Nymphs of the genus Amblyomma (Acari: Ixodidae) of Brazil: descriptions, redescriptions, and identification key

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Ticks and Tick-borne Diseases 1 (2010) 75–99

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Ticks and Tick-borne Diseases

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riginal article

ymphs of the genus Amblyomma (Acari: Ixodidae) of Brazil: descriptions,edescriptions, and identification key

hiago F. Martinsa, Valeria C. Onofriob, Darci M. Barros-Battestib, Marcelo B. Labrunaa,∗

Department of Preventive Veterinary Medicine and Animal Health, Faculty of Veterinary Medicine, University of São Paulo, São Paulo, SP, BrazilLaboratory of Parasitology, Butantan Institute, São Paulo, SP, Brazil

r t i c l e i n f o

rticle history:eceived 23 January 2010eceived in revised form 12 March 2010ccepted 14 March 2010vailable online 20 April 2010

eywords:mblyommaymphs

a b s t r a c t

Together with the larval stage, the nymphal stage of ticks of the genus Amblyomma are the most aggres-sive ticks for humans entering areas inhabited by wildlife and some domestic animals in Brazil. However,due to the absence of morphological descriptions of the nymphal stage of most Brazilian Amblyommaspecies, plus the lack of an identification key, little or nothing is known about the life history of Ambly-omma spp. nymphs in the country. In the present study, morphological description of the nymphal stage,illustrating important external characters through scanning electron microscopy, is provided for nymphsof 15 Amblyomma species that occur in Brazil, for which the nymphal stage had never been described: A.aureolatum, A. auricularium, A. calcaratum, A. coelebs, A. fuscum, A. humerale, A. incisum, A. latepunctatum,

escriptiondentification keyrazil

A. naponense, A. nodosum, A. ovale, A. pacae, A. pseudoconcolor, A. scalpturatum, A. varium. In addition,the nymphal stage of 12 Amblyomma species, which had been previously described, are redescribed: A.brasiliense, A. cajennense, A. dissimile, A. dubitatum, A. longirostre, A. oblongoguttatum, A. parkeri, A. parvum,A. romitii, A. rotundatum, A. tigrinum, A. triste. The descriptions and redescriptions totalized 27 species.Only 2 species (A. geayi, A. goeldii) out of the 29 Amblyomma species established in Brazil are not included

chotombly

in the present study. A dithe nymphal stage of 27 A

ntroduction

The genus Amblyomma (Acari: Ixodidae) is currently repre-ented by 130 valid tick species (Nava et al., 2009a, 2009b), 29 ofhich (22.3%) are established in Brazil (Dantas-Torres et al., 2009).hile the adult stage of these 29 species has been morphologi-

ally well described, resulting in good identification keys, the sameannot be said for the immature stages (Guglielmone et al., 2003;nofrio et al., 2006). The nymphal stage of 17 Amblyomma species

rom Brazil remains undescribed. This scenario has precluded theonstruction of identification keys. Consequently, taxonomic iden-ification to species level of field-collected Amblyomma nymphsas been a laborious and very difficult task, sometimes impossible.or example, species identification of field-collected Amblyommaymphs in recent studies in Brazil were possible only because these
ymphs were reared to the adult stage in the laboratory or becauseolecular methods were applied, such as DNA sequencing ofolecular markers previously determined for each species through
he adult stage (Labruna et al., 2007; Ogrzewalska et al., 2009).

∗ Corresponding author at: Faculdade de Medicina Veterinária e Zootecnia, Uni-ersidade de São Paulo, Av. Prof. Orlando Marques de Paiva 87, Cidade Universitária,5508-270 São Paulo, SP, Brazil. Tel.: +55 11 3091 1394; fax: +55 11 3091 7928.

E-mail address: [email protected] (M.B. Labruna).

877-959X/$ – see front matter © 2010 Elsevier GmbH. All rights reserved.oi:10.1016/j.ttbdis.2010.03.002

mous identification key is included to support taxonomic identification ofomma species established in Brazil.

© 2010 Elsevier GmbH. All rights reserved.

Herein, we describe the nymphal stage of 15 Amblyomma speciesfor the first time and redescribe the nymphal stage of another 12Amblyomma species from Brazil. In addition to providing scanningelectron micrographs of the nymphs, we also include an identifi-cation key for the 27 Amblyomma species evaluated in the presentstudy.

Materials and methods

During 2000–2009, tick colonies of 23 Amblyomma species fromBrazil were maintained in the laboratory of the Faculty of Veteri-nary Medicine of the University of São Paulo at different periodsfor different purposes. Unfed nymphs, 15–30 days old, of the firstlaboratory generation (F1) of each species were killed in hot water(70–80 ◦C) and immediately preserved in 70% alcohol until furtherprocessing for description or redescription. In addition, alcohol-preserved F1 unfed nymphs from laboratory colonies of 3 otherAmblyomma species were kindly provided by colleagues from otherinstitutions, as stated in the Acknowledgments. All these 26 tick
colonies started with field-collected ticks as stated in Table 1.Species identification of the tick colonies was undertaken duringthe adult stage (F0) using specific identification keys and corre-sponding morphological redescriptions available in the currentliterature (Robinson, 1926; Aragão and Fonseca, 1961; Jones et
http://www.sciencedirect.com/science/journal/1877959X

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76 T.F. Martins et al. / Ticks and Tick-borne Diseases 1 (2010) 75–99

Table 1Origin of tick colonies used to obtain F1 nymphs described or redescribed in the present study.

Tick species Laboratory colonies

Origin Nymphs used

Stage Host Locality Date Accession no. N

Amblyomma aureolatum Female Canis familiaris Bom Jesus dos Perdões, SP, Brazil 04 October 2007 CNC-1458 10A. auricularium Female Dasypus novemcinctus Mossoró, RN, Brazil 12 December 2007 CNC-1459 10A. calcaratum Female Myrmecophaga tridactyla Sorocaba, SP, Brazil. 19 February 2008 CNC-1460 10A. coelebs Female Tapirus terrestris Teodoro Sampaio, SP, Brazil 04 September 2007 CNC-1461 10A. fuscum Larvae, nymphs Didelphis aurita Guarujá, SP, Brazil 26 May 2008 CNC-1462 10A. humerale Female Geochelone denticulata Linhares, ES, Brazil 18 September 2006 CNC-1463 10A. incisum Males, females Free-living Ribeirão Grande, SP, Brazil 11 May 2005 CNC-1464 10A. latepunctatum Males, females Free-living Governador Jorge Teixeira, RO, Brazil 31 January 2008 CNC-1465 10A. naponense Female Tayassu pecari Teodoro Sampaio, SP, Brazil 29 November 2004 CNC-1466 10A. nodosum Female M. tridactyla Presidente Epitácio, SP, Brazil 23 March 2001 CNC-1467 10A. ovale Female C. familiaris Monte Negro, RO, Brazil 23 November 2001 CNC-1468 10A. pacae Female Cuniculus paca Monte Negro, RO, Brazil 23 February 2002 CNC-1469 10A. pseudoconcolor Female Euphractus sexcinctus Água Morta, Salta, Argentina 25 July 2008 CNC-1470 6A. scalpturatum Males, females Free-living Governador Jorge Teixeira, RO, Brazil 31 January 2008 CNC-1471 10A. varium Female Bradypus variegatus Sorocaba, SP, Brazil 20 April 2000 CNC-0291 10A. brasiliense Female Pecari tajacu Teodoro Sampaio, SP, Brazil 06 March 2007 CNC-1472 10A. cajennense Female Equus caballus Pirassununga, SP, Brazil 01 March 2007 CNC-1473 10A. dissimile Female Iguana iguana Cuiabá, MT, Brazil 04 October 2007 CNC-1474 10A. dubitatum Female Hydrochoerus hydrochaeris Miracatu, SP, Brazil 04 December 2007 CNC-1475 10A. longirostre Female Sphiggurus villosus São Paulo, SP, Brazil 25 June 2008 CNC-1476 10A. oblongoguttatum Female C. familiaris Monte Negro, RO, Brazil 04 October 2007 CNC-1477 10A. parvum Female C. familiaris Barão de Melgaco, MT, Brazil 27 September 2002 CNC-1478 10A. romitii Female H. hydrochaeris Rurópolis, PA, Brazil 17 April 2009 CNC-1479 10A. rotundatum Female Boa constrictor Porto Velho, RO, Brazil 08 February 2008 CNC-1480 10A. tigrinum Female C. familiaris Herval, RS, Brazil 03 July 2002 CNC-1481 10A. triste Female Blastocerus dichotomus Bataguassu, MS, Brazil 10 January 2001 CNC-1482 10A. parkeri Nympha Coendou sp. Cotia, SP, Brazil 15 March 1933 IBSP-4458 1

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NC: Tick collection “Colecão Nacional de Carrapatos da Faculdade de Medicinacarológica do Instituto Butantan, São Paulo”. N: No. of specimens used for descripa Only a single engorged nymph was available for redescription.

l., 1972; Estrada-Pena et al., 2002; Barros-Battesti et al., 2005,007; Labruna et al., 2005; Onofrio et al., 2006, 2008). In the casef A. parkeri Fonseca and Aragão, 1952, no laboratory colony wasvailable; therefore we used a single engorged nymphal specimenor redescription that had been described by Fonseca and Aragão1952) (Table 1).

The nymphal stage of the 27 Amblyomma species listed in Table 1ere described or redescribed based on optical microscopy, fol-

owing previous authors who described Amblyomma nymphs fromhe New World (Cooley and Kohls, 1944; Guglielmone et al., 1990;strada-Pena et al., 1993, 2002; Nava et al., 2009b). For this purpose,0 specimens of each species (except for A. pseudoconcolor Aragão,908, and A. parkeri, for which only 6 and 1 specimen, respectively,ere available) were measured using the Image-Pro Plus 5.1 pro-

ram for analysis of images and morphometry, fitted to an OlympusZX stereoscope microscope. In the descriptions/redescriptionshat follow, all measurements are given in millimeters; first the

ean ± standard deviation, followed by the range in parenthe-es. With the exception of A. parkeri, representative specimensf all Amblyomma species were prepared for scanning electronicroscopy (SEM) following techniques described by Corwin et al.

1979).Finally, we constructed a dichotomous identification key based

n morphological differences and similarities of the nymphal stagef the 27 Amblyomma species evaluated in the present study.

esults

escriptions

mblyomma aureolatum (Pallas, 1772) (Fig. 1)Idiosoma. Length from apices of scapula to posterior body mar-

in 1.287 ± 0.072 (1.104–1.365), maximum breadth 0.907 ± 0.045

inária e Zootecnia da Universidade de São Paulo”. IBSP: Tick collection “Colecãoredescription, deposited in the designated tick collection.

(0.794–0.956), outline oval, longilinear, with 11 festoons with-out tubercles. Scutum. Length 0.660 ± 0.038 (0.568–0.707), breadth0.731 ± 0.022 (0.679–0.766), breadth/length ratio 1.109 ± 0.035(1.081–1.194), inornate, surface extensively shagreened (rugose);few punctations, larger and deeper laterally. Eyes not orbitedat lateral scutal angles at the level of scutal midlength. Cervicalgrooves reaching the scutal midlength, deeper at the anterior half.Spiracular plate triangular with rounded angles, with a discretedorsal prolongation; length 0.053 ± 0.002 (0.050–0.057), breadth0.026 ± 0.002 (0.021–0.028). Gnathosoma (capitulum). Length frompalpal apices to posterior margin 0.306 ± 0.030 (0.236–0.337),breadth 0.297 ± 0.012 (0.273–0.316). Basis capituli triangular, pos-terior margin slightly concave, without cornua; posterior marginconvex ventrally, without auriculae, lateral margin projected lat-erally. Palpi length 0.230 ± 0.018 (0.184–0.246), article I withoutventral prolongation, article II 0.152 ± 0.010 (0.127–0.167) long,article III 0.055 ± 0.006 (0.045–0.064) long. Hypostome roundedapically; length 0.212 ± 0.026 (0.169–0.267); length of toothed por-tion 0.119 ± 0.011 (0.098–0.136); dentition 2/2 with 7–8 teeth perrow. Legs. Coxa I with 2 stout spurs close to each other, the exter-nal longer; coxae II–IV with a small triangular spur that graduallydecreases in size from II to IV. Trochanters without spur; tar-sus I 0.364 ± 0.023 (0.316–0.396) long, 0.122 ± 0.013 (0.105–0.149)broad; tarsus IV 0.286 ± 0.022 (0.253–0.319) long, 0.092 ± 0.017(0.073–0.125) broad.

Amblyomma auricularium (Conil, 1878) (Fig. 2)Idiosoma. Length from apices of scapula to posterior body mar-

gin 1.256 ± 0.076 (1.107–1.333), maximum breadth 0.951 ± 0.057(0.816–1.007); outline oval, slightly longilinear, with 11 festoonswithout tubercles. Scutum. Length 0.629 ± 0.043 (0.525–0.664),breadth 0.767 ± 0.039 (0.679–0.812), breadth/length ratio1.221 ± 0.029 (1.184–1.294), inornate; few medium punctations

T.F. Martins et al. / Ticks and Tick-borne Diseases 1 (2010) 75–99 77

Fig. 1. Amblyomma aureolatum nymph. (A) Dorsal gnathosoma (capitulum) (bar: 20 �m). (B) Ventral gnathosoma (bar: 20 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV(bar: 30 �m).

Fig. 2. Amblyomma auricularium nymph. (A) Dorsal gnathosoma (capitulum) (bar: 20 �m). (B) Ventral gnathosoma (bar: 20 �m). (C) Scutum (bar: 30 �m). (D) Coxae I–IV(bar: 30 �m).


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ig. 3. Amblyomma calcaratum nymph. (A) Dorsal gnathosoma (capitulum) (bar: 300 �m).

venly distributed, although deeper laterally. Eyes not orbitedt lateral scutal angles at the level of scutal midlength. Cervicalrooves deep, converging anteriorly then diverging as shallowepressions at the scutal median third. Spiracular plate triangularith rounded angles, with an evident dorsal prolongation; length

.053 ± 0.001 (0.050–0.055), breadth 0.028 ± 0.004 (0.023–0.035).nathosoma (capitulum). Length from palpal apices to poste-ior margin 0.244 ± 0.021 (0.202–0.271), breadth 0.267 ± 0.0150.241–0.284). Basis capituli rectangular, posterior margin slightlyoncave, without cornua; posterior margin convex ventrally,ithout auriculae. Palpi length 0.184 ± 0.017 (0.149–0.206), arti-

le I with a small ventral prolongation, article II 0.125 ± 0.0150.104–0.143) long, article III 0.045 ± 0.005 (0.033–0.053) long.ypostome rounded apically; length 0.188 ± 0.015 (0.154–0.208);

ength of toothed portion 0.107 ± 0.008 (0.088–0.119); dentition/2 with 6–7 teeth per row. Legs. Coxa I with 2 slender spurs, thexternal slightly longer; coxae II–IV with a small triangular spur.rochanters without spur; tarsus I 0.312 ± 0.026 (0.257–0.337)ong, 0.088 ± 0.008 (0.078–0.098) broad; tarsus IV 0.238 ± 0.0180.217–0.270) long, 0.070 ± 0.006 (0.060–0.079) broad.

mblyomma calcaratum Neumann, 1899 (Fig. 3)Idiosoma. Length from apices of scapula to posterior body mar-

in 1.432 ± 0.065 (1.281–1.525), maximum breadth 1.177 ± 0.0411.075–1.230), outline oval, with 11 festoons without tubercles.cutum. Length 0.658 ± 0.019 (0.624–0.683), breadth 0.829 ± 0.0400.786–0.928), breadth/length ratio 1.250 ± 0.010 (1.238–1.261),
nornate, surface extensively shagreened (rugose); medium punc-ations evenly distributed, deeper laterally. Eyes not orbited atateral scutal angles at the level of scutal midlength. Cervicalrooves long, reaching the scutal posterior third, deeper anteriorly.piracular plate triangular with rounded angles, with a discrete
(B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 30 �m). (D) Coxae I–IV (bar:

dorsal prolongation; length 0.066 ± 0.006 (0.053–0.075), breadth0.043 ± 0.006 (0.032–0.051). Gnathosoma (capitulum). Length frompalpal apices to posterior margin 0.262 ± 0.023 (0.208–0.300),breadth 0.276 ± 0.007 (0.262–0.289). Basis capituli sub-triangular,without cornua, although the posterior margin concave givesthe aspect of minute cornua at the posterolateral margins; pos-terior margin convex ventrally, without auriculae. Palpi length0.211 ± 0.016 (0.182–0.237), article I without ventral prolon-gation, article II 0.132 ± 0.018 (0.095–0.155) long, article III0.054 ± 0.004 (0.046–0.058) long. Hypostome rounded apically;length 0.189 ± 0.010 (0.176–0.206); length of toothed portion0.115 ± 0.006 (0.106–0.125); dentition 2/2 with 6–7 teeth perrow. Legs. Coxa I with 2 triangular spurs, the external longer andpointed; coxae II–IV with a small triangular spur. Trochanters with-out spur; tarsus I 0.402 ± 0.011 (0.391–0.419) long, 0.119 ± 0.012(0.103–0.147) broad; tarsus IV 0.287 ± 0.019 (0.263–0.316) long,0.088 ± 0.007 (0.078–0.102) broad.

Amblyomma coelebs Neumann, 1899 (Fig. 4)Idiosoma. Length from apices of scapula to posterior body mar-

gin 1.568 ± 0.055 (1.492–1.665), maximum breadth 1.309 ± 0.043(1.240–1.374), outline oval, with 11 festoons without tubercles.Scutum. Length 0.757 ± 0.033 (0.704–0.817), breadth 1.008 ± 0.034(0.942–1.051), breadth/length ratio 1.332 ± 0.019 (1.285–1.360);inornate, numerous large and deep punctations evenly distributed.Eyes not orbited at lateral scutal angles at the level of scu-tal midlength. Cervical grooves deep and short, ending as small
shallow depressions at the level of the posterior margin of theeyes. Spiracular plate triangular with rounded angles, with adiscrete dorsal prolongation; length 0.077 ± 0.007 (0.068–0.088),breadth 0.051 ± 0.008 (0.039–0.064). Gnathosoma (capitulum).Length from palpal apices to posterior margin 0.329 ± 0.033


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ig. 4. Amblyomma coelebs nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m0 �m).

0.274–0.386), breadth 0.361 ± 0.018 (0.336–0.394). Basis capit-li slightly hexagonal, posterior margin slightly concave, withoutornua; posterior margin convex ventrally, without auriculae.alpi length 0.258 ± 0.017 (0.236–0.285), article I with vestigialentral prolongation; article II 0.161 ± 0.013 (0.143–0.188) long,rticle III 0.076 ± 0.005 (0.067–0.083) long. Hypostome roundedpically; length 0.269 ± 0.014 (0.250–0.296); length of toothed por-ion 0.132 ± 0.010 (0.118–0.144); dentition 2/2 with 7–8 teeth perow. Legs. Coxa I with 2 pointed spurs, separated to each othery a space larger than one spur; the internal spur broader andhe external slightly longer; coxae II–IV with a small triangularpur, emerged from a ridge-like medial extension in coxae II andII. Trochanters without spur; tarsus I 0.468 ± 0.053 (0.331–0.521)ong, 0.108 ± 0.008 (0.088–0.117) broad, tarsus IV 0.343 ± 0.0140.315–0.361) long, 0.086 ± 0.005 (0.077–0.093) broad.

mblyomma fuscum Neumann, 1907 (Fig. 5)Idiosoma. Length from apices of scapula to posterior body mar-

in 1.654 ± 0.082 (1.498–1.762), maximum breadth 1.350 ± 0.0671.201–1.444), outline oval, with 11 festoons without tubercles.cutum. Length 0.840 ± 0.034 (0.806–0.900), breadth 1.024 ± 0.0520.906–1.082), breadth/length ratio 1.109 ± 0.035 (1.081–1.194),nornate; small and shallow punctations evenly distributed; pos-erolateral margin somewhat sinuous. Eyes not orbited at lateralcutal angles at the level of scutal midlength. Cervical groovesong, reaching the scutal posterior third, deeper at the anteriorhird. Spiracular plate triangular with rounded angles, with an
vident dorsal prolongation; length 0.090 ± 0.008 (0.095–0.104),readth 0.057 ± 0.007 (0.053–0.068). Gnathosoma (capitulum).ength from palpal apices to posterior margin 0.382 ± 0.0370.337–0.450), breadth 0.359 ± 0.021 (0.311–0.387). Basis capit-li pentagonal; posterior margin slightly concave, without cornua;
Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV (bar:

posterior margin convex ventrally, without auriculae. Palpi length0.296 ± 0.024 (0.267–0.340), article I without ventral prolongation;article II 0.214 ± 0.022 (0.192–0.264) long, article III 0.060 ± 0.005(0.048–0.068) long, article IV apically projected from article III.Hypostome rounded apically; length 0.295 ± 0.021 (0.268–0.330);length of toothed portion 0.142 ± 0.008 (0.128–0.153); dentition2/2 with 7–8 teeth per row. Legs. Coxa I with 2 sub-equal trian-gular spurs, the external slightly stouter; coxae II–IV with a smalltriangular spur similar to the coxa I external spur. Trochanters with-out spur; tarsus I 0.439 ± 0.021 (0.407–0.477) long, 0.109 ± 0.010(0.095–0.135) broad; tarsus IV 0.339 ± 0.032 (0.279–0.388) long,0.089 ± 0.011 (0.073–0.104) broad.

Amblyomma humerale Koch, 1844 (Fig. 6)Idiosoma. Length from apices of scapula to posterior body mar-

gin 1.519 ± 0.100 (1.336–1.670), maximum breadth 1.292 ± 0.086(1.125–1.428), outline oval, with 11 festoons without tubercles.Scutum. Length 0.743 ± 0.042 (0.677–0.791), breadth 0.970 ± 0.051(0.859–1.028), breadth/length ratio 1.305 ± 0.022 (1.269–1.347),inornate, numerous large and deep punctations evenly distributed.Eyes not orbited at lateral scutal angles at the level of scutalmidlength. Cervical grooves deep, reaching the scutal midlength.Spiracular plate triangular with rounded angles, with a dis-crete dorsal prolongation; length 0.072 ± 0.006 (0.059–0.080),breadth 0.048 ± 0.008 (0.034–0.061). Gnathosoma (capitulum).Length from palpal apices to posterior margin 0.360 ± 0.036(0.296–0.413), breadth 0.332 ± 0.014 (0.309–0.348). Basis capituli
sub-triangular, posterior margin slightly concave, without cor-nua; posterior margin convex ventrally, without auriculae. Palpilength 0.272 ± 0.022 (0.234–0.310), article I without ventral pro-longation; article II 0.184 ± 0.014 (0.162–0.204) long, article III:0.066 ± 0.008 (0.053–0.080) long. Hypostome rounded apically;


Fig. 5. Amblyomma fuscum nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m). (B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV (bar:30 �m).

Fig. 6. Amblyomma humerale nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m). (B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV (bar:30 �m).


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ig. 7. Amblyomma incisum nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m0 �m). (E) Ventral festoons showing tubercles at the posterointernal angles (bar: 2

ength 0.256 ± 0.013 (0.233–0.276); length of toothed portion.123 ± 0.006 (0.112–0.131); dentition 2/2 with 7–8 teeth per row.egs. Coxa I with 2 triangular spurs, the external slightly longer;oxae II–III with 2 small spurs, the external triangular and longer,he internal rounded and almost obsolete, especially in coxa III.oxa IV with a small triangular spur. Trochanters without spur; tar-us I 0.409 ± 0.035 (0.368–0.465) long, 0.108 ± 0.008 (0.096–0.125)road; tarsus IV 0.297 ± 0.028 (0.242–0.332) long, 0.099 ± 0.0130.085–0.128) broad.

mblyomma incisum Neumann, 1906 (Fig. 7)Idiosoma. Length from apices of scapula to posterior body mar-

1.510–1.683), outline oval, with 11 festoons with well-developed,hitish tubercles at the posterointernal angles. Scutum. Length
.872 ± 0.031 (0.825–0.917), breadth 1.174 ± 0.060 (1.108–1.295),readth/length ratio 1.345 ± 0.029 (1.310–1.412), inornate; fewedium and shallow punctations evenly distributed, deeper lat-

Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV (bar:).

erally; posterolateral margin somewhat sinuous. Eyes not orbited,slightly convex, at lateral scutal angles at the level of scu-tal midlength. Cervical grooves short and deep at the anteriorthird, then diverging posteriorly as shallow depressions. Spirac-ular plate triangular with rounded angles, with an evidentdorsal prolongation; length 0.082 ± 0.009 (0.070–0.102), breadth0.050 ± 0.005 (0.044–0.064). Gnathosoma (capitulum). Length frompalpal apices to cornua apices 0.374 ± 0.047 (0.318–0.452), breadth0.411 ± 0.016 (0.383–0.431). Basis capituli rectangular, posteriormargin slightly concave, with broad triangular cornua; poste-rior margin convex ventrally, without auriculae. Palpi length0.273 ± 0.027 (0.239–0.315), article I with vestigial ventral pro-longation; article II 0.169 ± 0.022 (0.144–0.207) long, article III
0.085 ± 0.007 (0.076–0.097) long. Hypostome rounded apically;length 0.307 ± 0.014 (0.287–0.330); length of toothed portion0.156 ± 0.006 (0.149–0.170); dentition 2/2 with 8–9 teeth per row.Legs. Coxa I with 2 separated stout spurs, the external longerbut less than twice as long; coxae II–IV with a small triangular


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ig. 8. Amblyomma latepunctatum nymph. (A) Dorsal gnathosoma (capitulum) (barbar: 100 �m).

pur, slightly shorter in coxa IV. Trochanters without spur; tar-us I 0.499 ± 0.051 (0.424–0.568) long, 0.130 ± 0.006 (0.118–0.137)road; tarsus IV 0.386 ± 0.041 (0.349–0.461) long, 0.119 ± 0.0080.100–0.130) broad.

mblyomma latepunctatum Tonelli-Rondelli, 1939 (Fig. 8)Idiosoma. Length from apices of scapula to posterior body mar-

in 1.780 ± 0.121 (1.575–1.958), maximum breadth 1.426 ± 0.1291.197–1.561); outline oval, with 11 festoons without tubercles.cutum. Length 0.896 ± 0.043 (0.828–0.954), breadth 1.122 ± 0.0750.972–1.230), breadth/length ratio 1.251 ± 0.032 (1.172–1.288),nornate; medium and shallow punctations evenly distributed.yes not orbited, slightly convex, at lateral scutal angles at theevel of scutal midlength. Cervical grooves short and deep at thenterior third, then diverging posteriorly as shallow depressionsnding at the posterolateral margin. Spiracular plate triangularith rounded angles, with a discrete dorsal prolongation; length

.079 ± 0.005 (0.071–0.089), breadth 0.046 ± 0.003 (0.041–0.052).nathosoma (capitulum). Length from palpal apices to cornua apices.391 ± 0.037 (0.330–0.462), breadth 0.376 ± 0.015 (0.336–0.393).asis capituli rectangular, posterior margin slightly concave,ith triangular cornua; posterior margin convex ventrally, with-

ut auriculae. Palpi length 0.288 ± 0.024 (0.253–0.340), articlewith vestigial ventral prolongation; article II 0.181 ± 0.024

0.148–0.228) long, article III 0.081 ± 0.004 (0.072–0.091) long.ypostome rounded apically; length 0.286 ± 0.016 (0.264–0.306);

ength of toothed portion 0.146 ± 0.011 (0.133–0.165); dentition
/2 with 8–9 teeth per row. Legs. Coxa I with 2 stout spurs sep-rated by a space shorter than the breadth of the external spurase, the internal spur stouter, the external slender and longerut less than twice as long; coxae II–IV with a small triangularpur, slightly shorter in coxa IV. Trochanters without spur; tar-
). (B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV

sus I 0.529 ± 0.031 (0.467–0.568) long, 0.130 ± 0.008 (0.122–0.150)broad; tarsus IV 0.384 ± 0.038 (0.321–0.419) long, 0.100 ± 0.005(0.092–0.106) broad.

Amblyomma naponense (Packard, 1869) (Fig. 9)Idiosoma. Length from apices of scapula to posterior body mar-

gin 1.624 ± 0.104 (1.486–1.837), maximum breadth 1.308 ± 0.049(1.223–1.406), outline oval, with 11 festoons without tubercles.Scutum. Length 0.728 ± 0.047 (0.638–0.789), breadth 0.963 ± 0.046(0.902–1.045), breadth/length ratio 1.324 ± 0.038 (1.287–1.413),inornate; few small punctations evenly distributed, deeper inthe shallow depressions of the cervical grooves. Eyes notorbited, slightly convex, at lateral scutal angles at the levelof scutal midlength. Cervical grooves deep at the anteriorthird, then diverging posteriorly as shallow depressions end-ing at the posterolateral margin. Spiracular plate triangular withrounded angles, with a discrete dorsal prolongation; length0.069 ± 0.004 (0.062–0.075), breadth 0.043 ± 0.006 (0.034–0.053).Gnathosoma (capitulum). Length from palpal apices to cornua apices0.370 ± 0.064 (0.281–0.458), breadth 0.422 ± 0.018 (0.390–0.441).Basis capituli hexagonal, posterior margin straight with pro-nounced triangular cornua; posterior margin convex ventrally,without auriculae. Palpi length 0.261 ± 0.037 (0.201–0.320), articleI with a pronounced ventral prolongation; article II 0.165 ± 0.025(0.120–0.202) long, article III 0.077 ± 0.011 (0.063–0.096) long.Hypostome rounded apically; length 0.268 ± 0.022 (0.221–0.298);length of toothed portion 0.129 ± 0.012 (0.115–0.149); dentition2/2 with 9–10 teeth per row. Legs. Coxa I with 2 spurs, the exter-
nal long, the internal short and robust; coxae II–IV with a smalltriangular spur that gradually decreases in size from II to IV.Trochanters without spur; tarsus I 0.462 ± 0.055 (0.365–0.532)long, 0.139 ± 0.013 (0.119–0.160) broad; tarsus IV 0.325 ± 0.037(0.246–0.383) long, 0.096 ± 0.005 (0.088–0.109) broad.


F m). (3

A

g(S(iptlaw0Gr(sta(Hl2tro(0

A

g

ig. 9. Amblyomma naponense nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �0 �m).

mblyomma nodosum Neumann, 1899 (Fig. 10)Idiosoma. Length from apices of scapula to posterior body mar-

in 1.279 ± 0.063 (1.201–1.414), maximum breadth 1.054 ± 0.0670.967–1.197), outline oval, with 11 festoons without tubercles.cutum. Length 0.554 ± 0.036 (0.498–0.600), breadth 0.776 ± 0.0440.712–0.843), breadth/length ratio 1.400 ± 0.029 (1.358–1.451),nornate, surface extensively shagreened (rugose); few mediumunctations evenly distributed. Eyes not orbited at lateral scu-al angles at the level of scutal midlength. Cervical groovesong, deep at the scutal anterior third, shallow at the mediumnd posterior thirds of the scutum. Spiracular plate triangularith rounded angles, with a discrete dorsal prolongation; length

.061 ± 0.006 (0.053–0.073), breadth 0.037 ± 0.006 (0.028–0.048).nathosoma (capitulum). Length from palpal apices to poste-ior margin 0.238 ± 0.037 (0.171–0.292), breadth 0.270 ± 0.0220.219–0.296). Basis capituli sub-triangular, posterior marginlightly concave, without cornua; posterior margin convex ven-rally, without auriculae. Palpi length 0.191 ± 0.028 (0.133–0.226),rticle I without ventral prolongation; article II 0.110 ± 0.0190.071–0.135) long, article III 0.060 ± 0.008 (0.044–0.072) long.ypostome rounded apically; length 0.164 ± 0.015 (0.140–0.186);

ength of toothed portion 0.090 ± 0.011 (0.068–0.106); dentition/2 with 6–7 teeth per row. Legs. Coxa I with 2 triangular spurs,he external slightly longer and pointed, the internal short andobust; coxae II–IV with a small triangular spur. Trochanters with-ut spur; tarsus I 0.313 ± 0.064 (0.201–0.383) long, 0.099 ± 0.0170.075–0.132) broad; tarsus IV 0.245 ± 0.025 (0.209–0.296) long;.068 ± 0.009 (0.051–0.081) broad.

mblyomma ovale Koch, 1844 (Fig. 11)Idiosoma. Length from apices of scapula to posterior body mar-


B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV (bar:

(0.859–0.956), outline oval, longilinear, with 11 festoons with-out tubercles. Scutum. Length 0.635 ± 0.018 (0.606–0.668), breadth0.714 ± 0.028 (0.659–0.754), breadth/length ratio 1.123 ± 0.019(1.087–1.148), inornate; few punctations, larger and deeper lat-erally. Eyes not orbited at lateral scutal angles at the level ofthe posterior third of the scutum. Cervical grooves long, reach-ing the scutal posterior fourth, deeper in the anterior fourth.Spiracular plate triangular with rounded angles, with a dis-crete dorsal prolongation; length 0.055 ± 0.005 (0.050–0.068),breadth 0.030 ± 0.003 (0.025–0.035). Gnathosoma (capitulum).Length from palpal apices to posterior margin 0.300 ± 0.023(0.269–0.348), breadth 0.360 ± 0.013 (0.328–0.376). Basis capit-uli triangular, posterior margin slightly concave, without cornua;posterior margin convex ventrally, auriculae as small posterolat-eral rounded projections, lateral margin projected laterally. Palpilength 0.230 ± 0.011 (0.213–0.246), article I without ventral pro-longation; article II 0.159 ± 0.019 (0.137–0.206) long, article III0.055 ± 0.006 (0.041–0.062) long. Hypostome rounded apically;length 0.219 ± 0.007 (0.210–0.233); length of toothed portion0.127 ± 0.006 (0.116–0.134); dentition 2/2 with 7–8 teeth per row.Legs. Coxa I with 2 stout spurs, the external longer and broader;coxae II–IV with a small triangular spur, which is rudimentarin coxae III–IV. Trochanters without spur; tarsus I 0.388 ± 0.019(0.357–0.412) long, 0.108 ± 0.013 (0.095–0.135) broad; tarsusIV 0.281 ± 0.031 (0.228–0.320) long, 0.079 ± 0.004 (0.073–0.091)broad.

Amblyomma pacae Aragão, 1911 (Fig. 12)Idiosoma. Length from apices of scapula to posterior body mar-

gin 1.566 ± 0.095 (1.345–1.706), maximum breadth 1.297 ± 0.076(1.136–1.395); outline oval, with 11 festoons without tubercles.Scutum. Length 0.741 ± 0.039 (0.673–0.786), breadth 1.037 ± 0.056


Fig. 10. Amblyomma nodosum nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m). (B) Ventral gnathosoma (bar: 20 �m). (C) Scutum (bar: 30 �m). (D) Coxae I–IV (bar:30 �m).

Fig. 11. Amblyomma ovale nymph. (A) Dorsal gnathosoma (capitulum) (bar: 20 �m). (B) Ventral gnathosoma (bar: 20 �m). (C) Scutum (bar: 30 �m). (D) Coxae I–IV (bar:30 �m).


F ). (B)1

(ilaaew0Gr(cwc(Hl2sa(Ib

A

g(S(ia

ig. 12. Amblyomma pacae nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m00 �m).

0.967–1.114), breadth/length ratio 1.398 ± 0.029 (1.348–1.435),nornate, numerous large punctations evenly distributed, deeperaterally, smaller centrally. Eyes large, not orbited, at lateral scutalngles at the level of scutal midlength. Cervical grooves long, deepert the scutal anterior third, followed by a rugose shallow depressionnding forward of the level of the eyes. Spiracular plate triangularith rounded angles, with a discrete dorsal prolongation; length

.086 ± 0.009 (0.066–0.098), breadth 0.053 ± 0.008 (0.039–0.068).nathosoma (capitulum). Length from palpal apices to poste-ior margin 0.327 ± 0.025 (0.281–0.374); breadth 0.337 ± 0.0110.316–0.356). Basis capituli rectangular, posterior margin slightlyoncave, without cornua; posterior margin convex ventrally,ithout auriculae. Palpi length 0.244 ± 0.017 (0.212–0.273), arti-

le I with vestigial ventral prolongation; article II 0.161 ± 0.0100.147–0.182) long, article III 0.065 ± 0.006 (0.051–0.072) long.ypostome rounded apically; length 0.246 ± 0.021 (0.212–0.280);

ength of toothed portion 0.128 ± 0.009 (0.116–0.143); dentition/2 with 7–8 teeth per row. Legs. Coxa I with 2 slender pointedpurs, the external slightly longer; coxae II–IV with a small tri-ngular spur. Trochanters without spur; tarsus I 0.446 ± 0.0270.401–0.483) long, 0.114 ± 0.010 (0.097–0.134) broad; tarsusV 0.317 ± 0.019 (0.285–0.351) long, 0.088 ± 0.008 (0.071–0.098)road.

mblyomma pseudoconcolor Aragão, 1908 (Fig. 13)Idiosoma. Length from apices of scapula to posterior body mar-

1.043–1.298); outline oval, with 11 festoons without tubercles.cutum. Length 0.689 ± 0.037 (0.637–0.734), breadth 0.936 ± 0.0690.866–1.045), breadth/length ratio 1.356 ± 0.036 (1.322–1.422),nornate; rare punctations. Eyes large, not orbited, at lateral scutalngles at the level of scutal midlength. Cervical grooves long, reach-

ing the scutal posterolateral margin, deeper at the anterior third.Spiracular plate triangular with rounded angles, with an evidentdorsal prolongation; length 0.063 ± 0.007 (0.053–0.073), breadth0.038 ± 0.006 (0.030–0.044). Gnathosoma (capitulum). Length frompalpal apices to posterior margin 0.270 ± 0.029 (0.234–0.317);breadth 0.314 ± 0.013 (0.302–0.339). Basis capituli rectangular,posterior margin slightly convex, without cornua; posterior mar-gin convex ventrally, without auriculae. Palpi length 0.208 ± 0.023(0.180–0.247), article I with a small ventral prolongation, article II0.133 ± 0.008 (0.125–0.144), article III 0.059 ± 0.010 (0.041–0.073).Hypostome rounded apically; length 0.176 ± 0.016 (0.151–0.199);length of toothed portion 0.098 ± 0.018 (0.075–0.127); dentition2/2 with 6–7 teeth per row. Legs. Coxa I with 2 slender spurs,the external longer; coxae II–IV with a small triangular spur.Trochanters without spur; tarsus I 0.302 ± 0.036 (0.241–0.348)long, 0.091 ± 0.011 (0.074–0.106) broad; tarsus IV 0.265 ± 0.028(0.221–0.298) long, 0.077 ± 0.008 (0.067–0.091) broad.

Amblyomma scalpturatum Neumann, 1906 (Fig. 14)Idiosoma. Length from apices of scapula to posterior body mar-

gin 1.695 ± 0.202 (1.425–2.098), maximum breadth 1.306 ± 0.174(1.071–1.615), outline oval, with 11 festoons with discrete,brownish tubercles at the posterointernal angles. Scutum. Length0.790 ± 0.086 (0.751–0.944), breadth 0.988 ± 0.121 (0.827–1.205),breadth/length ratio 1.248 ± 0.028 (1.196–1.288), inornate;medium and shallow punctations evenly distributed; posterolat-eral margin not sinuous. Eyes not orbited, slightly convex, at lateral
scutal angles at the level of scutal midlength. Cervical groovesshort and deep at the anterior third, then diverging posteriorlyas shallow depressions. Spiracular plate triangular with roundedangles, with a discrete dorsal prolongation; length 0.074 ± 0.007(0.061–0.088), breadth 0.044 ± 0.005 (0.032–0.053). Gnatho-


F ar: 20I

s0Bwwa(Hl2tcIl(

A

g(S(iemtw0Gr(s

ig. 13. Amblyomma pseudoconcolor nymph. (A) Dorsal gnathosoma (capitulum) (b–IV (bar: 30 �m).

oma (capitulum). Length from palpal apices to cornua apices.393 ± 0.032 (0.346–0.455), breadth 0.367 ± 0.037 (0.302–0.431).asis capituli rectangular, posterior margin slightly concave,ith broad triangular cornua; posterior margin convex ventrally,ithout auriculae. Palpi length 0.267 ± 0.019 (0.237–0.300),

rticle I without ventral prolongation; article II: 0.168 ± 0.0180.144–0.209) long, article III: 0.075 ± 0.006 (0.062–0.083) long.ypostome rounded apically; length 0.274 ± 0.021 (0.232–0.312);

ength of toothed portion 0.138 ± 0.009 (0.117–0.151); dentition/2 with 8–9 teeth per row. Legs. Coxa I with 2 separated spurs,he internal broader, the external slender about twice as long;oxae II–IV with a pointed triangular spur, slightly shorter in coaxV. Trochanters without spur; tarsus I 0.501 ± 0.053 (0.391–0.602)ong, 0.130 ± 0.007 (0.119–0.140) broad; tarsus IV 0.371 ± 0.0480.297–0.473), 0.097 ± 0.009 (0.084–0.112) broad.

mblyomma varium Koch, 1844 (Fig. 15)Idiosoma. Length from apices of scapula to posterior body mar-

in 1.660 ± 0.085 (1.472–1.946), maximum breadth 1.313 ± 0.1591.169–1.550); outline oval, with 11 festoons without tubercles.cutum. Length 0.728 ± 0.045 (0.699–0.808), breadth 0.970 ± 0.1020.875–1.116), breadth/length ratio 1.329 ± 0.076 (1.252–1.436),nornate, numerous deep punctations evenly distributed, larger lat-rally. Eyes not orbited at lateral scutal angles at the level of scutalidlength. Cervical grooves short and slightly deep at the scu-

al anterior third, shallow posteriorly. Spiracular plate triangularith rounded angles, with a discrete dorsal prolongation; length

.096 ± 0.010 (0.088–0.115), breadth 0.060 ± 0.004 (0.055–0.068).nathosoma (capitulum). Length from palpal apices to poste-ior margin 0.328 ± 0.035 (0.271–0.368), breadth 0.306 ± 0.0210.282–0.342). Basis capituli slightly hexagonal, posterior marginlightly concave, without cornua; posterior margin convex ven-

�m). (B) Ventral gnathosoma (bar: 20 �m). (C) Scutum (bar: 100 �m). (D) Coxae

trally, without auriculae. Palpi length 0.273 ± 0.028 (0.243–0.318),article I with vestigial ventral prolongation, article II 0.168 ± 0.028(0.151–0.218) long, article III 0.087 ± 0.013 (0.068–0.106) long.Hypostome rounded apically; length 0.215 ± 0.018 (0.203–0.246);length of toothed portion 0.108 ± 0.011 (0.100–0.127); dentition2/2 with 6–7 teeth per row. Legs. Coxa I with 2 pointed spurs,the internal short and triangular, the external about twice aslong; coxae II–IV with a small triangular spur that graduallydecreases in size from II to IV. Trochanters without spur; tar-sus I 0.407 ± 0.047 (0.360–0.480) long, 0.112 ± 0.010 (0.097–0.123)broad; tarsus IV 0.320 ± 0.017 (0.298–0.339) long, 0.080 ± 0.016(0.062–0.098) broad.

Redescriptions

Amblyomma brasiliense Aragão, 1908 (Fig. 16)Aragão (1908): description of the nymph.Sanches et al. (1908): redescription of the nymph.Idiosoma. Length from apices of scapula to posterior

body margin 1.652 ± 0.067 (1.563–1.745), maximum breadth1.286 ± 0.047 (1.212–1.388), outline oval, with 11 festoonswith tubercles at the posterointernal angles. Scutum. Length0.678 ± 0.044 (0.604–0.743), breadth 0.897 ± 0.023 (0.858–0.926),breadth/length ratio 1.326 ± 0.054 (1.246–1.420), inornate; fewmedium and shallow punctations evenly distributed, deeper later-ally. Eyes not orbited, slightly convex, at lateral scutal angles at thelevel of scutal midlength. Cervical grooves broad and long, cover-
ing the first 2 thirds of the scutum, deeper at the anterior third.Spiracular plate triangular with rounded angles, with a discretedorsal prolongation; length 0.065 ± 0.003 (0.059–0.070), breadth0.031 ± 0.003 (0.025–0.037). Gnathosoma (capitulum). Lengthfrom palpal apices to cornua apices 0.295 ± 0.031 (0.251–0.350),


F 30 �m( bar: 3

bpp0l0l0rssIb(

A

ig. 14. Amblyomma scalpturatum nymph. (A) Dorsal gnathosoma (capitulum) (bar:bar: 30 �m). (E) Ventral festoons showing tubercles at the posterointernal angles (

readth 0.335 ± 0.010 (0.316–0.348). Basis capituli rectangular,osterior margin slightly concave, with broad triangular cornua;osterior margin convex ventrally, without auriculae. Palpi length.210 ± 0.019 (0.179–0.244), article I with a small ventral pro-

ongation; article II 0.133 ± 0.017 (0.112–0.165) long, article III.062 ± 0.004 (0.055–0.072) long. Hypostome rounded apically;

ength 0.205 ± 0.014 (0.182–0.228); length of toothed portion.105 ± 0.008 (0.093–0.117); dentition 2/2 with 8–9 teeth perow. Legs. Coxa I with 2 spurs, the internal broader, the externallender about twice as long; coxae II–IV with a pointed triangularpur, slightly shorter in coxa IV. Trochanters without spur; tarsus0.391 ± 0.078 (0.218–0.462) long, 0.133 ± 0.014 (0.115–0.150)road; tarsus IV 0.323 ± 0.041 (0.253–0.371) long, 0.092 ± 0.008
0.079–0.107) broad.
mblyomma cajennense (Fabricius, 1787) (Fig. 17)Cooley and Kohls (1944): description of the nymph.

). (B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV0 �m).

Idiosoma. Length from apices of scapula to posterior body mar-gin 1.282 ± 0.056 (1.218–1.363); maximum breadth 1.041 ± 0.038(0.985–1.082); outline oval, with 11 festoons without tubercles.Scutum. Length 0.574 ± 0.048 (0.489–0.633), breadth 0.819 ± 0.033(0.774–0.884), breadth/length ratio 1.433 ± 0.082 (1.374–1.595),inornate; large and deep punctations evenly distributed. Eyes notorbited at lateral scutal angles at the level of scutal midlength.Cervical grooves deep in the scutal anterior third, followedby a rugose shallow depression in the scutal median third.Spiracular plate triangular with rounded angles, with an evi-dent dorsal prolongation; length 0.071 ± 0.007 (0.059–0.084),breadth 0.045 ± 0.004 (0.035–0.052). Gnathosoma (capitulum).Length from palpal apices to posterior margin 0.293 ± 0.049
(0.222–0.357), breadth 0.282 ± 0.008 (0.267–0.294). Basis capit-uli rectangular, posterior margin slightly concave, without cornua;posterior margin convex ventrally, without auriculae. Palpi length0.222 ± 0.035 (0.161–0.265), article I with a vestigial ventral pro-longation, article II 0.141 ± 0.026 (0.100–0.170) long, article III


F m). (B3

0l0reewIb(

A

g(S(itoCodbL(sp0l0

ig. 15. Amblyomma varium nymph. (A) Dorsal gnathosoma (capitulum) (bar: 20 �0 �m).

.063 ± 0.011 (0.045–0.079) long. Hypostome rounded apically;ength 0.242 ± 0.013 (0.219–0.262); length of toothed portion.127 ± 0.006 (0.115–0.138); dentition 2/2 with 7–8 teeth perow. Legs. Coxa I with 2 pointed spurs separated by a spacequal or greater than the breadth of the external spur base, thexternal spur about twice as long than the internal; coxae II–IVith a small triangular spur. Trochanters without spur; tarsus0.362 ± 0.065 (0.223–0.425) long, 0.101 ± 0.006 (0.092–0.113)road; tarsus IV 0.259 ± 0.033 (0.211–0.302) long, 0.070 ± 0.0040.061–0.080) broad.

mblyomma dissimile Koch, 1844 (Fig. 18)Cooley and Kohls (1944): description of the nymph.Idiosoma. Length from apices of scapula to posterior body mar-

in 1.622 ± 0.100 (1.403–1.775), maximum breadth 1.306 ± 0.0801.115–1.413), outline oval, with 11 festoons without tubercles.cutum. Length 0.841 ± 0.038 (0.771–0.886), breadth 1.054 ± 0.0440.943–1.104), breadth/length ratio 1.253 ± 0.024 (1.223–1.298),nornate, punctations moderate in number, larger and deeper inhe lateral fields, posterolateral margin slightly sinuous. Eyes notrbited at lateral scutal angles at the level of scutal midlength.ervical grooves reaching the scutal midlength, deep anteri-rly. Spiracular plate triangular with rounded angles, with aiscrete dorsal prolongation; length 0.093 ± 0.005 (0.084–0.102),readth 0.058 ± 0.009 (0.043–0.070). Gnathosoma (capitulum).ength from palpal apices to posterior margin 0.428 ± 0.0450.329–0.474), breadth 0.377 ± 0.017 (0.338–0.404). Basis capituli
ub-triangular, posterior margin slightly convex, without cornua;osterior margin straight ventrally, without auriculae. Palpi length.312 ± 0.026 (0.264–0.342), article I with an evident ventral pro-
ongation; article II 0.207 ± 0.021 (0.167–0.234) long, article III.084 ± 0.007 (0.067–0.094) long. Hypostome rounded apically;

) Ventral gnathosoma (bar: 20 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV (bar:

length 0.282 ± 0.015 (0.255–0.304); length of toothed portion0.141 ± 0.010 (0.120–0.156); dentition 2.5/2.5 (3/3 apically, 2/2at the base) with 7–8 teeth in external and median rows, 4–5teeth in internal row. Legs. Coxa I with 2 separated spurs, theinternal short and triangular, the external longer; coxae II–IVwith a small triangular spur. Trochanters without spur; tarsusI 0.455 ± 0.024 (0.414–0.480) long, 0.099 ± 0.008 (0.087–0.112)broad; tarsus IV 0.337 ± 0.028 (0.271–0.380) long, 0.074 ± 0.004(0.067–0.080) broad.

Amblyomma dubitatum Neumann, 1899 (Fig. 19)Estrada-Pena et al. (2002): description of the nymph.Idiosoma. Length from apices of scapula to posterior body mar-

gin 1.699 ± 0.072 (1.577–1.800), maximum breadth 1.352 ± 0.048(1.269–1.435); outline oval, with 11 festoons without tubercles.Scutum. Length 0.794 ± 0.029 (0.736–0.840), breadth 1.002 ± 0.028(0.972–1.053), breadth/length ratio 1.261 ± 0.024 (1.235–1.319);inornate, deep punctations evenly distributed, larger laterally,smaller centrally. Eyes large, not orbited, at lateral scutal angles atthe level of scutal midlength. Cervical grooves long and deep reach-ing the scutal posterior third. Spiracular plate rounded with anevident dorsal prolongation; length 0.095 ± 0.005 (0.084–0.107),breadth 0.054 ± 0.003 (0.046–0.059). Gnathosoma (capitulum).Length from palpal apices to posterior margin 0.352 ± 0.044(0.264–0.418); breadth 0.360 ± 0.011 (0.336–0.372). Basis capitulislightly hexagonal, posterior margin slightly concave, without cor-nua; posterior margin convex ventrally, without auriculae. Palpi
length 0.269 ± 0.027 (0.214–0.321), article I with vestigial ventralprolongation; article II 0.179 ± 0.017 (0.143–0.206) long, articleIII 0.070 ± 0.008 (0.055–0.085) long. Hypostome rounded api-cally; length 0.256 ± 0.023 (0.212–0.285); length of toothed portion0.139 ± 0.011 (0.120–0.154); dentition 2/2 with 7–8 teeth per row.


F 0 �m( bar: 2

LIsb(

A

o

g(S(itar

ig. 16. Amblyomma brasiliense nymph. (A) Dorsal gnathosoma (capitulum) (bar: 2bar: 30 �m). (E) Ventral festoons showing tubercles at the posterointernal angles (

egs. Coxa I with 2 pointed spurs, the external slightly longer; coxaeI–IV with a small triangular spur. Trochanters without spur; tar-us I 0.457 ± 0.052 (0.361–0.511) long, 0.113 ± 0.007 (0.102–0.123)road; tarsus IV 0.337 ± 0.040 (0.263–0.377) long, 0.088 ± 0.0070.080–0.103) broad.

mblyomma longirostre (Koch, 1844) (Fig. 20)Cooley and Kohls (1944): description of the nymph as Ambly-

mma avecolens.Idiosoma. Length from apices of scapula to posterior body mar-

in 1.852 ± 0.151 (1.494–2.030), maximum breadth 1.426 ± 0.1141.197–1.589); outline oval, with 11 festoons without tubercles.cutum. Length 1.049 ± 0.076 (0.899–1.136), breadth 1.097 ± 0.056
0.963–1.166), breadth/length ratio 1.047 ± 0.030 (1.081–1.005),nornate, surface extensively shagreened (rugose); few puncta-ions, larger and deeper laterally. Eyes not orbited, at lateral scutalngles at the level of scutal midlength. Cervical grooves deep ante-iorly followed by a shallow depression. Spiracular plate triangular
). (B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV0 �m).

with rounded angles, with a discrete dorsal prolongation; length0.090 ± 0.010 (0.079–0.109), breadth 0.044 ± 0.006 (0.035–0.055).Gnathosoma (capitulum). Length from palpal apices to poste-rior margin 0.616 ± 0.046 (0.557–0.683), breadth 0.457 ± 0.017(0.428–0.476). Basis capituli triangular, posterior margin straight,without cornua; posterior margin convex ventrally, auriculae assmall posterolateral rounded projections, lateral margin projectedlaterally. Palpi length 0.482 ± 0.034 (0.437–0.532), article I withoutventral prolongation; article II 0.318 ± 0.019 (0.291–0.344) long,article III 0.134 ± 0.017 (0.106–0.161) long. Hypostome sharplypointed apically (lanceolate); length 0.444 ± 0.030 (0.403–0.490);length of toothed portion 0.298 ± 0.022 (0.266–0.332); dentition2/2 with 9–10 teeth per row. Legs. Coxa I with 2 short spurs, the
external broader and longer, the internal obsolete; coxae II–IVwith an obsolete ridge-like spur. Trochanters without spur; tar-sus I 0.539 ± 0.029 (0.490–0.562) long, 0.126 ± 0.009 (0.114–0.138)broad; tarsus IV 0.421 ± 0.036 (0.362–0.461) long, 0.094 ± 0.005(0.088–0.101) broad.


Fig. 17. Amblyomma cajennense nymph. (A) Dorsal gnathosoma (capitulum) (bar: 20 �m). (B) Ventral gnathosoma (bar: 20 �m). (C) Scutum (bar: 30 �m). (D) Coxae I–IV(bar: 30 �m).

Fig. 18. Amblyomma dissimile nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m). (B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV (bar:100 �m).


Fig. 19. Amblyomma dubitatum nymph. (A) Dorsal gnathosoma (capitulum) (bar: 20 �m). (B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV(bar: 30 �m).

Fig. 20. Amblyomma longirostre nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m). (B) Ventral gnathosoma (bar: 100 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV(bar: 100 �m).


F (bar:I

A

g(S(iaobr0G0Bwwc(Hltusa(Ib

A

ig. 21. Amblyomma oblongoguttatum nymph. (A) Dorsal gnathosoma (capitulum)–IV (bar: 30 �m).

mblyomma oblongoguttatum Koch, 1844 (Fig. 21)Nuttall (1912): description of the nymph as Amblyomma darlingi.Idiosoma. Length from apices of scapula to posterior body mar-

in 1.323 ± 0.059 (1.203–1.416), maximum breadth 1.153 ± 0.0421.075–1.219); outline oval, with 11 festoons without tubercles.cutum. Length 0.654 ± 0.025 (0.585–0.681), breadth 0.902 ± 0.0350.852–0.960), breadth/length ratio 1.378 ± 0.039 (1.339–1.455),nornate, large punctations evenly distributed, deeper later-lly. Eyes not orbited at lateral scutal angles at the levelf scutal midlength. Cervical grooves short and deep, endingefore the scutal midlength. Spiracular plate triangular withounded angles, with an evident dorsal prolongation; length.069 ± 0.005 (0.061–0.075), breadth 0.040 ± 0.005 (0.034–0.052).nathosoma (capitulum). Length from palpal apices to cornua apices.366 ± 0.022 (0.337–0.395); breadth 0.303 ± 0.015 (0.280–0.329).asis capituli sub-triangular, posterior margin slightly concaveith minute triangular cornua, posterior margin convex ventrally,ithout auriculae. Palpi length 0.284 ± 0.018 (0.256–0.306), arti-

le I with vestigial ventral prolongation; article II 0.182 ± 0.0140.160–0.199) long, article III 0.076 ± 0.005 (0.065–0.083) long.ypostome rounded apically; length 0.290 ± 0.019 (0.259–0.325);

ength of toothed portion 0.146 ± 0.009 (0.133–0.160); den-ition 2/2 with 7–8 teeth per row. Legs. Coxa I with 2nequal spurs, the internal shorter, the external longer andlender, slightly curved laterally; coxae II–IV with a small tri-ngular spur. Trochanters without spur; tarsus I 0.461 ± 0.0320.384–0.491) long, 0.105 ± 0.004 (0.100–0.115) broad; tarsusV 0.326 ± 0.033 (0.262–0.380) long, 0.082 ± 0.004 (0.077–0.094)
road.
mblyomma parvum Aragão, 1908 (Fig. 22)Guglielmone et al. (1990): description of the nymph.

30 �m). (B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 30 �m). (D) Coxae

Idiosoma. Length from apices of scapula to posterior body mar-gin 1.180 ± 0.031 (1.108–1.217), maximum breadth 0.824 ± 0.021(0.794–0.852), outline oval, longilinear, with 11 festoons with-out tubercles. Scutum. Length 0.617 ± 0.042 (0.523–0.670), breadth0.653 ± 0.025 (0.622–0.698), breadth/length ratio 1.060 ± 0.048(1.031–1.188); inornate; few medium punctations evenly dis-tributed. Eyes elongated, not orbited, slightly convex, at lateralscutal angles at the level of scutal midlength, delimitated medi-ally by a groove formed by small deep punctations. Cervicalgrooves long, deep anteriorly. Spiracular plate triangular withrounded angles, with a discrete dorsal prolongation; length0.048 ± 0.005 (0.043–0.059), breadth 0.027 ± 0.005 (0.021–0.039).Gnathosoma (capitulum). Length from palpal apices to cornua apices0.249 ± 0.023 (0.208–0.278), breadth 0.260 ± 0.011 (0.237–0.275).Basis capituli sub-rectangular, posterior margin slightly concavewith minute triangular cornua; posterior margin convex ventrally,without auriculae. Palpi length 0.180 ± 0.012 (0.157–0.196), arti-cle I with vestigial ventral prolongation; article II 0.118 ± 0.012(0.096–0.131) long, article III 0.049 ± 0.003 (0.045–0.054) long.Hypostome rounded apically; length 0.195 ± 0.004 (0.190–0.204);length of toothed portion 0.101 ± 0.005 (0.093–0.112); denti-tion 2/2 with 6–7 teeth per row. Legs. Coxa I with 2 pointedspurs, the external about twice as long the internal; coxae II–IVwith a small triangular spur. Trochanters without spur; tarsusI 0.315 ± 0.015 (0.298–0.343) long, 0.088 ± 0.006 (0.075–0.102)broad; tarsus IV 0.251 ± 0.022 (0.198–0.274) long, 0.070 ± 0.004(0.064–0.078) broad.

Amblyomma romitii Tonelli-Rondelli, 1939 (Fig. 23)Tonelli-Rondelli (1939): description of the nymph.Idiosoma. Length from apices of scapula to posterior

body margin 1.985 ± 0.134 (1.775–2.170), maximum breadth


F m). (B3

1d0bomslrs0Gr(sta(Hl3eTl(

A

g(S

ig. 22. Amblyomma parvum nymph. (A) Dorsal gnathosoma (capitulum) (bar: 10 �0 �m).

.637 ± 0.109 (1.455–1.782), outline oval. Dorsum of bodyensely pilose; 11 festoons without tubercles. Scutum. Length.799 ± 0.081 (0.676–0.907), breadth 1.187 ± 0.082 (1.025–1.295),readth/length ratio 1.489 ± 0.054 (1.411–1.581), inornate, numer-us punctations evenly distributed, larger laterally; posterolateralargin somewhat sinuous. Eyes large, not orbited at lateral

cutal angles at the level of scutal midlength. Cervical groovesong, almost reaching the scutal posterior margin, deeper ante-iorly. Spiracular plate triangular with rounded angles, marginslightly festooned, with a prominent dorsal prolongation; length.097 ± 0.002 (0.093–0.098), breadth 0.055 ± 0.002 (0.052–0.057).nathosoma (capitulum). Length from palpal apices to poste-ior margin 0.423 ± 0.061 (0.340–0.487), breadth 0.381 ± 0.0280.310–0.414). Basis capituli sub-triangular, posterior marginlightly concave, without cornua; posterior margin convex ven-rally, without auriculae. Palpi length 0.328 ± 0.037 (0.250–0.366),rticle I without ventral prolongation; article II 0.200 ± 0.0300.139–0.242) long, article III 0.104 ± 0.010 (0.089–0.120) long.ypostome rounded apically; length 0.314 ± 0.030 (0.272–0.355);

ength of toothed portion 0.191 ± 0.012 (0.176–0.212); dentition/3 with 7–8 teeth per row. Legs. Coxa I with 2 short spurs, thexternal slightly longer; coxae II–IV with a small triangular spur.rochanters without spur; tarsus I 0.586 ± 0.069 (0.429–0.660)ong, 0.120 ± 0.007 (0.108–0.129) broad; tarsus IV 0.357 ± 0.0530.285–0.455) long, 0.106 ± 0.007 (0.093–0.115) broad.

mblyomma rotundatum Koch, 1844 (Fig. 24)
Keirans and Oliver (1993): description of the nymph.Idiosoma. Length from apices of scapula to posterior body mar-
in 1.636 ± 0.078 (1.454–1.750), maximum breadth 1.322 ± 0.0541.190–1.402), outline oval, with 11 festoons without tubercles.cutum. Length 0.864 ± 0.031 (0.794–0.900), breadth 1.014 ± 0.036


(0.939–1.061), breadth/length ratio 1.174 ± 0.009 (1.162–1.189),inornate, moderate punctations, larger and deeper laterally; pos-terolateral margin somewhat sinuous. Eyes not orbited at lateralscutal angles at the level of scutal midlength. Cervical groovesending at the level of scutal midlength, deeper at the ante-rior half. Spiracular plate triangular with rounded angles, with adiscrete dorsal prolongation; length 0.091 ± 0.005 (0.079–0.098),breadth 0.061 ± 0.005 (0.052–0.070). Gnathosoma (capitulum).Length from palpal apices to posterior margin 0.425 ± 0.040(0.368–0.514), breadth 0.368 ± 0.007 (0.352–0.381). Basis capitulisub-triangular, posterior margin straight, without cornua; pos-terior margin straight ventrally, without auriculae. Palpi length0.300 ± 0.025 (0.265–0.357), article I without ventral prolon-gation; article II 0.201 ± 0.014 (0.183–0.232) long, article III0.078 ± 0.008 (0.064–0.095) long. Hypostome rounded apically;length 0.284 ± 0.013 (0.255–0.304); length of toothed portion0.147 ± 0.006 (0.133–0.155); dentition 2.5/2.5 (3/3 apically, 2/2 atthe base) with 7–8 teeth at the external and median rows, 4–5teeth at the internal row. Legs. Coxa I with 2 triangular spurs ofsimilar size; coxa II with short spurs, the external triangular andlonger, the internal rounded and almost obsolete; coxa III with anexternal triangular spur, and an internal spur obsolete or absent;coxa IV with a small triangular spur. Trochanters without spur; tar-sus I 0.445 ± 0.012 (0.421–0.462) long, 0.098 ± 0.006 (0.089–0.108)broad; tarsus IV 0.327 ± 0.030 (0.291–0.381) long, 0.069 ± 0.004(0.059–0.075) broad.

Amblyomma tigrinum Koch, 1844 (Fig. 25)Estrada-Pena et al. (1993): description of the nymph.Idiosoma. Length from apices of scapula to posterior body mar-

gin 1.265 ± 0.099 (1.113–1.448), maximum breadth 0.781 ± 0.052(0.701–0.884); outline oval, clearly longilinear, with 11 festoons


F m). (B1

wb0tatrdgdbL(tpdP

ig. 23. Amblyomma romitii nymph. (A) Dorsal gnathosoma (capitulum) (bar: 100 �00 �m). (E) Alloscutum (densely pilose) (bar: 100 �m).

ithout tubercles. Scutum. Length 0.652 ± 0.051 (0.575–0.765),readth 0.629 ± 0.042 (0.550–0.698), breadth/length ratio.964 ± 0.020 (0.912–0.982); inornate; rare punctations; pos-erolateral margin not sinuous. Eyes not orbited at lateral scutalngles at the level of the beginning of the posterior third ofhe scutum. Cervical grooves long, reaching the scutal poste-ior fourth, deeper at the anterior fourth; presence of a rugoseepression parallel and external to the posterior half of cervicalrooves. Spiracular plate triangular with rounded angles, with aiscrete dorsal prolongation; length 0.064 ± 0.005 (0.053–0.071),readth 0.030 ± 0.005 (0.025–0.039). Gnathosoma (capitulum).ength from palpal apices to posterior margin 0.332 ± 0.026
0.307–0.368), breadth 0.330 ± 0.020 (0.291–0.372). Basis capituliriangular, posterior margin slightly concave, without cornua;osterior margin convex ventrally, auriculae as posteriorlyirected spurs, not curved; lateral margin projected laterally.alpi length 0.251 ± 0.016 (0.228–0.279), article I without ventral

prolongation; article II 0.163 ± 0.011 (0.149–0.183) long, article III0.064 ± 0.006 (0.056–0.076) long. Hypostome rounded apically;length 0.230 ± 0.023 (0.201–0.269), length of toothed portion0.121 ± 0.011 (0.107–0.140); dentition 2/2 with 8–9 teeth per row.Legs. Coxa I with a long, narrow spur, its base fusing to coxa beforethe internal coxal margin; coxa II with a short triangular spur;coxae III–IV with a very short bridge-like spur. Trochanters with-out spur; tarsus I 0.316 ± 0.021 (0.283–0.362) long, 0.098 ± 0.006(0.088–0.111) broad; tarsus IV 0.236 ± 0.016 (0.213–0.265) long,0.070 ± 0.005 (0.061–0.077) broad.

Amblyomma triste Koch, 1844 (Fig. 26)
Estrada-Pena et al. (2002): description of the nymph.Idiosoma. Length from apices of scapula to posterior body mar-
gin 1.279 ± 0.049 (1.151–1.319), maximum breadth 0.795 ± 0.019(0.748–0.812); outline oval, clearly longilinear, with 11 festoonswithout tubercles. Scutum. Length 0.646 ± 0.022 (0.612–0.676),


Fig. 24. Amblyomma rotundatum nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m). (B) Ventral gnathosoma (bar: 30 �m). (C) Scutum (bar: 100 �m). (D) Coxae I–IV(bar: 30 �m).

Fig. 25. Amblyomma tigrinum nymph. (A) Dorsal gnathosoma (capitulum) (bar: 30 �m). (B) Ventral gnathosoma (bar: 20 �m). (C) Scutum (bar: 30 �m). (D) Coxae I–IV (bar:30 �m).


F3

ig. 26. Amblyomma triste nymph. (A) Dorsal gnathosoma (capitulum) (bar: 20 �m). (B)0 �m).

Fig. 27. Amblyomma parkeri nymph. (A) Dorsal gnathosoma (capitulum). (B) Vent


ral gnathosoma. (C) Scutum. (D) Coxa I. (E) Coxa II. (F) Coxa III. (G) Coxa IV.

ick-bo

b1tstproa(s0Bwlpa(Hl2swf(0b

A

Siptaorl0wscIkitslsb

rctbiinttc2TC

15. -Hypostomal dentition 3/3 apically 16

T.F. Martins et al. / Ticks and T

readth 0.648 ± 0.018 (0.616–0.680), breadth/length ratio.002 ± 0.010 (0.989–1.021); inornate; rare punctations; pos-erolateral margin somewhat sinuous. Eyes not orbited at lateralcutal angles at the level of the beginning of the posteriorhird of the scutum. Cervical grooves long, reaching the scutalosterior fourth, deeper at the anterior fourth; presence of augose depression parallel and external to the posterior halff cervical grooves. Spiracular plate triangular with roundedngles, with a discrete dorsal prolongation; length 0.064 ± 0.0060.052–0.077), breadth 0.028 ± 0.005 (0.021–0.037). Gnatho-oma (capitulum). Length from palpal apices to posterior margin.346 ± 0.035 (0.290–0.391), breadth 0.364 ± 0.010 (0.348–0.381).asis capituli triangular, posterior margin slightly concave,ithout cornua; posterior margin convex ventrally, auricu-

ae as posteriorly directed spurs, not curved; lateral marginrojected laterally. Palpi length 0.266 ± 0.018 (0.236–0.297),rticle I without ventral prolongation; article II 0.187 ± 0.0190.146–0.223) long, article III 0.058 ± 0.005 (0.050–0.068) long.ypostome rounded apically; length 0.265 ± 0.012 (0.245–0.288),

ength of toothed portion 0.137 ± 0.006 (0.128–0.151); dentition/2 with 8–9 teeth per row. Legs. Coxa I with a long, robustpur, its base reaching the internal coxal margin; coxae II–IVith a short bridge-like spur that gradually decreases in size

rom II to IV. Trochanters without spur; tarsus I 0.329 ± 0.0130.306–0.350) long, 0.096 ± 0.005 (0.090–0.106) broad; tarsus IV.242 ± 0.018 (0.214–0.265) long, 0.071 ± 0.003 (0.066–0.077)road.

mblyomma parkeri Fonseca e Aragão, 1952 (Fig. 27)Fonseca and Aragão (1952): description of the nymph.Idiosoma. Outline oval, with 11 festoons without tubercles.

cutum. Length 0.980, breadth 1.207, breadth/length ratio 1.231;nornate, surface slightly shagreened, numerous large and deepunctations evenly distributed. Eyes not orbited at lateral scu-al angles at the level of scutal midlength. Cervical grooves deept the anterior third, followed by shallow depressions forwardf the level of scutal midlength. Spiracular plate triangular withounded angles; length 0.086, breadth 0.041. Gnathosoma (capitu-um). Length from palpal apices to posterior margin 0.521, breadth.466. Basis capituli hexagonal, posterior margin slightly concave,ithout cornua; posterior margin convex ventrally, auriculae as

mall posterolateral rounded projections. Palpi length 0.402, arti-le I without ventral prolongation; article II 0.260 long, articleII 0.122 long. Hypostome is not redescribed because it was bro-en in the single specimen available for redescription. However,n its original description, Fonseca and Aragão (1952) reported ito be rounded apically, with a 2/2 dentition. Legs. Coxa I with 2hort spurs, the external longer and broader, the internal triangu-ar; coxae II–IV with a small triangular spur. Trochanters withoutpur; tarsus I 0.532 long, 0.132 broad; tarsus IV 0.430 long, 0.137road.

The single engorged nymphal specimen used for the presentedescription [previously described by Fonseca and Aragão (1952)],oupled with the identification key provided below, allowed uso identify a second A. parkeri engorged nymph collected from arown howler monkey (Alouatta fusca) in São Paulo municipal-

ty, state of São Paulo, in January 2010. The pictures providedn Fig. 27 refer to this new engorged nymph, since the firstymphal specimen was very dark, precluding good pictures. Likehe first nymph, this second nymph also had a broken hypos-
ome. Species identification of this second nymph was definitelyonfirmed by leaving the engorged nymph alive in a cabinet at5 ◦C and 85% RH, where the tick molted to an A. parkeri female.his female has been deposited in the CNC (accession numberNC-1600).
rne Diseases 1 (2010) 75–99 97

Key to nymphs of the genus Amblyomma of Brazil

1. -Coxa I with 1 spur, auriculae as posteriorly directedspurs, not curved

2

-Coxa I with 2 spurs 3

2. -Spur on coxa I robust, its base reaching the internalcoxal margin; scutum with a somewhat sinuousposterolateral margin

A. triste (Fig. 26)

-Spur on coxa I narrow, its base fusing to coxa beforethe internal coxal margin; scutum lacking a sinuousposterolateral margin

A. tigrinum(Fig. 25)

3. -Coxa II with 2 spurs, the internal very small 4-Coxa II with only one spur 5

4. -Scutum with deep punctations evenly distributed;hypostomal dentition 2/2

A. humerale (Fig. 6)

-Scutum with deep punctations concentrated in thelateral fields; hypostomal dentition 2.5/2.5

A. rotundatum(Fig. 24)

5. -Cornua present 6-Cornua absent 12

6. -Tubercles present at posterointernal angles offestoons

7

-Tubercles absent at posterointernal angles of festoons 9

7. -Scutum length >0.75 mm 8-Scutum length <0.75 mm A. brasiliense

(Fig. 16)

8. -Festoons with well-developed, whitish tubercles atthe posterointernal angles; scutum broader,breadth/length ratio >1.3, with a somewhat sinuousposterolateral margin

A. incisum (Fig. 7)

-Festoons with poorly developed, brownish tuberclesat the posterointernal angles; scutum slightly longer,breadth/length ratio <1.3, lacking a sinuousposterolateral margin

A. scalpturatum(Fig. 14)

9. -Coxa IV with a small triangular spur as long asbroad; internal spur on coxa I shorter than the spuron coxa II

10

-Coxa IV with a small triangular spur longer thanbroad; internal spur on coxa I as long as the spur oncoxa II

11

10. -Dorsal basis capituli hexagonal; pronouncedtriangular cornua

A. naponense(Fig. 9)

-Dorsal basis capituli sub-rectangular; minutetriangular cornua; idiosoma longilinear

A. parvum (Fig. 22)

11. -Distinct triangular cornua; scutal length >0.8 mm A. latepunctatum(Fig. 8)

-Minute triangular cornua; scutal length <0.7 mm A.oblongoguttatum(Fig. 21)

12. -Auriculae present as small posterolateral roundedprojections

13

-Auriculae absent 15

13. -Coxa I with 2 short spurs; eyes located at the levelof the scutal midlength

14

-Coxa I with 2 medium spurs; eyes located at the levelof the posterior third of the scutum; idiosomalongilinear; hypostome rounded apically

A. ovale (Fig. 11)

14. -Scutal surface extensively shagreened (rugose)(this character is visualized under opticalmicroscopy better than under scanning electronmicroscopy) with few large and deep punctations inthe lateral fields; hypostome sharply pointed

A. longirostre(Fig. 20)

-Scutal surface slightly shagreened, with many largeand deep punctations evenly distributed; hypostomerounded apically (The single specimen of A. parkerithat was available for examination had a brokenhypostome. However, according to Fonseca andAragão (1952), the A. parkeri nymph has ahypostome rounded apically, with a dentition 2/2.)

A. parkeri (Fig. 27)

-Hypostomal dentition 2/2 throughout 17

16. -Dorsum of body lacking dense pilosity; scutumsparsely punctated; spiracular plate not festooned;hypostomal dentition 2.5/2.5

A. dissimile(Fig. 18)

9 ick-bo

D

aogccAlAssb

8 T.F. Martins et al. / Ticks and T

-Dorsum of body densely pilose; scutum denselypunctated; spiracular plate slightly festooned;hypostomal dentition 3/3

A. romitii (Fig. 23)

17. -Coxa I with 2 unequal spurs, the external longerthan the internal

18

-Coxa I with 2 sub-equal short spurs; dorsal basiscapituli pentagonal; palpal article IV projectedapically

A. fuscum (Fig. 5)

18. -Deep punctations rarely present on scutum orwhen present they are concentrated in the lateralfields

19

-Scutum moderately punctate, deep punctationsevenly distributed

23

19. -Dorsal basis capituli triangular or sub-triangular;scutal surface extensively shagreened (rugose) (Thischaracter is visualized under optical microscopybetter than under scanning electron microscopy.)

20

-Dorsal basis capituli broadly rectangular; scutalsurface slightly shagreened

22

20. -Coxa I with well separated spurs, the externallonger and pointed

21

-Coxa I with stout spurs, close to each other; idiosomalongilinear

A. aureolatum(Fig. 1)

21. -Scutum broader, breadth/length ratio >1.3, deeppunctations rare

A. nodosum(Fig. 10)

-Scutum slightly longer, breadth/length ratio <1.3,deep punctations concentrated in the lateral fields

A. calcaratum(Fig. 3)

22. -Scutum broader, breadth/length ratio >1.3,cervical groove extending to the scutal posteriorborder

A. pseudoconcolor(Fig. 13)

-Scutum slightly longer, breadth/length ratio <1.3,cervical groove extending to the level of scutalmidlength

A. auricularium(Fig. 2)

23. -Coxa I with external spur twice as long as internalspur

24

-Coxa I with external spur longer than internal,however less than twice as long

25

24. -Dorsal basis capituli slightly hexagonal; theexternal spur on coxa I less than twice as long ascoxa II spur

A. varium (Fig. 15)

-Dorsal basis capituli rectangular; coxa I with externalspur at least twice as long as coxa II spur

A. cajennense(Fig. 17)

25. -Cervical groove long, well forward of the level ofthe posterior margin of the eyes; scutum with largeand deep punctations in the lateral fields, smallerpunctations in the central field

26

-Cervical groove short, ending as a small shallowdepression at the level of the posterior margin of theeyes; scutum with large and deep punctations inboth lateral and central fields

A. coelebs (Fig. 4)

26. -Cervical groove deep in its anterior convergenthalf, and as a shallow large depression on itsposterior divergent half; scutum broader,breadth/length ratio >1.33

A. pacae (Fig. 12)

-Cervical groove deep throughout; lacking a shallowand large depression in its posterior divergent half;scutum slightly longer, breadth/length ratio <1.33

A. dubitatum(Fig. 19)

iscussion

Members of the genus Amblyomma have considerable medicalnd veterinary importance in South America. Different membersf this genus are the main vectors and reservoirs of spotted feverroup rickettsioses to humans and animals in different parts of theontinent (Labruna, 2009). Amblyomma spp. (especially nymphs)omprise the vast majority of human-biting tick reports in Southmerica (Guglielmone et al., 2006; Szabó et al., 2006). However,

ife-history information for most Amblyomma species from South
merica is still incomplete and has been affected by the lack ofufficient host records for the immature stages. With the presenttudy, we hope that morphology-based species identification cane applied to field-collected Amblyomma nymphs in Brazil, con-
rne Diseases 1 (2010) 75–99

tributing to a better knowledge of the life histories of Amblyommaticks and the disease agents they transmit in the country.

Among the 29 Amblyomma species established in Brazil, onlyA. goeldii Neumann, 1899, and A. geayi Neumann, 1899 were notincluded in the present study. The nymphal stage of these 2 rarespecies remains undescribed (Onofrio et al., 2006). Since both A.goeldii and A. geayi in Brazil appear to be restricted to the northernregion (Amazonia) (Aragão, 1936; Labruna et al., 2009), our iden-tification key with 27 species comprises all Amblyomma speciesfrom the remaining regions of the country. Hopefully, nymphalspecimens of A. goeldii and A. geayi will be available for morpholog-ical description in the near future, so they can be included in theidentification key.

Finally, it must be stressed that our study evaluated nymphsderived from a single geographic population of each tick species.Some of these species, like A. cajennense and A. parvum, have awide distribution in the New World, with significant intraspecificgenetic differences reported among different populations (Beati etal., 2007; Nava et al., 2008). At least one of these species, A. cajen-nense, is now considered to be a complex of several distinct speciesyet to be determined (Beati et al., 2007). Thus, it is quite possi-ble that some of our descriptions/redescriptions based on Brazilianspecimens will not fit perfectly with presumably conspecific pop-ulations from other geographic areas of the New World. Furtherstudies based on both morphological and genetic methods shouldtest this possibility.

Acknowledgments

We are indebted to the following colleagues who provided uswith unfed nymphs from their laboratory colonies for the presentstudy: J.L.H. Faccini and E.M. Piranda from Federal Rural Univer-sity of Rio de Janeiro (Seropédica, Rio de Janeiro, Brazil) for theA. auricularium nymphs; M.P.J. Szabó from Federal University ofUberlândia (Uberlândia, Minas Gerais, Brazil) for the A. incisumnymphs; and A.A. Guglielmone and S. Nava from Instituto Nacionalde Tecnologia Agropecuaria (Rafaela, Santa Fe, Argentina) for the A.pseudoconcolor nymphs. We are extremely grateful to A.A. Gugliel-mone for revising earlier versions of this study, and providingvaluable suggestions to improve the identification key. Thanks toGustavo Fernochi for his technical support in editing photographs.This study was supported by Fundacão de Amparo à Pesquisa doEstado de São Paulo (FAPESP) and Conselho Nacional de Desen-volvimento Científico e Tecnológico (CNPq).

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Nymphs of the genus Amblyomma (Acari: Ixodidae) of Brazil: descriptions, redescriptions, and identification key

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