Zootaxa 3579: 80–88 (2012)
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Copyright © 2012 · Magnolia Press
ISSN 1175-5326 (print edition)
Article
ZOOTAXA
ISSN 1175-5334 (online edition)
urn:lsid:zoobank.org:pub:A0710F65-8CF1-4853-ACB7-EE5575E3A3BF
Redescription of the larva of Amblyomma oblongoguttatum Koch, 1844
(Acari: Ixodidae) by light and scanning electron microscopy
FABIO S. BARBIERI1, LUCIANA G. BRITO1, DARCI M. BARROS-BATTESTI2, KÁTIA M. FAMADAS3,
MARCELO B. LABRUNA4 & LUIS MARCELO A. CAMARGO5
1
Embrapa Rondônia, 78900-970, Porto Velho, RO, Brazil. E-mail: [email protected]
Laboratório Especial de Coleções Zoológicas, Instituto Butantan, 05503-900, São Paulo, SP, Brazil
3
Departamento de Parasitologia Animal, UFRRJ, 23890-000, Seropédica, RJ, Brazil
4
Faculdade de Medicina Veterinária e Zootecnia, USP, 05508-900 São Paulo, SP, Brazil
5
Instituto de Ciências Biomédicas 5, USP, 78965-000, Monte Negro, RO, Brazil
2
Abstract
The larval stage of Amblyomma oblongoguttatum Koch is redescribed using optical and scanning electron microscopy.
Unfed larvae were obtained from a colony of A. oblongoguttatum originated from engorged females collected on domestic
pigs from Monte Negro municipally (10°29’S, 63°32’W), State of Rondônia, Western Amazon, Brazil. Several characters
are presented including the chaetotaxy of the idiosoma, palpi and Haller’s organ, as well as morphological features of the
idiosoma, gnathosoma and legs. In addition, the porotaxy (topographical and numerical patterns of integumentary structures) were presented by using a new nomenclature recently proposed. The chaetotaxy of the larvae of A. oblongoguttatum, in general, is similar to other Neotropical Amblyomma species. Three types of integumentary structures were observed
on the idiosoma: lyrifissures, small glands, and large wax glands. Topographic and numerical patterns of the integumentary structures consisted of 5 pairs of large wax glands (1 dorsal/ 4 ventral), 24 pairs of lyrifissures (11 dorsal/ 13 ventral),
and 49 pairs of small glands (28 dorsal/ 21 ventral). These topographic and numerical patterns found for A. oblongoguttatum show only minor differences when compared with patterns of other Amblyomma larvae, however, a few key features
can be used for identification of these species.
Key words: Amblyomma oblongoguttatum, larva, description, morphology, chaetotaxy, porotaxy
Introduction
A total of 59 Amblyomma species are recorded from the Neotropical region, of which 30 are considered established
in Brazil (Dantas-Torres et al. 2009). Morphological descriptions for these Neotropical Amblyomma species have
been published for the adult stage but are seldom available for immature stages, with the exception of an illustrated
key for nymphs from Brazil (Martins et al. 2010). This oversight occurs because most species are primarily
associated with native wildlife, making it difficult to obtain larvae and nymphs from females correctly identified
through field collections.
Morphological description of immature tick stages is fundamental to studies of tick ecology and disease
relationships, since the correct identification of immature stages, and the specific host species, are important points
in the epidemiology of tick-borne disease. In tick morphology, including the genus Amblyomma, some
integumentary structures such as setae and pores have been used in systematic studies of larvae (Dinnik & Zumpt
1949; Nawar & Madbouly 1985; Klompen et al. 1996; Barbieri et al. 2007). Sensory setae are the most studied
integumentary structure, however their topographical distribution is highly conserved among Neotropical
Amblyomma species; therefore these setae are of poor systematic value for specific level comparisons (Barbieri et
al. 2007).
Some authors have utilized the integumentary pores distributed on the idiosoma of the larvae for specific
identification of ticks. On the larval idiosoma there are three types of the pores: lyrifissures, small glands, and large
80
Accepted by O. Seeman: 5 Nov. 2012; published: 11 Dec. 2012
wax glands. These structures form topographical and numerical patterns that are intraspecifically stable and also
less variable between species of the same genus; therefore they can be used to identify the larval stage to species
(Barbieri et al. 2007). Recently, Barbieri et al. (2007) proposed a new nomenclature for integumentary pores based
on the larval porotaxy of four Neotropical Amblyomma species, and described the larvae of Amblyomma ovale
Koch (Barbieri et al. 2008a) and Amblyomma pacae Aragão (Barbieri et al. 2008b) based on this new
methodology, which was an adaptation of a methodology previously proposed by Klompen et al. (1996).
The tick Amblyomma oblongoguttatum Koch is distributed from Mexico to northern Brazil and is also found in
central-western, southeastern and southern Brazil (Arzua et al. 2005). This species is found parasitizing tapirs,
jaguars, domestic dogs, wild and domestic pigs, and humans (Labruna et al. 2005; Guglielmone et al. 2006). The
larval stage of A. oblongoguttatum was described by Nutall (1912) under the name of Amblyomma darlingi,
considered later as a synonym; therefore, the present study redescribes the morphology of the larva of A.
oblongoguttatum by means of conventional light and scanning electron microscopy, emphasizing chaetotaxy and
porotaxy.
Material and methods
Ticks used in this study were from the progenies of three engorged females of A. oblongoguttatum collected on
domestic pigs from Monte Negro municipally (10°29’S, 63°32’W), State of Rondônia, Western Amazon, Brazil.
The females were determined to species using the keys and redescriptions of Robinson (1926), Aragão and Fonseca
(1961), and Onofrio et al. (2006).
The females oviposited under laboratory conditions (27±1°C, 80 ±10% RH) in BOD, and the egg masses were
individually placed in modified disposable syringes. After all the larvae had hatched, a sample of 30 individuals
was kept without meal for 15 days, so that the consolidation of the exoskeleton took place. The specimens were
subsequently placed in water at 70 ± 10°C and preserved 70% ethanol. From these, 15 were prepared for light
microscopy, as previously described by Barbieri et al. (2007), and another five specimens were processed for
scanning electron microscopy according to the method proposed by Keirans et al. (1976).
Larval chaetotaxy terminology is that of Clifford and Anastos (1960), Hess and Vlimant (1983) and Wooley
(1983). For porotaxy we have used the nomenclature proposed by Barbieri et al. (2007). For determination of the
frequency of integumentary structures, each idiosomal side (left and right) was analyzed independently, according
to Klompen et al. (1996).
Larval illustrations and measurements were made using a Leica DM 1000 light microscope equipped with a
drawing tube and both ocular and slide micrometers. All measurements are given in millimeters. The average is
followed by the standard deviation and the interval represents a sample of 15 specimens.
For scanning electron microscopy four larvae were dehydrated for 30 minutes in each of the following
concentrations of alcohol: 70%, 80%, 90% and 100% (three times). They were then kept in acetone until
undergoing critical point drying. Micrographs were taken using a Digital Scanning Microscope ZEISS/LEO 440
from the Laboratório de Microscopia Eletrônica do Museu de Zoologia da Universidade de São Paulo. Two larval
specimens were deposited in the Acari Collection of the Instituto Butantan (IBSP 9689), São Paulo, Brazil.
Redescription
Amblyomma oblongoguttatum Koch 1844, LARVA
(Fig. 1–10)
IDIOSOMA: Dorsal surface (fig. 4). Length from apices of scapulae to posterior margin of body 0.631 ± 0.020
(0.592–0.662); greatest width 0.618 ± 0.017 (0.588–0.649); outline oval, with 11 festoons. Setae: 2 central dorsal
pairs (Cd1, Cd2), 8 marginal dorsal pairs (Md1–Md8), with Md1 and Md2 before a large wax gland, present in the
segment VIII (WdVIII1), and Md3 pair located on the inner side behind of these glands; Md4 – Md8 pairs posterior
to large wax gland, each one on a different festoon. Scutum (Fig. 4, 8): outline subtriangular; length 0.247 ± 0.010
(0.230–0.262) along median line; width 0.400 ± 0.016 (0.377–0.441) at the level of eyes; with few punctuations
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(Fig. 8). Eyes slightly bulging and shallow; cervical grooves distinct extending parallel to the proximities of setae
Sc3, but slightly diverging at the end. Setae: 3 scutal pairs—Sc1, Sc2, Sc3 (fig. 4, 8).
Ventral surface (figs. 4, 9): Anal aperture on central portion of opisthosoma. Setae: 3 sternal pairs (St1, St2, St3),
2 preanal pairs (Pa1, Pa2), 4 premarginal pairs (Pm1–Pm4), 5 marginal ventral pairs (Mv1–Mv5), and 1 pair anal A1.
GNATHOSOMA: Dorsal (figs. 1, 5). Basis capituli triangular in outline; length from posterior margin of
trochanter to posterior margin of capituli 0.095 ± 0.005 (0.083–0.103), width 0.178 ± 0.003 (0.171–0.183).
Posterior margin straight, cornua absent. Basis capituli on median line with 1 sensillum pair. Palpal grooves
segments well-defined (fig. 5, 6). Palpi length from apices of tibiotarsal segment to posterior margin of trochanter
0.140 ± 0.005 (0.133–0.147), width 0.049 ± 0.002 (0.045–0.054); trochanter length 0.021 ± 0.007 (0.016–0.044),
width 0.039 ± 0.002 (0.037–0.043); femur length 0.059 ± 0.003 (0.054–0.064), width 0.047 ± 0.003 (0.041–0.051);
genu length 0.062 ± 0.003 (0.056–0.069), width 0.041 ± 0.002 (0.035–0.044). Femur with sensillum near seta Fa1
(Fig. 1).
FIGURE 1. Amblyomma oblongoguttatum larva. Gnathosoma dorsal (right) and ventral (left) views. Abbreviations: d—dorsal;
v—ventral; a—antiaxial; p—paraxial; t—terminal; F—femur; G—genu; Tt—tibiotarsus.
Ventral (Figs. 1, 6): Basis capituli as illustrated. Hypostome – compact, spatulate, length from apices to post
hypostomal seta 0.117 ± 0.004 (0.111–0.124), dental formula 2/2 in file teeth, apical corona usually with 9
denticles (fig. 7); 1 pair of hypostomal setae (Ph1).
Palpal setae (Figs. 1, 5, 6): 10 setae on tibiotarsus, 6 terminal (Ttt1 – Ttt6), 2 paraxial (Ttp1, Ttp2) and 2 antiaxial
(Tta1, Tta2) (fig. 7); 6 genual setae, 1 paraxial (Gp1), 1 antiaxial (Ga1), 3 dorsal (Gd1, Gd2, Gd3), and 1 ventral (Gv1);
6 femoral setae, 1 paraxial (Fp1), 2 antiaxial (Fa1, Fa2), 1 dorsal (Fd1), and 2 ventral (Fv1, Fv2); trochanter 0.
LEGS: Coxa I with 2 triangular short spurs, the external slightly larger; coxa II and III each with 1 short
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BARBIERI ET AL.
triangular spur (Fig. 9). Setae: 3 on coxa I, 1 anterior (CIa), 1 posterior (CIp) and 1 paraxial (CIpa); coxa II and III,
each with 2 setae, 1 anterior (CIIa, CIIIa) and 1 posterior (CIIp, CIIIp). Trochanter lacking spur. Tarsus I length
0.200 ± 0.007 (0.184–0.207), width 0.075 ± 0.004 (0.067 – 0.082). Setae (figs. 2, 3, 10): Dorsal, 2 in dorsal I group
(dI1, dI2); 5 dorsal II (dII1 – dII5); 2 dorsal III (dIII1, dIII2); 2 dorsal IV (dIV1, dIV2); 0 dorsal V; 2 dorsal VI (dVI1,
dVI2); Ventral, 2 ventral I (vI1, vI2), 2 in II group (vII1, vII2) and 2 in III (vIII1, vIII2); Lateral anterior, 1 in lateral
anterior I group (laI1), and 3 in laII group (laII1, laII2, laII3); Lateral posterior, 1 in lateral posterior I group (lpI1) and
3 in lpII group (lpII1, lpII2, lpII3). Ambulacrum as illustrated (Fig. 2, 3, 10).
FIGURE 2. Amblyomma oblongoguttatum larva. Tarsus I dorsal views. Abbreviations: d—dorsal; a— antiaxial; p—paraxial;
la—lateral anterior; lp—lateral posterior.
FIGURE 3. Amblyomma oblongoguttatum larva. Tarsus I ventral views. Abbreviations: v—ventral; a— antiaxial; p—paraxial;
la—lateral anterior; lp—lateral posterior.
Porotaxy: Dorsal (Fig. 4): Large wax gland—1 pair located on the lateral margin of the idiosoma, on segment
VIII, WdVIII1 (30/30 = 30 idiosomal sides containing the pore/ 30 observed idiosomal sides). Lyrifissures – 11
pairs: 1 pair on segment III, LdIII5 (30/30); 1 in IV, LdIV2 (29/30); 2 in V, LdV1 (23/30) and LdV6 (30/30); 1 in VI,
LdVI1 (26/30); 2 in VIII, LdVIII4 (25/30) and LdVIII6 (25/30); 2 in IX, LdIX4 (29/30) and LdIX6 (30/30); 2 in XII,
LdXII4 (27/30) and LdXII7 (30/30). Small glands – 28 pairs, 3 in scutum, 2 in segment III, SdIII3 (21/30) and SdIII5
(30/30); 1 in IV, SdIV6 (29/30); and one additional asymmetric small gland on the posterior central margin of the
scutum, SdV7 (15/15). In alloscutum, 25 pairs on segments: 1 in IV, SdIV2 (27/30); 3 in V, SdV2 (20/30), SdV4 (29/
30), SdV6 (26/30); 4 in VI, SdVI1 (23/30), SdVI2 (10/30), SdVI4 (29/30), SdVI6 (27/30); 3 in VIII, SdVIII4 (29/30),
SdVIII6 (29/30), SdVIII7 (27/30); 3 in IX, SdIX1 (21/30), SdIX2 (29/30), SdIX7 (30/30); 3 in X, SdX1 (13/30), SdX3
(29/30), SdX5 (11/30), and dorsal fovea (dorsal series 7–30/30); 2 in XI, SdXI3 (28/30), SdXI7 (30/30); 2 in XII,
SdXII1 (29/30), SdXII7 (29/30); 3 in XIII, SdXIII1 (30/30), SdXIII3 (28/30), SdXIII5 (27/30); 1 in XIV, SdXIV1 (27/
30).
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FIGURE 4. Amblyomma oblongoguttatum larva. Segmentation model of the idiosoma. Segments are indicated by Roman
numbers (III–VI e VIII–XIV) and delimitated by dashed lines (----); series are indicated by Arabic numbers and delimitated by
dotted lines (........). Integumentary structures are illustrated.
Ventral (Fig. 4): Large wax glands—4 pairs, one behind each coxa in segments III, IV and V, WvIII1 (30/30),
WvIV4 (30/30), WvV5 (30/30); 1 pair located on the 5th. festoon, segment XIII, WvXIII1 (30/30). Lyrifissures—13
pairs on segments: 1 in IV, LvIV6 (28/30); 1 in V, LvV6 (30/30); 1 in VIII, LvVIII5 (26/30); 4 in IX, LvIX1 (28/30),
LvIX3 (30/30), LvIX5 (28/30) and LvIX6 (28/30); 1 in X, LvX2 (30/30); 1 in XI, LvXI2 (30/30); 1 in XII, LvXII3
(30/30); 2 in XIII, LvXIII2 (30/30) and LvXIII6 (30/30); 1 in XIV, LvXIV2 (29/30).
Small gland. 21 pairs on segments: 1 in III, SvIII6 (30/30); 1 in IV, SvIV6, (30/30); 2 in V, SvV1 (30/30), SvV5
(29/30); 1 in VI, SvVI2 (30/30); 1 in VIII, SvVIII1 (30/30); 3 in IX, SvIX2 (29/30), SvIX4 (30/30), SvIX4’ (29/30); 3
in X, SvX1 (30/30), SvX3 (29/30), SvX4 (23/30); 3 in XI, SvXI1 (30/30), SvXI3 (29/30), SvXI4 (16/30); 2 in XII,
SvXII2 (30/30), SvXII4 (30/30); 2 in XIII, SvXIII3 (30/30), SvXIII6 (30/30); 2 in XIV, SvXIV1 (29/30), SvXIV4 (13/
30).
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Asymmetric lyrifissures found at low frequency, LvXIII6 (01/30), and small glands, SdX6 (01/30), SdXII3 (01/
30), SdXII4 (01/30), SdXIII6 (02/30), and SvVIII4 (01/30).
FIGURE 5–10. Amblyomma oblongoguttatum larva. 5. Gnathosoma, dorsal view (30 µm); 6 Gnathosoma, ventral view (30
µm); 7. Detail of tibiotarsus and hypostome (30 µm); 8. Detail of scutum (30 µm); 9. Coxae I–III (20 µm); 10.Tarsus I, dorsal
view (30 µm).
Discussion
The chaetotaxy of the idiosoma, palpi and tarsus I of the A. oblongoguttatum larva was generally similar to other
Amblyomma species, except for the tibiotarsal segment and group dII of the tarsus I. The tibiotarsal segment of the
palpi presented 10 setae (Figs.7), as also observed in the larvae of Amblyomma varium Koch, Amblyomma
dubitatum Neumann, Amblyomma parvum Aragão and Amblyomma pseudoparvum Guglielmone, Mangold and
Keirans (Amorim & Serra Freire 1996; Amorim & Serra Freire 1999a; Guglielmone et al. 1990). The larvae of
Amblyomma rotundatum Koch (Amorim & Serra Freire 1995) showed seven setae on this segment, larvae of A.
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ovale and A. pacae 11 setae (Barbieri et al. 2008a; Barbieri et al. 2008b), and the larvae of Amblyomma cajennense
(Fabricius), Amblyomma longirostre (Koch) and Amblyomma brasiliense Aragão had 12 setae (Famadas et al.
1997; Barros-Battesti et al. 2005; Sanches et al. 2009).
Tarsus I of the larvae of A. oblongoguttatum is in accordance with the genus Amblyomma as stated by Clifford
and Anastos (1960), with the tarsal formula 2:2:2:2:2 of the dorsal setae. However, we found five setae in group dII
of the larvae of A. oblongoguttatum (Fig. 2, 10), as in A. rotundatum (Amorim & Serra Freire 1995), A. varium
(Amorim & Serra Freire 1996), A. ovale (Barbieri et al. 2008a), and A. pacae (Barbieri et al. 2008b). In contrast,
six setae were found in group dII of the tarsus of A. parvum and A. pseudoparvum (Guglielmone et al. 1990) and A.
dubitatum (Amorim & Serra Freire 1999a), whereas seven setae were reported for A. cajennense, A. longirostre,
and A. brasiliense (Famadas et al. 1997; Barros-Battesti et al. 2005; Sanches et al. 2009).
The idiosoma of the A. oblongoguttatum larva contained lyrifissures, small glands, and large wax glands.
These structures presented typical morphological features as described in the literature (Schulze 1942; Dinnik &
Zumpt 1949; Barbieri et al. 2007), and were found isolated or associated over the entire idiosoma, except on the
scutum where large wax glands were not observed.
Large wax glands are the less numerous integumentary structures, but they are the most stable, with minimal
frequency variation between specimens when compared to lyrifissures and small glands. This stability in the
frequency of large wax glands was also observed in the larva of A. cajennense, A. longirostre, A. parvum, A.
rotundatum, A. ovale and A. pacae (Barbieri et al. 2007; Barbieri et al. 2008a; Barbieri et al. 2008b). In A.
oblongoguttatum, we found four pairs on the ventral surface, three located behind each coxa (segments III, IV and
V), and one on the 5th festoon (segment XIII), and a single dorsal pair located on the posterior lateral margin of the
body (segment VIII) (fig. 4).
The large wax arrangement of A. oblongoguttatum is in accordance with Clifford and Anastos (1960) for the
genus Amblyomma, and confirmed for A. cajennense (Famadas et al. 1997; Barbieri et al. 2007), Amblyomma
glauerti Keirans, King and Sharrad, Amblyomma variegatum (Fabricius), Amblyomma americanum (Linnaeus)
(Klompen et al. 1996), A. parvum, A. rotundatum (Barbieri et al. 2007), A. pacae (Barbieri et al. 2008b), and
Amblyomma brasiliense (Sanches et al. 2009). However, the larvae of Amblyomma tuberculatum Marx,
Amblyomma geomydae (Cantor), Amblyomma babirussae Schulze (Klompen et al. 1996), Amblyomma aureolatum
(Pallas) (Arzua, 2002), A. longirostre (Barros-Battesti et al. 2005; Barbieri et al. 2007) and A. ovale (Barbieri et al.
2008a) possess one additional ventral pair on the 4th festoon (segment XII); moreover, the larva of A. geomydae has
an additional pair on segment V; A. barbirussae has two additional pairs on segment V and one pair on X, and A.
longirostre has an additional dorsal pair (WdV1) on segment V.
The larval idiosoma of A. oblongoguttatum possesses 11 and 13 pairs of lyrifissures in the dorsal and ventral
surfaces, respectively (fig. 4). The numerical and topographic pattern of lyrifissures in A. oblongoguttatum is
similar to those observed in the larvae of A. cajennense, A. longirostre, A. rotundatum, A. ovale and A. pacae
(Barbieri et al. 2007; Barbieri et al. 2008a; Barbieri et al. 2008b). In the A. parvum larva there are 12 pairs of
lyrifissures on the ventral surface and the lyrifissure of the segment XIV is absent.
Small glands are the most numerous structures on the idiosomal segments of the A. oblongoguttatum larva,
including on the dorsal scutum, however they are less stable. According to Barbieri et al. (2007) this variation
appears to be a function of the greater number of small glands present on the idiosoma, since the least variation was
observed in the large wax glands, which have the lowest number on the idiosoma. This pattern of variation has
previously been reported for larvae of other species within the genera Rhipicephalus Koch (Nawar & Madbouly
1985), Amblyomma, Dermacentor Koch and Ixodes Latreille (Klompen et al. 1996), as well as in A. cajennense, A.
longirostre, A. parvum, A. rotundatum (Barbieri et al. 2007), A. ovale (Barbieri et al. 2008a) and A. pacae (Barbieri
et al. 2008b).
The patterns of lyrifissures, small glands, and large wax glands observed in A. oblongoguttatum are similar to
patterns found in other Neotropical Amblyomma species (Barbieri et al. 2007; Barbieri et al. 2008a; Barbieri et al.
2008b). However, when a particular group of these structures is compared between species, clear differences can be
observed. The arrangement of the large wax glands is similar in A. oblongoguttatum, A. cajennense, A. parvum, A.
rotundatum, A. pacae, A. aureolatum and A. brasiliense, but is distinct for larvae of A. longirostre and A. ovale by
the presence of two pairs of large wax glands on the 4th and 5th festoons (segment XII and XIII) (Arzua 2002;
Barbieri et al. 2007; Barbieri et al. 2008a; Barbieri et al. 2008b; Sanches et al. 2009). Similar patterns are also
observed for small glands between Amblyomma species, but some differences can aid differentiation of species,
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such as the number of small glands distributed on the idiosoma and the association between two types of structure
in the same segment. For example, segment IX series 6 in the ventral surface of the A. oblongoguttatum larva has
the lyrifissure (LvIX6) only, and the same is seen in A. cajennense and A. parvum; on the other hand, in A.
longirostre, A. rotundatum, A. ovale and A. pacae this segment and series has the lyrifissure and small gland
(LvIX6; SvIX6) (Barbieri et al. 2007; Barbieri et al. 2008a; Barbieri et al. 2008b).
Currently, larval morphological descriptions are available for only 17 Amblyomma species out of the 30
species from the Brazilian tick fauna (Amorim & Serra-Freire 1994a; Amorim & Serra-Freire 1994b; Amorim &
Serra-Freire 1995; Amorim & Serra-Freire 1996; Amorim & Serra-Freire 1999a; Amorim & Serra-Freire 1999b;
Amorim & Serra-Freire 2000; Arzua 2002; Barros-Battesti et al. 2005; Estrada-Peña et al. 1993; Famadas et al.
1997; Guglielmone et al. 1990; Barbieri et al. 2008a; Barbieri et al. 2008b; Sanches et al. 2009). While all these
descriptions provide general measurements of the body and chaetotaxy, complete porotaxy (lyrifissures, small
glands, and large wax glands) has been reported for only six species (A. cajennense, A. parvum, A. rotundatum, A.
longirostre, A. ovale, and A. pacae) (Barbieri et al. 2007; Barbieri et al. 2008a; Barbieri et al. 2008b). Since
chaetotaxy is generally highly conserved among Amblyomma spp. (Barbieri et al. 2007), porotaxy is indeed a
valuable target to be included in a taxonomic key for larvae of the Neotropical Amblyomma species, yet to be
constructed.
References
Amorim, M & Serra-Freire, N.M. (1994a) Amblyomma nodosum Neumann, 1899 descrição morfológica do estádio de larva.
Revista Brasileira de Parasitologia Veterinária, 3, 131–142.
Amorim, M. & Serra-Freire, N.M. (1994b) Descrição morfológica do estádio de larva de carrapato (Acari: Ixodidae). 4.
Amblyomma dissimile Koch, 1844. Boletim Museu Paraense Emílio Goeldi, série Zoológica, 10, 273–288.
Amorim, M & Serra-Freire, N.M. (1995) Descrição morfológica do estádio de larva de carrapato (Acari: Ixodidae). 1.
Amblyomma rotundatum Koch, 1844. Parasitologia al Dia, 19, 9–19.
Amorim, M. & Serra-Freire, N.M. (1996) Morphological description of tick larval stage (Acari: Ixodida). 3. Amblyomma
varium Koch, 1844. Entomologia y Vectores, 3, 67–81.
Amorim, M. & Serra-Freire, N.M. (1999a) Descrição morfológica do estádio de larva de carrapato (Acari: Ixodidae). 6.
Amblyomma cooperi Nuttall & Warburton, 1907. Entomologia y Vectores, 6, 126–155.
Amorim, M. & Serra-Freire, N M. (1999b) Descrição morfológica do estádio de larva de carrapato (Acari: ixodidae). 8.
Amblyomma geayi Neumann, 1899. Entomologia y Vectores, 6, 517–554.
Amorim, M. & Serra-Freire, N.M. (2000) Morphological description of tick larval stage (Acari: Ixodidae). 7. Amblyomma
auriculare (Conil, 1878). Entomologia y Vectores, 7, 297–310.
Aragão, H.P. & Fonseca, F. (1961) Notas de Ixodologia.VIII. Lista e chave dos representantes da fauna ixodológica brasileira.
Memórias do Instituto Oswaldo Cruz, 59, 115–129.
Arzua, M. (2002) Bioecologia do parasitismo de carrapatos (Acari: Ixodidae) em aves silvestres do Bosque Reinhard Maack,
Curitiba, Paraná. Descrição, diagnóstico morfológico e molecular da larva de Amblyomma aureolatum (Pallas, 1772).
Curitiba. Universidade Federal do Paraná, MSc Dissertation. 88 pp.
Arzua, M., Onofrio, V. C. & Barros-Battesti, D.M. (2005) Catalogue of the tick collection (Acari: Ixodida) of the Museu de
História Natural Capão da Imbuia, Curitiba, Paraná, Brazil. Revista Brasileira de Zoologia, 22, 623–632.
Barbieri, F.S., Chacón, S.C., Labruna, M.B., Barros-Battesti, D.M., Faccini, J.L.H. & Famadas, K.M. (2007) Topographical and
numerical study of the idiosomal integumental structures of the larva of four Neotropical Amblyomma (Acari: Ixodidae)
species. Systematic Parasitology, 68, 57–70.
Barbieri, F.S., Brito, L.G., Labruna, M.B., Barros-Battesti, D.M., Camargo, L.M.A. & Famadas, K.M. (2008a) Description of
the larva of Amblyomma ovale Koch, 1844 (Acari: Ixodidae) by light and scanning electron microscopy. Systematic and
Applied Acarology, 13, 109–119.
Barbieri, F.S., Brito, L.G., Labruna, M.B., Barros-Battesti, D.M., Camargo, L.M.A. & Famadas, K.M. (2008b) Description of
the larva of Amblyomma pacae Aragão, 1911 (Acari: Ixodidae) by light and scanning electron microscopy. Systematic and
Applied Acarology, 13, 195–203.
Barros-Battesti, D.M.; Arzua M.; Rebello, V.M.M.; Barbieri, F.S. & Famadas, K.M. (2005) Description of the larva of
Amblyomma longirostre (Koch 1844) (Acari: Ixodidae) by light and scanning electron microscopy. Revista Brasileira de
Parasitologia Veterinária, 14, 51–57.
Clifford, C.M. & Anastos, G. (1960) The use of chaetotaxy in the identification of larval ticks (Acarina: Ixodidae). Journal of
Parasitology, 46, 567–578.
Dantas-Torres, F., Onofrio, V. C. & Barros-Battesti, D.M. (2009) The ticks (Acari: Ixodida: Argasidae, Ixodidae) of Brazil.
Systematic and Applied Acarology, 14, 30–46.
Dinnik, J. & Zumpt, F. (1949) The integumentary sense organs of the larvae of Rhipicephalinae (Acarina). Psyche, 56, 1–17.
THE LARVA OF AMBLYOMMA OBLONGOGUTTATUM
Zootaxa 3579 © 2012 Magnolia Press ·
87
Estrada-Peña, A., Guglielmone, A.A., Mangold, A.J. & Castellá, J. (1993) A description of Amblyomma tigrinum Koch, A.
neumanni Ribaga and A. testudinis (Conil) immatures (Acarina: Ixodidae). Folia Parasitologica, 40, 147–153.
Famadas, K.M., Serra-Freire, N.M. & Lanfredi, R.M. (1997) Redescription of the larva of Amblyomma cajennense (Fabricius)
(Acari: Ixodidae) using optical and scanning electron microscopy. Acarologia, 38, 101–109.
Guglielmone, A.A., Mangold, A.J. & Keirans, J.F. (1990) Redescription of the male and female of Amblyomma parvum
Aragão, 1908 and description of nymph and larva, and description of all stages of Amblyomma pseudoparvum n.sp. (Acari:
Ixodida: Ixodidae). Acarologia, 32, 143–159.
Guglielmone, A.A., Beati, L., Barros-Battesti, D.M., Labruna, M.B., Nava, S., Venzal, J.M., Mangold, A.J., Szabó, M.P.J.,
Martins, J.R., González-Acuña, D. & Estrada-Peña, A. (2006) Ticks (Ixodidae) on humans in South America.
Experimental and Applied Acarology, 40, 86–100.
Hess, E. & Vlimant, M. (1983) The tarsal sensory system of Amblyomma variegatum Fabricius (Ixodidae: Metastriata). III.
Mapping of sensory hairs and evolution of relative importance of sensory modalities during post-embryonic development.
Revue Suisse Zoologie, 90, 887–897.
Keirans, J.E., Clifford, C.M. & Corwin, D. (1976) Ixodes sigelos n. sp. (Acarina: Ixodidae), a parasite of rodents in Chile, with
a method for preparing ticks for examination by scanning electron microscopy. Acarologia, 18, 217–225.
Klompen, J.S.H., Keirans, J.E., Filippova, N.A. & Oliver, J.H. JR. (1996) Idiosomal lyrifissures, setae, and small glands as
taxonomic characters and potencial indicators of ancestral segmentation patterns in larval Ixodidae (Acari: Ixodida).
International Journal of Acarology, 22, 113–134.
Labruna, M.B., Jorge, R.S., Sana, D.A., Jácomo, A.T., Kashivakura, C.K., Furtado, M.M., Ferro, C., Perez, S.A., Silveira, L.,
Santos Jr, T.S., Marques, S.R., Morato, R.G., Nava, A., Adania, C.H., Teixeira, R.H., Gomes, A.A., Conforti, V.A.,
Azevedo, F.C., Prada, C.S., Silva, J.C., Batista, A.F., Marvulo, M.F., Morato, R.L., Alho, C.J., Pinter, A., Ferreira, P.M.,
Ferreira, F. & Barros-Battesti, D.M. (2005) Ticks (Acari: Ixodida) on wild carnivores in Brazil. Experimental and Applied
Acarology 36, 149–163.
Nawar, M.S. & Madbouly, M.H. (1985) The possible use of integumentary sense organs as specific diagnostic mean for larvae
and nynphs of Rhipicephalus sanguineus and Rhipicephalus turanicus. Journal Egyptian Society Parasitology, 15,
607–621.
Martins, T.F., Onofrio, V. C., Barros-Battesti, D.M. & Labruna M.B. (2010) Nymphs of the genus Amblyomma (Acari:
Ixodidae) of Brazil: descriptions, redescriptions, and identification key. Ticks and Tick-borne Diseases, 1, 75–99.
Onofrio, V.C., Labruna, M.B., Pinter, A., Giacomin, F.G. & Barros-Battesti, D.M. (2006) Comentários e chaves para as espécies
do gênero Amblyomma. In: Barros-Battesti, D.M., Arzua, M. & Bechara, G.H. (Eds.) Carrapatos de importância médicoveterinária da Região Neotropical: um guia ilustrado para identificação de espécies. São Paulo, Brasil, Vox/ICTTD-3/
Butantan. pp. 53–113.
Robinson, L.E. (1926) Ticks. A monograph of the Ixodoidea IV.—The genus Amblyomma. Cambridge University Press,
London, 302 pp.
Sanches, G.S., Bechara, G.H. & Camargo-Mathias, M.I. (2009) Morphological description of Amblyomma brasiliense Aragão,
1908 (Acari: Ixodidae) larvae and nymphs. Revista Brasileira de Parasitologia Veterinária, 18, 15–21.
Schulze, P. (1942) Ueber die Hautsinnesorgane der Zecken, besonders uber eine bisher unbekannte Art von Arthropodensinnesorgane, die Krobylophoren. Zeitschrift fuer Morphologie und Okologie der Tiere, 39, 1–20.
Wooley, T.A. (1988) Acarology. Mites and human welfare. A Wiley-Interscience Publications, Wiley, J. & Sons Ed, New York,
484 pp.
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