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Determination of the karyotype of the giant anteater

Myrmecophaga tridactyla (Myrmecophagidae:
Xenarthra) using classical and molecular cytogenetic
techniques
a a a a
L. F. Rossi , M. G. Chirino , J. P. Luaces & M. S. Merani
a
Laboratorio de Biologa Cromosmica, Facultad de Medicina, Universidad de Buenos Aires,
Buenos Aires, Argentina
Published online: 31 Jul 2014.

To cite this article: L. F. Rossi, M. G. Chirino, J. P. Luaces & M. S. Merani (2014): Determination of the karyotype of the giant
anteater Myrmecophaga tridactyla (Myrmecophagidae: Xenarthra) using classical and molecular cytogenetic techniques,
Italian Journal of Zoology, DOI: 10.1080/11250003.2014.943308

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 Italian Journal of Zoology, 2014, 16
http://dx.doi.org/10.1080/11250003.2014.943308

Determination of the karyotype of the giant anteater Myrmecophaga

tridactyla (Myrmecophagidae: Xenarthra) using classical and
molecular cytogenetic techniques

L. F. ROSSI, M. G. CHIRINO, J. P. LUACES, & M. S. MERANI

Laboratorio de Biologa Cromosmica, Facultad de Medicina, Universidad de Buenos Aires, Buenos Aires, Argentina
Downloaded by [University of Newcastle (Australia)] at 12:04 29 August 2014

(Received 10 March 2014; accepted 30 June 2014)

Abstract
The karyotype of the giant anteater Myrmecophaga tridactyla was studied throughout the species Argentine range. Images of
the chromosome complement clearly revealed the karyotype and identied previously misinterpreted chromosomes. The
peripheral blood lymphocytes of 26 adult animals (11 males and 15 females) were cultured and used to obtain mitotic
metaphases. These preparations were subjected to conventional staining, G- and C-banding, and Fluorescent in situ
Hibridization (FISH). Spermatocyte pachytene microspreads for one adult were examined for synaptonemal complexes.
The karyotype (2n = 60 XX/XY; Fundamental Number (FN) = 110 without XY) showed an autosomal complement of six
metacentric, 18 submetacentric, and six acrocentric chromosome pairs. The X chromosome (4.67 0.29% of the haploid
set) was shown to be large and submetacentric, similar in size to autosomes 35. The Y chromosome was submetacentric
(2.60 1.08% of the haploid set). It is, however, larger than the Y chromosome of the closely related armadillos. Pairs 1, 3,
4, 5, 6, 9, 11, 14, 15, 23, 25 and 29 showed small masses of heterochromatin in the pericentromeric region. These masses
were slightly larger in chromosome pairs 8, 10 and 18. Pairs 2, 7, 16, 17, 19, 20, 21, 22, 24, 26, 27 and 28 were entirely C-
negative. Analysis of the telomeric sequences by FISH involving a Cy3-conjugated peptide nucleic acid pantelomeric probe
revealed no signals from the interstitial regions. Ag-NORs were located on chromosomes 5, 10, 11, 21 and 23. The
spermatocyte pachytene microspreads conrmed the morphology and size of both sex chromosomes. The present results,
obtained via the analysis of a large number of specimens, provide an in-depth characterization of the chromosomal
complement of this species.

Keywords: Chromosome, Xenarthra, anteaters, banding pattern, lymphocyte cultures

Introduction
The extant members of Xenarthra eminently
Central and South American placental mammals
form three phylogenetic groups: the arboreal sloths
(Folivora: Megalonychidae and Bradypodidae), the
shelled armadillos (Cingulata: Dasypodidae) and the
toothless anteaters (Vermilingua: Myrmecophagidae
and Cyclopedidae) (Gardner 2008). Chromosomal
polymorphisms, pairs of heteromorphic autosomes
of different lengths (Benirschke & Wurster 1969;
Jorge et al. 1978; Rossi et al. 2014), and unusual
sex chromosome systems (Corin-Frederic 1969;
Jorge et al. 1985) have been described in some spe-
cies of Xenarthra. A number of cytogenetic studies
have been performed but none have involved large
numbers of specimens from different locations.
The family Myrmecophagidae contains two gen-
era: Myrmecophaga and Tamandua. Myrmecophaga
comprises only one species, the giant anteater
Myrmecophaga trydactyla, which has, according to
preliminary reports, 60 chromosomes, some of
which are acrocentric (Hsu 1965; Pereira et al.
2004). Tamandua has two species of vested antea-
ter, Tamandua tetradactyla and Tamandua mexi-
cana, neither of which have acrocentric
chromosomes. Two chromosome numbers have
been reported for T. tetradactyla (Pereira et al.
2004). Diploid numbers for the family
Myrmecophagidae range from 2n = 54 to
2n = 60. Reciprocal translocations and reversible
fusions/ssions may explain these differences
(Jorge et al. 1985).

Correspondence: M. S. Merani, Laboratorio de Biologa Cromosmica, Facultad de Medicina, Universidad de Buenos Aires, Paraguay 2155 P10, C1121ABG
Buenos Aires, Argentina. Tel: +54 11 5950 9500 ext. 2153. Fax: +54 11 5950 9612. Email: mmerani@fmed.uba.ar

2014 Unione Zoologica Italiana

 2 L. F. Rossi et al.

Myrmecophaga tridactyla the giant anteater is Table I. List of studied individuals with their respective sex and
recognized as vulnerable by the ISCN (Shaffer et al. localities of provenance.
2009). Although the karyotype of this species has been Provenance
described in two studies, both involved only conven-
tional staining techniques and just a few individuals of Individual no. Locality Province Sex
unknown provenance living in captivity (Hsu 1965; 1 Dto. Alberdi Sgo. del Estero M
Pereira et al. 2004). For example, no G-banding 2 Monte Rico Sgo. del Estero M
patterns were reported that might allow the sex 3 Monte Quemado Sgo. del Estero M
chromosomes to be identied. Thus, if the karyotype 4 Pampa de los Guanacos Sgo. del Estero F
5 Campo Gallo Sgo. del Estero M
of this species is to be reliably understood, studies
6 Copo Sgo. del Estero M
involving cytogenetic and resolution banding techni- 7 Dto. Alberdi Sgo. del Estero F
ques are required. This study reports the complete 8 El Cabur Sgo. del Estero M
cytogenetic characterization of M. tridactyla. Mitotic 9 Pje Monte Rico Sgo. del Estero F
spreads were examined by G- and C-banding, 10 Los Pirpintos Sgo. del Estero F
11 Sacha yo Sgo. del Estero F
Nucleolus Organizer Regin (NOR) staining and
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12 Copo Sgo. del Estero F

Fluorescent in situ Hibridization (FISH), and meiotic 13 Pampa del Inerno Chaco M
spreads via synaptonemal complex (SC) analysis. 14 Roque Saenz Pea Chaco M
15 Roque Saenz Pea Chaco F
16 Gral. Pinedo Chaco M
Materials and methods 17 Roque Saenz Pea Chaco F
18 Quitilipi Chaco F
Sampled specimens 19 L. Cabral Salta M
20 La esperanza Salta F
The animals studied were adult specimens (11 21 San Agustn Salta F
males and 15 females) of M. tridactyla housed at 22 Curva del Turco Salta F
various zoos, breeding centres and wildlife reintro- 23 El Mistol Salta F
duction facilities in different parts of Argentina 24 Las Flores Salta F
25 Las Lajitas Salta F
(Table I). After immobilization using Telazol 26 Tostado Santa Fe M*
(4 mg/kg) with a Telinject (remote injection) sys-
tem, peripheral blood samples were collected with * Meiotic bivalent and sex chromosome analysis.
disposable heparinized syringes. All handling was
performed according to the Guidelines of the
American Society of Mammalogists for the Use of Silver nitrate staining of mitotic and meiotic mate-
Wild Mammals in Research (Sikes et al. 2011). rial was performed according to Sciurano et al.
(2006). All preparations were photographed using a
Leitz DMRB microscope (Leica Microsystems,
Mitotic preparations Wetzlar, Germany) and a Leica DFC 300 FX digital
camera (Leica Microsystems, Cambridge, UK).
Lymphocytes were cultured for 72 h at 34C in RPMI Ideograms were prepared for all chromosomes.
1640 Medium (Roswell Park Memorial Institute, The morphology of the individual chromosomes of
Gibco) according to Moorhead et al. (1960). three cells was characterized and karyotypes
Metaphase chromosome spreads were prepared assembled in the conventional manner (Shaffer
following the usual air-drying methods, and then et al. 2009).
stained with carbol fuchsine (Carr & Walker 1961).
G-banding was performed following the method of
Wang and Fedoroff (1972), which involves trypsin
Fluorescence in situ hybridization
treatment and subsequent staining with 5% Giemsa
solution at pH 6.8. C-banding was performed accord- The telomeric distribution of two complementary
ing to Sumner (1972). The slides were incubated with oligonucleotides (Telo1, TTAGGG7 Oligo number
a supersaturated solution of barium hydroxide 203006A623H01 , and Telo2, GGGTTA7 Oligo
(BaOH) at 50C, and then further incubated for 1 h number 20306A623H02 2/2) was analysed by FISH
in 2XSSC solution at 60C. They were then stained in metaphase spreads, using a Cy3-conjugated pep-
with 1.5% Giemsa solution at pH 6.8 for 15 min tide nucleic acid (PNA) pantelomeric probe (Biofab
(Sumner 1972). G-banding was carried out in not Research, Roma, Italy). FISH of the repetitive
less than ve metaphases for each specimen. At least sequences was performed following standard proce-
10 G- and 10 C-banded metaphases were selected to dures (Lichter et al. 1992). The slides were then
be used in the production of ideograms. conventionally stained with propidium iodide and
 Classical and molecular cytogenetic of Myrmecophaga tridactyla
mounted in Vectashield medium (Vector
Laboratories, Peterborough, UK). DAPI (4,6-dia-
midino-2-phenylindole) counterstaining facilitated
the identication of homologues. Signals were
observed at 1000 using a Leica DM epiuores-
cence microscope (Leica Microsystems, Wetzlar,
Germany) equipped with an HBO 50 mercury
lamp and lters for DAPI and Cy3 (Chroma
Technology, Bellows Falls, USA). A Leica DFC
300 FX digital camera (Leica Microsystems) was
used for all photography. Images were processed
using Adobe Photoshop CS software (Adobe
Systems Inc.).
Downloaded by [University of Newcastle (Australia)] at 12:04 29 August 2014
Meiotic bivalent and sex chromosome analysis
Testicular material was obtained from an animal
(specimen number 25) that died of natural causes.
Spermatocyte microspreads for SC analysis were
prepared as described by Sciurano et al. (2012).
Some slides were stained with 4% phosphotungstic
acid in ethanol or silver nitrate (Howell & Black
1980), while others were kept at 70C until examina-
tion by immunouorescence microscopy. The immu-
nolocalization of meiotic proteins was performed
according to Sciurano et al. (2012). The following
primary antibodies were used: rabbit anti-SMC3
(1:500) (Merck Millipore, Billerica, MA, USA) to
recognize components of the cohesion complex and
thus identify bivalents; and human CREST serum
that binds to kinetochores (1:10) (Laboratorios IFI,
Buenos Aires, Argentina). Slides were examined
using a LEICA DM epiuorescence microscope
(Leica Microsystems) and photographed with a Leica
DFC 300 FX digital camera (Leica Microsystems).
Separate images were superimposed using Adobe
Photoshop CS software (Adobe Systems Inc.).
Figure 1. (a), Karyotype of a male Myrmecophaga tridactyla from Alberdi, Province of Santiago del Estero. (b), G-banded karyotype of
M. tridactyla from Copo, Province of Santiago del Estero. (c), Ideogram of a male M. tridactyla based on the G-banded karyotype Scale bars:
10 m.
3
Results
Mitotic studies
All individuals showed a chromosome number of 60
(58 + XX/XY; female/male), with six metacentric
and 18 submetacentric chromosome pairs, all clearly
two-armed, plus six acrocentric pairs (Figure 1a).
The fundamental number (FN) was 110. The only
difference in the karyotypes was the morphology of
chromosome pairs 2 and 3, which were hetero-
morphic in 65% of individuals. This polymorphism,
involving the absence of small arms, is probably the
result of pericentromeric inversion.
The X chromosome (4.67 0.29% of the haploid
set) was large and submetacentric, and similar in size
to autosomes 35. The Y chromosome was submeta-
centric (2.60 1.08% of the haploid set) and it is
between medium and smaller chromosomes
(Figure 1a). G-banding accurately identied all the
autosomal pairs and the X and Y chromosomes. The
homology between the two elements of each pair and
the X and Y chromosome was checked by G-banding
comparisons (Figure 1b). The ideogram (Figure 1c)
is a composite representation of all the G-bands seen
in 10 well-banded karyotypes (it indicates the average
location of each of the bands detected).
The C-banding pattern in pairs 1, 3, 4, 5, 6, 9,
11, 14, 15, 23, 25 and 29 showed small masses of
heterochromatin in the pericentromeric region;
these were slightly larger in pairs 8, 10 and 18.
Other pairs were entirely C-negative (pairs 2, 7,
16, 17, 19, 20, 21, 22, 24, 26, 27 and 28).
C-banding analysis revealed no pericentromeric
constitutive heterochromatin in the X and Y chro-
mosomes (Figure 2a).
Five nucleolar organizing regions (Ag-NORs), one
on each of chromosome pairs 5, 10, 11, 21 and 23,
 4 L. F. Rossi et al.
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Figure 2. (a), C-banded karyotype of a male Myrmecophaga tridactyla from the Province of Santiago del Estero, showing centromeric C+
heterochromatin. (b), NOR-banded karyotype of a male M. tridactyla from the Province of Salta. Identication of the NORs (Arrowhead).
(c), SC karyotype of a male M. tridactyla from Tostado, Province of Santa Fe. Each bivalent is represented by 1 SC immunolabeled with
anti-SMC3 (red), anti-kinetochore (yellow). (d), Metaphase plate of a male M. tridactyla from the Province of Salta, showing the location of
the telomeric probes. The chromosomes were counterstained with DAPI. Scale bars: 10 m.

were observed in most cells (Figure 2b). The SC
karyotype showed 29 autosomal SCs, 24 autosomal
SCs with meta or submetacentric components and
ve autosomal SCs with acrocentric elements. The
location of the centromeres (kinetochores) was
observed and veried by inmunolocalization
(Figure 2c). The spermatocyte pachytene micro-
spreads corroborated the chromosomal morphology
and FN (Figure 2c) and clearly showed the sex
chromosome kinetochores. Hybridization of the
probe with the telomeric sequences revealed no
regions with interstitial signals (Figure 2d).
Discussion
The present work ascertained the karyotype of M.
tridactyla using classical and molecular cytogenetic
techniques. All chromosomes were identied, includ-
ing some that had previously been misinterpreted.
All the animals examined had a chromosome com-
plement of 2n = 60, conrming the results of other
authors (Hsu 1965; Pereira et al. 2004). Pereira et al.
(2004) reported the giant anteater to have ve pairs
of small- and medium-sized acrocentric
chromosomes. In the present work, however, six
pairs of acrocentric chromosomes were seen; one of
the medium-sized submetacentric pairs described by
the latter authors was in fact a medium-sized acro-
centric pair.
The sex chromosomes showed morphologies dif-
ferent to those described by Pereira et al. (2004) for
giant anteaters from Brazil. In the present work, the
X chromosome was large and submetacentric, and
the Y chromosome was also submetacentric. It was
larger than that of the closely related armadillos; in
armadillos, the Y chromosome is the smallest of the
entire complement (Sciurano et al. 2012; Rossi et al.
2014). Pereira et al. (2004) reported the Y chromo-
some to be small and acrocentric. G- and C- band-
ings conrmed the presence of chromosome
designations and identied those erroneously paired
by these authors.
The differences in the distribution of the C-positive
regions in Myrmecophagidae species reect the varia-
tion in the content, location and distribution of the
constitutive heterochromatin in this family and within
Xenarthra (Dobigny et al. 2005; Liu et al. 2011; Rossi
et al. 2014). M. tridactyla is characterized by the
 Classical and molecular cytogenetic of Myrmecophaga tridactyla
presence of small masses of heterochromatin in most
of its chromosomes, whereas T. tetradactyla has
ve pairs of chromosomes with large centromeric
heterochromatin regions (Dobigny et al. 2005).
Interestingly, no telomeric signals were found in M.
tridactyla, whereas T. tetradactyla has ve pairs of
chromosomes that show interstitial telomeric signals
(unpublished results). It is well known that clusters of
different sequences of repetitive DNA, including telo-
meric repeats in subtelomeric and interstitial posi-
tions, are useful for characterizing the breakpoints of
recurrent chromosomal rearrangements (Azzalin
et al. 2001; Nergadze et al. 2004). The C-banding
pattern and telomeric sequence of M. tridactyla may
clarify the chromosomal mechanisms underlying the
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karyotype evolution of the family Myrmecophagidae.
The difference in chromosome number between M.
tridactyla and T. tetradactyla species might be
explained by Robertsonian translocation.
Species with more primitive karyotypes show the
smallest number of NORs (Chiarelli & Capanna
1973). It is well documented that different groups
of species have acquired more NORs over time (Hall
& Parker 1995; Hirai et al. 1996). Among the
Xenarthrans, Dasypus hybridus (2n = 64, FN = 79)
(Saez et al. 1964) has one NOR (Sciurano et al.
2006), Chaetophractus villosus has three (Sciurano
et al. 2006; Rossi et al. 2014) and M. tridactyla has
ve (as revealed by the Ag-NOR technique).
The SCs in spermatocytes examined during early
prophase showed the X and Y chromosomes of M.
tridactyla to be submetacentric, with a clearly dened
short arm. This was not reported by previous authors
(Hsu 1965; Pereira et al. 2004).
The present results update the available informa-
tion on the karyotype of the giant anteater. Modern
cytogenetic techniques may be able to show
how changes in the chromosomes affected the nal
complement of this species and the family
Myrmecophagidae.
Acknowledgements
M.S. Merani was supported by grant PICT 1198
from the Agencia Nacional de Promocin Cientca y
Tecnolgica and grant PIP 0204 from the Consejo
Nacional de Investigaciones Cientcas y Tcnicas.
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