. albiventris (Barros-Battesti & Knysak 1999) .

Ceará, Coité (07º25´S 38º43´W), F on Didelphis sp. (Barros-Battesti & Knysak 1999, the host is most probably D. albiventris ).

Goiás, Anápolis (16º19´S 48º57´W), MF on D. albiventris , F on Philander sp. (Barros-Battesti & Knysak 1999), MF on Didelphis sp. (Cooley & Kohls 1945, the host is most probably D. albiventris ), F on Metachirus opossum (Linnaeus) (= P. opossum ) (Fonseca & Aragão 1952).

Note: the Didelphis sp. hosts above are considered to be D. albiventris because it is the only species for the genus established in that locality.

Mato Grosso do Sul, Parque Estadual do Prosa (20º26´S 54º38´W), MFNL on D. albiventris (Miziara et al. 2008) .

Minas Gerais, Araxá (19º35´S 46º56´W), F on Didelphis sp. (most probably D. albiventris ), M on Marmosa sp. (Barros-Battesti & Knysak 1999); Belo Horizonte (19º49´S 43º57´W), F on D. albiventris (Schumaker et al. 2000) ; Mar de Espanha (20º52´S 43º00´W), MF on D. marsupialis (Keirans 1982, but the host is surely D. aurita because D. marsupialis is not established in this locality).

Paraíba, João Eugenio (coordinates unknown), F on Didelphis sp. (Barros-Battesti & Knysak 1999).

Paraná, Antonina (25º26´S 48º42´W), F on P. opossum (= P. frenatus ) (Arzua et al. 2005); Caiobá (25º50´S 48º33´W), M on M. opossum (= P. frenatus ) (Guimaraes 1945); Campo Largo (25º28´S 49º32´W), F on Didelphis sp.; Castro (24º39´S 50º04´W), F on D. albiventris ; Curitiba (25º25´S 49º17´W), M on D. albiventris , F on D. marsupialis but most probably D. aurita ; Pinhao (25º42´S 51º40´W), MN on D. albiventris ; MN on P. opossum (= P. frenatus ) (Arzua et al. 2005); Matinhos (25º49´S 48º33´W), FNL on P. opossum (= P. f re n a t u s) (Barros & Baggio 1992; Arzua et al. 2005); Ponta Grossa (25º06´S 50º10´W), MFN on D. albiventris , MFN on D. marsupialis but most probably D. aurita , MFN on L. crassicaudata (Barros & Baggio 1992) ; Ponta Grossa "Parque Estadual Vila Velha" (25º12´S 50º05´W), MF on D. albiventris , MFL on D. marsupialis but most probably D. aurita , FN on L. crassicaudata ; Quatro Barras "Morro do Anhangava" (25º23´S 48º59´W), MF on P. opossum (= P. fre n a tu s); Río Ivaí (coordinates unknown), MF on Didelphis sp., MF on P. opossum (= P. frenatus ); Tunas do Paraná "Parque Estadual de Campinhos" (25º02´S 49º03´W), F on P. opossum (= P. frenatus ) (Arzua et al. 2005).

Pernambuco, Caruarú (08º16´S 35º58´W), F on D. albiventris ; Garanhuns (08º52´S 36º29´W), MF on D. albiventris (Barros-Battesti & Knysak 1999) ; Reserva Ecológica de Gurjaú (08º14´S 35º00´W), A on Didelphis sp., A on D. albiventris , A on D. marsupialis but most probably D. aurita , N on Monodelphis americana (Müller) , N on Marmosa sp.; N on Marmosa murina (Linnaeus) (Botelho et al. 2004) .

Rio de Janeiro, Gavea (22º58´S 43º14´W), F on Didelphis sp. (Cooley & Kohls 1945); Teresópolis (22º24´S 42º57´W), F on Didelphis sp. (Labruna et al. 2002b); Tijuca (22º56´S 43º15´W), F on Didelphis sp. (Cooley & Kohls 1945). The hosts are most probably D. aurita .

Rio Grande do Sul, LU, NL on Microdelphys sorex (Hensel) (= Monodelphis sorex (Hensel) (Neumann 1899) ; Eldorado do Sul (30º05´S 51º25´W), M on D. marsupialis ; Guaiba (30º04´S 51º44´W), M on D. marsupialis (Evans et al. 2000) ; Pelotas (31º46´S 52º20´W), MFNL on D. albiventris , FN on D. marsupialis , F on L. crassicaudata (Barros-Battesti & Knysak 1999; Brum et al. 2003; Muller et al. 2005). Most probably the hosts identified as D. marsupialis in Rio Grande do Sul are in fact D. aurita . If this is confirmed the southern distribution of D. aurita in Cerqueira and Tribe (2008) will increase about 240 km.

Santa Catarina, Ilha de Ratones Grande (27º29´S 48º 34´W), MF on D. aurita ( Salvador et al. 2007); Joinville (26º18´S 48º50´W), F on D. marsupialis (Keirans 1985, the host is most probably D. aurita ).

São Paulo, Caçapava (23º06´S 45º42´W), A on D. aurita (Souza et al. 2006) ; Cotia (23º36´S 46º55´W), F on D. marsupialis (Labruna et al. 2002b, but the host is most probably D. aurita ); Francisco Morato (23º15´S 46º45´W), F on Didelphis sp. (Barros-Battesti & Knysak 1999); Fundação Parque Zoológico São Paulo (23º38´S 43º38´W), A on D. aurita (Labruna et al. 2004) ; Ipiranga (21º48´S 47º42´W), F on D. aurita (Aragão 1918) ; Itapevi (23º33´S 46º56´W), F on D. aurita (Barros-Battesti et al. 2000, several adult ticks were named as Ixodes didelphidis Fonseca and Aragão, 1952); Mogi das Cruzes (23º38´S 46º11´W), F on D. aurita (Horta et al. 2006) , A on D. albiventris and-or D. aurita (Horta et al. 2007) ; Mogi das Cruzes-Serra de Itapety (23º26´S 46º09´W), F on D. marsupialis but most probably D. aurita , N on Marmosa sp., MF on P. opossum (= P. frenatus ) (Labruna et al. 1997); Monte Alegre do Sul (22º39´S 46º41´W), A on D. albiventris (Souza et al. 2006) ; Pirassununga (21º59´S 47º25´W), F on Didelphis sp. (Labruna et al. 2002b), A on D. albiventris and-or D. aurita (Horta et al. 2007) ; Ribeirão Pires (23º43´S 46º24´W), F on Marmosa sp. (Barros-Battesti & Knysak 1999); Sampaio Moreira (coordinates unknown), F on Marmosa sp. (Barros-Battesti & Knysak 1999); São Paulo (23º31´S 46º37´W), F on Didelphis sp., F on D. albiventris , MF on D. marsupialis but most probably D. aurita , M on Marmosa sp. (Barros- Battesti & Knysak 1999), A on D. albiventris and-or D. aurita (Horta et al. 2007) ; Serra do Mar State Park (23º52´S 46º26´W), A on D. aurita (Sabatini et al. 2010) .

Colombia. In the tick collection of the Instituto Nacional de Tecnología Agropecuaria, Estación Experimental Agropecuaria Rafaela (INTA), Rafaela, Argentina there is a female labeled as Ixodes loricatus , from Metachirus nudicaudatus , 25 noviembre 1970, La Tirana (06º50´N 75º34´W), Antioquia, gift from the United States National Tick Collection (USNTC), Statesboro, U.S.A. It is part of a lot of eight female ticks maintained in the USNTC under accession number RML57488. We compared this specimen with bona fide I. loricatus females from Argentina and Brazil and with the descriptions contained in Marques et al. (2004) and Onofrio et al. (2009) because the types of I. loricatus are not available (Guglielmone et al. 2003b). The results are in the note below.

Note: the Colombian specimen, almost unfed, resembles at first sight under low magnification I. loricatus although it is pale brown and the coxae not as dark as in Argentinian and Brazilian specimens. However, this situation changed drastically when morphological details were compared under high stereoscopy magnification. The Colombian specimen differs from the Argentinian and Brazilian ticks as follows: pronounced ridges lateral to each porose area surpassing the posterior border of the basis capituli giving the impression of sharp cornua, a feature absent in I. loricatus ; scutum obviously longer than wide (ratio 1.5) while it is just longer than wide (ratio 1.2) in I. loricatus ; coxae I with two acute spurs with the external longer than the internal, while the spurs are less acute in I. loricatus ; hypostome pointed, while it is blunt in I. loricatus ; presence of a dark elevated borders on the ventral basis capituli which is absent from the Argentinian and Brazilian I. loricatus . We conclude that the alleged specimen of I. loricatus from Colombia is not a representative of this species. Lorenza Beati examined the specimens in RML57488 (USNTC) and found that they are not I. loricatus and their morphologies are similar to the specimen deposited in INTA. Additional studies are needed to determine if these specimens belong to a known species or represent a still undescribed taxon.

Guatemala. Monroy Lefebre and Cejas González (1988) state that I. loricatus is established in this country, but locality, hosts and tick stages found on them are not provided. This is a controversial record that will be further discussed below.

Panama, Note: Fairchild et al. (1966) allegedly found a bona fide female of I. loricatus in Darién, Tacarcuna Station (08º05´N 77º17´W) on Metachirus nudicaudatus on September 1, 1958. This specimen is kept in the United States Tick Collection under accession number RML37477 where it was examined by Valeria C. Onofrio, who concluded that it is not I. loricatus and species identification is under way. Therefore, Panama is excluded form the range of this species.

Paraguay, LU, A on D. albiventris (Whitaker & Abrell 1987).

Cordillera, probably from Sapucay (25º19´S 56º 55´W), MF on Didelphis sp. (Keirans 1982), but we were unable to locate this place with the coordinates provided by the author.

Presidente Hayes, Fortín Juan de Zalazar (23º06´S 59º18´W), MF on D. albiventris (Valeria C. Onofrio, personal communication). These ticks are deposited in the Gorgas Memorial Laboratory Tick Collection, and the locality is named “Juan de Zalazar” and allegedly located in the Boquerón Department. However, there is no Paraguayan locality named “Juan Zalazar” but “Fortín Juan de Zalazar” often named as “Fortín Zalazar” is located in the Department Presidente Hayes, close to the borderline with the Department Boquerón.

Uruguay, LU, MFN on D. albiventris, NL on M. dimidiata (Venzal et al. 2003).

Maldonado, Barra del Arroyo Maldonado (34º58´S 54º56´W), F on D. albiventris (Nava et al. 2004) .

Montevideo, Paso de la Arena (34º53´S 56º09´W), MF on L. crassicaudata (Wolffhugel 1933) .

Venezuela, Araguá, Rancho Grande Biological Station (10º21´N 67º36´W), NL on Monodelphis brevicaudata (Erxleben) (Jones et al. 1972) . See the note about M. brevicaudata in I. luciae .

Miranda, east of Caracas (10º29´N 66º44´W), L on M. robinsoni (Jones et al. 1972) .

Note: Jones et al. (1972) consider the precedent diagnosis of I. loricatus as tentative. There are other records from Venezuelan marsupials in Vogelsang and Cordero (1940) but the tick stages found on hosts and morphological support for the identification are not provided, bringing uncertainty about the presence of this tick in the country.

Additional geographical information from parasitism on non-marsupial or undetermined hosts

Argentina, Buenos Aires, Hudson (34º45´S 58º06´W), LN on Akodon azarae (Fischer) ; Punta Lara (34º47´S 58º01´W), L on A. azarae, Ramallo (33º32´S 59º52´W), N on Oligoryzomys flavescens (Waterhouse) ; San Nicolás (33º20´S 60º13´W), NL on A. azarae (Nava et al. 2004) .

Salta, Parque Nacional El Rey (24º41´S 64º40´W), NL on Calomys sp. (Nava et al. 2004).

Santa Fe, Reserva Natural de la Universidad Nacional del Litoral (31º23´S 60º55´W), L in nests of sigmodontine rodents (Manzoli et al., 2006); Santa Clara de Saguier (31º20´S 61º49´W), NL on Akodon sp. (Nava et al. 2004).

Tierra del Fuego, LU, F from an unknown hosts (Neumann 1901). This is a controversial record that will be further discussed below.

Brazil, Minas Gerais, Pacau (15º14´S 43º41´W), F on Agouti paca (Linnaeus) (= Cuniculus paca (Linnaeus)) (Aragão 1918) .

São Paulo, Araraquara (21º47´S 48º10´W), F on undetermined host; Barueri (23º29´S 46º51´W), M on undetermined host; Biritiba Mirim (23º34´S 46º01´W), F on undetermined host; Bragança Paulista (22º57´S 46º32´W), M on undetermined host; Taubaté (23º02´S 45º33´W) (Marques et al. 2004).

Note: Aragão (1936) has a record of I. loricatus on an undetermined host at Curralinho (01º43´S 49º43´W) in the state of Pará that represents the northernmost record of this species in Brazil. However, it should be noted that this diagnosis was made before the description of I. luciae in 1940. The armature of coxae (crucial for differential diagnosis between these species) was not considered in the key used by this author bringing some uncertainty about the identification. We can not say that this diagnosis is erroneous but being cautious it is considered that it at least requires confirmation.

Mexico, Tabasco, Frontera (18º32´N 92º38´W), MF on Ateles geoffroyi Kuhl (Keirans 1982, 1985).

Note: this Mexican record of I. loricatus is controversial. This tick species has not been found yet on Mexican marsupials or rodents but only once on this unusual monkey host for I. loricatus . We doubt about the origin of the specimens (see also the discussion section).

Uruguay, Canelones, Solymar Norte (34º47´S 55º56´W), L on O. flavescens (Nava et al. 2004) .

32) Ixodes luciae Sénevet, 1940 is a Neotropical species close morphologically and in host usage to the precedent species, I. loricatus . The only study on the ecology of I. luciae shows the relevance of sigmodontin rodents as hosts for larvae and nymphs of this tick (Díaz et al. 2007, 2009) because all specimens but two were found on them. However, other authors present several records of sub-adults ticks on Didelphidae (see below).

Argentina, LU, M on D. paraguayensis (= D. albiventris) (Cooley & Kohls 1945).

Salta, road to Isla de las Cañas (22º57´S 64º33´W), N on Micoureus constantiae (Thomas) (= Marmosa constantiae Thomas ), N on Thylamys sp., N on Thylamys venustus (Thomas) , F on L. crassicaudata (Autino et al. 2006) .

Tucumán, El Siambón (26º40´S 65º24´W), N on Thylamys cinderella Thomas (Autino et al. 2006) .

Note: the Argentinian record for I. luciae in Buenos Aires province, Delta del Paraná INTA (34º25´S 58º35´W) on D. albiventris and L. crassicaudata in Ivancovich and Luciani (1992) was changed to I. loricatus after examining the specimens as stated in the note of I. loricatus in Argentina (see details in the note under I. loricatus in Argentina).

Belize, Middlesex (17º01´N 88º30´W), MF on Didelphis mesamericana Allen (= D. marsupialis ) (Cooley & Kohls 1945).

Bolivia, Santa Cruz, Buen Retiro (17º16´S 63º42´W), MFNL on D. paraguayensis (= D. albiventris ) (Fonseca 1959).

Brazil, Acre, Assis (10º56´S 69º34´W), A on D. marsupialis (Marcelo B. Labruna, personal communication).

Pará, Igarapé Açú (01º07´S 47º37´W), MF on P. opossum (Barros-Battesti & Knysak 1999) .

Rondônia, LU, MF on Monodelphis sp. (Labruna et al. 2005 a, most probably the host is Monodelphis glirina (Wagner); Jamari River headwaters (10º18´S 63º 14´W), F on D. marsupialis (Labruna et al. 2009).

Colombia. There are records of I. luciae in Wells et al. (1981) and Osorno-Mesa (1942 as I. loricatus spinosus Nuttall, 1910 , a preoccupied name replaced by I. loricatus vogelsangi Santos Dias, 1954 ) on South American marsupials, but there is no information about tick stages found on hosts.

Costa Rica. Guglielmone et al. (2003b) state that I. luciae is present in the country but no host and tick stages are provided.

Ecuador. Guglielmone et al. (2003b) state that I. luciae is found in the country, but no host and tick stages are provided.

French Guiana, Cayena, Cayenne, (04º56´N 52º19´W), F on D. marsupialis (Floch & Fauran 1958) .

Guatemala, Izabal, Bobos (15º21´N 88º48´W), MF on D. mesamericana (= D. marsupialis ); Escobas (15º41´N 88º38´W), F on D. mesamericana (= D. marsupialis ) (Cooley & Kohls 1945).

Honduras. Onofrio et al. (2002) state that I. luciae is established in this country, but hosts and tick stages found on them are not provided.

Mexico, Chiapas, Finca Germania (15º04´N 92º31´W), MF on Didelphis sp. (Vázquez 1946 as Ixodes scuticrenatus Vázquez, 1946 ).

Colima, La Barragana (19º20´N 103º45´W), MF on D. marsupialis (Guzmán-Cornejo et al. 2007) .

Tabasco, Frontera (18º32´N 92º38´W), F on Didelphis sp. (Keirans 1982).

Veracruz, North-East of las Minas (19º41´N 97º09´W), M on Didelphis sp. (Guzmán-Cornejo et al. 2007).

Nicaragua. Jones et al. (1972) state that I. luciae is established in this country, but hosts and tick stages found on them are not provided.

Panama, Bocas del Toro, Almirante (09º18´N 82º24´W), N on M. robinsoni (Fairchild et al. 1966) .

Colón, Alhajuela (09º11´N 79º38´W), MF D. mesamericana (= D. marsupialis ) (Cooley & Kohls 1945), M on D. marsupialis (Fairchild 1943) ; Camp Piña (09º16´N 80º03´W), MF on D. marsupialis , F on P. opossum ; Fort Gulick (currently named Fuerte Espinar) (09º18´N 79º53´W), F on P. opossum (Fairchild et al. 1966) .

Comarca de Kuna Yala (previously known as Comarca San Blas), Río Mandinga (09º29´N 79º05´W), M on D. marsupialis (Fairchild et al. 1966) .

Darién, Tacarcuna Station (08º10´N 77º18´W), A on D. marsupialis (Fairchild et al. 1966) .

Panama, Cerro Azul (09º12´N 79º21´W), A on D. marsupialis , F on P. opossum ; Cerro Campana (08º41´N 79º56´W), A on D. marsupialis (Fairchild et al. 1966) ; Panamá City (08º58´N 79º32´W), F on D. marsupialis (Bermúdez et al. 2010) .

Peru, Loreto, 13 de Febrero (04º02´S 73º26´W), F on P. opossum ; Belén (03º50´S 73º13´W), F on Caluromys lanatus (Olfers) ; Ex Petroleros (04º05´S 73º27´W), F on P. opossum ; Fundo San Martín (03º58´S 73º24´W), F on P. opossum ; Los Delfines (03º51´S 73º21´W), F on P. opossum ; Moralillo (03º54´S 73º21´W), F on P. opossum , F on Marmosops sp., now identified as Marmosops impavidus Tschudi by one of the authors (MMD) Palo Seco (04º00´S 73º26´W), F on M. nudicaudatus , L on Micoureus Lesson (= Marmosa ) sp. (most probably Micoureus regina (Thomas) , F on Philander andersoni Pine ; Paujil (04º04´S 73º27´W), F on D. marsupialis , F on P. opossum ; Peña Negra (03º52´S 73º20´W), MF on P. opossum ; San Juan (03º59´S 73º25´W), F on P. opossum , F on P. andersoni ; San Lucas (04º06´S 73º22´W), F on P. andersoni (Díaz et al. 2007) .

Surinam, LU, F on Didelphis sp. (Keirans 1985).

Trinidad and Tobago, Trinidad, Bush Bush Forest (10º24´N 61º00´W), A on D. marsupialis , A on Marmosa sp. (Aitken et al. 1968).

Venezuela, Amazonas, Boca Mavaca (02º31´N 65º16´W), F on P. opossum ; Capibara (02º36´N 66º20´W), F on P. opossum (Jones et al. 1972) ; Cerro Tamacuari (01º41´N 64º26´W), MF on P. opossum (Guerrero 1996) ; Tamatama (03º09´N 65º50´W), FN on M. robinsoni , F on P. opossum (Jones et al. 1972) ; however these localities are out of the distribution range of P. opossum as stated by (Patton & Silva 2007) and the hosts should be either P. mondolfii or P. andersoni .

Araguá, Rancho Grande Biological Station (10º21´N 67º36´W), NL on M. brevicaudata (Jones et al. 1972) .

Barinas, Altamira (08º49´N 70º30´W), M on D. marsupialis , N on M. brevicaudata ; south-west of Altamira (08º47´N 70º31´W), NL on M. brevicaudata (Jones et al. 1972) .

Carabobo, Aguirre (10º11´N 68º19´W), NL on M. brevicaudata ; Montalbán (10º12´N 68º19´W), N on M. robinsoni , NL on M. brevicaudata ; south-east of Montalbán (10º10´N 68º20´W), F on D. marsupialis , L on M. robinsoni (Jones et al. 1972) .

Distrito Federal, south-west of Caracas (10º27´N 67º09´W), M on D. marsupialis (Jones et al. 1972) .

Falcón, La Pastora (11º12´N 68º35´W), MF D. marsupialis , NL on M. brevicaudata (Jones et al. 1972) ; San Esteban (11º28´N 69º22´W), M on D. marsupialis (Keirans 1982) .

Guárico, south-east of Caracas (09º56´N 66º32´W), MF on D. marsupialis (Jones et al. 1972) .

Miranda, near Turgua (10º21´N 66º44´W), N on Marmosa cinerea (Temminck) (= Marmosa demerarae (Thomas) , because the name cinerea was preoccupied); Naiguatá (10º29´N 66º44´W), MF on Didelphis sp. (Méndez Arocha & Ortiz, 1958, the host is most probably D. marsupialis ); South of Río Chico (10º15´N 65º59´W), N on M. brevicaudata (Jones et al. 1972, but the host is doubtful because there is no evidence of its presence in Miranda).

Yaracuy, Aroa (10º25´N 68º53´W), MF D. marsupialis ; north-west of Urama (10º34´N 68º26´W), M on D. marsupialis , N on M. cinerea (= M. demerarae ) (Jones et al. 1972).

Zulia, south-west of Machiques (09º58´N 72º42´W), N on M. brevicaudata (Jones et al. 1972) , but the host is most probably Monodelphis palliolata (Osgood) .

Note: the following Venezuelan records are from Jones et al. (1972) who considered the specimens as probable I. luciae without further comments. In the light of the many records of I. luciae in Venezuela we treat these records as provisionally valid.

Amazonas, Río Cunucunuma at North-North West of La Esmeralda (03º39´N 65º21´W), L on Caluromys philander (Linnaeus) , L on M. murina .

Distrito Federal, South-West of Caracas (10º27´N 67º20´W), L on M. brevicaudata .

Falcón, North-West of La Pastora (11º12´N 68º36´W), NL on M. brevicaudata ; Mirimire (11º09´N 68º53´W), M on D. marsupialis .

Guárico, South-East of Caracas (09º58´N 66º33´W), L on M. brevicaudata , but the host is most probably M. palliolata or a related species instead of M. brevicaudata .

Miranda, South of Caracas (10º25´N 66º55´W), L on M. robinsoni .

Trujillo, West-North West of Valera (09º23´N 70º40´W), L on M. robinsoni .

Note: all records of M. brevicaudata have to be considered cautiously because considerable controversy exists about it and related species (Pine & Handley 2007).

Additional geographical information from parasitism on non-marsupial or undetermined hosts

Argentina, Salta, West South West of Pulares (25º05´S 65º37´W), N on Calomys callosus (probably) from Autino et al. (2006), but host and locality position reevaluated.

Brazil, Amazonas, Santa Isabel do Río Negro (02º23´S 65º04´W), F on undetermined host (Onofrio et al. 2010).

Mato Grosso do Sul, Dos Irmãos do Buriti (20º41´S 55º17´W), F on undetermined host (Onofrio et al. 2010).

Pará, Belém (01º 26´S 48º 29´W), MF on undetermined host; Mirabá (05º20´S 49º07´W), F on undetermined host; São João de Pirabas (00º46´S 47º10´W), F on undetermined host; Tucuruí (03º40´S 49º42´W), M on undetermined host (Onofrio et al. 2010).

Rondônia, Amorim Farm (10º38´S 63º31´W), N on undetermined host (Labruna et al. 2005a); Campo Novo de Rondônia (10º36´S 63º37´W), MF on undetermined host (Onofrio et al. 2010); Line C (10º26´S 63º21´W), N on undetermined host (Labruna et al. 2005a); Monte Negro (10º02´S 63º08´W), NL on undetermined host; Porto Velho (08º45´S 63º54´W), MF on undetermined host (Onofrio et al. 2010).

Panamá, Darién, Cerro Pirre (07º51´N 77º44´W), N on Oryzomys sp. (Fairchild et al. 1966).

Perú, Loreto, Ninarumi (03º51´S 73º22´W), N on Hylaeamys perenensis (Allen) ; Varillal (03º53´S 73º22´W), N on H. perenensis (Díaz et al. 2007) .

Venezuela, Distrito Federal, Caracas (10º29´N 66º55´W), MF on dog (Méndez Arocha & Ortiz 1958).

33) Ixodes pararicinus Keirans and Clifford, 1985 in Keirans et al. (1985) is a South American species with several hosts for all parasitic stages (Venzal et al. 2005; Autino et al. 2006).

Argentina, Salta, Parque Nacional El Rey (24º15S, 64º40`W), L on T. venustus (Beldoménico et al. 2003) . Peru, Loreto, Fundo San Martín (03º58´S 73º24´W), L on Monodelphis adusta (Thomas) (Díaz et al. 2007) .

34) Ixodes rubidus Neumann, 1901 is a Nearctic-Neotropical species but it is not established in South America. Their usual hosts are carnivore mammals (Guglielmone et al. 2003b).

Guatemala, Chimaltenango, Yepocapa (14º29´N 90º56´W), F on Didelphis sp. (Fairchild et al. 1966).

35) Ixodes venezuelensis Kohls, 1953 is a Neotropical tick species feeding on Didelphimorphia and Rodentia with Sigmodontinae playing a relevant role as hosts for all parasitic stages (Durden & Keirans 1994).

Costa Rica, Heredia, LU (10º25´N 84º00´W), N on Marmosa mexicana Merriam, N on P. opossum (Durden & Keirans 1994) .

Panama, Darién, LU, F on M. adusta (Durden & Keirans 1994) .

Venezuela, northern region, LU, FN on M. brevicaudata (Durden & Keirans 1994) .

Amazonas, LU, L on Philander sp. (Durden & Keirans 1994); Boca Mavaca (02º 31´N 65 º16´W), NL on D.

marsupialis ; Casiquiare Canal Capibara (02º36´N 66º21´W), L on P. opossum ; Río Cunucuma north-north west of La Esmeralda (03º39´N 65º21´W), NL on P. opossum (Jones et al. 1972) . Most probably the species of Philander in Jones et al. (1972) are P. andersoni or P. mondolfii .

Araguá, Sierra Maestra (coordinates unknown), FN on M. brevicaudata (Kohls 1953) .

Barinas, Altamira (08º49´N 70º30´W), NL on M. brevicaudata (Jones et al. 1972) .

Falcón, LU, N on M. brevicaudata (Durden & Keirans 1994).

Táchira, LU, L on Marmosa sp. (Durden & Keirans 1994).

Trujillo, West-Northwest of Valera (09º23´N 70º40´W), FNL on M. brevicaudata (Jones et al. 1972) ; these last authors have a record considered “probable I. venezuelensis ” for this state at east of Trujillo (09º22´N 70º17´W), L on Marmosa dryas (Thomas) (= Gracilinanus dryas (Thomas) .

See the note about M. brevicaudata in I. luciae .

Additional geographical information from parasitism on non-marsupial hosts

Colombia, Antioquía, Valdivia (07º11´N 75º27´W), F on Melanomys caliginosus (Tomes) (Kohls 1953) .

Costa Rica, Alajuela, LU (10º30´N 84º30´W), F on Heteromys desmarestianus Gray (Durden & Keirans 1994) .

Venezuela, Amazonas, South East of Boca Mavaca (02º22´N 65º08´W), N on Myoprocta pratti Pocock ; North of Raya (05º25´N 67º36´W), N on Proechimys guyannensis (Geoffroy) (Jones et al. 1972) .

Apure, North of Nula (07º18´N 71º54´W), N on Proechimys semispinosus (Tome) (Jones et al. 1972) , but this host is not established in Venezuela and the species of Proechimys found in the state of Apure are uncertain (Gómez-Laverde et al. 2008).

Aragua, Rancho Grande (10º21´N 67º36´W), F on Ichthyomys pittieri Handley and Mondolfi (Guerrero 1996) .

Bolívar, South East of El Dorado (06º13´N 61º28´W), N on P. guyannensis ; North East of Icaburú (04º34´N 61º30´W), N on P. guyannensis (Jones et al. 1972) .

Carabobo, Montalbán (10º12´N 68º20´W), NL on Dasyprocta aguti (Linnaeus) (= Dasyprocta leporina (Linnaeus) (Jones et al. 1972) ; Bejuma (10º10´N 68º15´W), F on I. pittieri (Guerrero 1996) .

Mérida, East of Tabay (08º38´N 71º02´W), Oryzomys minutus (Tomes) (= Microryzomys minutus (Tomes) (Jones et al. 1972) .

Miranda, South of Caracas (10º25´N 66º55´W), F on Heteromys anomalus (Thompson) (Jones et al. 1972) .

Trujillo, East of Trujillo (09º22´N 70º18´W), NL on O. minutus (= M. minutus ) (Jones et al. 1972).

36) Rhipicephalus microplus (Canestrini, 1888) is an extremely important tick species that affects the cattle industry in most tropical and sub-tropical areas of the world, and is found occasionally on mammalian hosts different to artiodactyls.

Argentina. Boero (1954) and Boero and Boehringer (1967) present vague information of infestation of R. microplus on South American marsupials.

37) Rhipicephalus sanguineus (Latreille, 1806) species complex has a world wide distribution feeding mainly on domestic dogs, but there are also records on a wide variety of hosts. The number of species that forms this complex is undetermined.

Argentina and Peru. Santa Cruz et al. (1998) and Need et al. (1991) present records of ticks of R. sanguineus complex on South American marsupials for Argentina and Peru, respectively, without stating tick stages found on hosts.