Parasitol Res (2009) 105:1629–1636
DOI 10.1007/s00436-009-1605-8
ORIGINAL PAPER
Morphotypes of Varroa destructor collected in Apis mellifera
colonies from different geographic locations of Argentina
Matías D. Maggi & Norma H. Sardella &
Sergio R. Ruffinengo & Martín J. Eguaras
Received: 30 June 2009 / Accepted: 14 August 2009 / Published online: 29 August 2009
# Springer-Verlag 2009
Abstract Parasites display considerable phenotypic plasticity
in life-history traits such as, body size. Varroa destructor is an
ectoparasitic mite of the western honey bee Apis mellifera.
Several studies have reported that in V. destructor, there is a
wide phenotypic plasticity within a population of mites.
However, it is unknown if there are morphologic variations
in V. destructor populations affecting different A. mellifera
populations. A morphometric study of V. destructor pop-
ulations was conducted to provide information concerned to
the relationships among parasite populations found in
different geographic locations from A. mellifera colonies of
Argentina. The hypothesis tested was different morphotypes
of V. destructor populations parasitizing different A. mellifera
populations from Argentina exist. A discriminant analysis
employing eight morphologic variables revealed that it is
M. D. Maggi : M. J. Eguaras
Laboratorio de Artrópodos, Facultad de Ciencias Exactas y
Naturales, Universidad Nacional de Mar del Plata,
Funes, 3350,
7600 Mar del Plata, Argentina
M. D. Maggi (*) : N. H. Sardella : M. J. Eguaras
CONICET, Consejo Nacional de Investigaciones Científicas y
Técnicas, Rivadavia 1917,
C1033AJ Buenos Aires, Argentina
e-mail: biomaggi@gmail.com
N. H. Sardella
Laboratorio de Parasitología, Facultad de Ciencias Exactas y
Naturales, Universidad Nacional de Mar del Plata,
Funes 3350,
7600 Mar del Plata, Argentina
S. R. Ruffinengo
Apicultura, Facultad de Ciencias Agrarias,
Universidad Nacional de Mar del Plata,
Ruta Nacional 226, Km. 73.5,
CC 276 (B7620ZAA) Balcarce, Argentina
possible to differentiate morphotypes of mites in Argentina.
However, the level of discrimination detected among mites
population varied according to the grouping of mite’s
population. Possible causes explaining the morphometric
variability in the V. destructor populations were discussed.
Introduction
Phenotypic plasticity is classically defined as occurring
when the phenotype expressed by a given genotype is
altered as environmental conditions change (Pigliucci 2005;
Nussey et al. 2007). Plasticity allows an organism to “fit”
its phenotype to the changeable environment; when the
optimal phenotype varies with environmental conditions
experienced, the evolution of phenotypic plasticity is
predicted (Van Tienderen 1991). In parasite populations,
adjustments of life-history strategies such as body size to
changing environmental conditions can take place like a
response to habitats highly variable spatiality or over short
time periods (Stearns 1992). According to Poulin (1998),
the habitat of parasites is not spatially continuous but
consists of discrete cells or islands. Hosts represent patches
of suitable habitat in an otherwise inhospitable environ-
ment. Since parasite habitat is not uniform, they have to
affront the imposed selective pressures established by the
host ambient adjusting its biology to ensure the continuity
of its life (Poulin 1998; Vizoso and Ebert 2005).
The genus Varroa (Acari: Varroidae) is currently repre-
sented by three highly specialized species of obligate
ectoparasitic mites that feed on the hemolymph of social
cavity-nesting bees (Apis spp.) in Asia. Varroa jacobsoni has
the widest distribution and is defined as the species that
parasitizes both, the Asiatic Apis cerana and the European
Apis mellifera bees, being difficult to establish why the same
1630
mite species produces different virulence on both bee species
(Delfinado-Baker 1988). The first investigations about these
differences were focused on the characterization of mite
populations affecting different bee species. Parasite popula-
tions could be discriminated when morphometric studies
were done (Delfinado-Baker and Houck 1989). Later, with
the development of molecular techniques, it could be
demonstrated that V. jacobsoni was a complex of at least
two different species affecting the genus Apis disseminated
around the world (Anderson and Fuchs 1998; De Guzman et
al. 1999; Warrit et al. 2004; Zhou et al. 2004). Anderson and
Trueman (2000) established that V. jacobsoni is an exclusive
parasite of A. cerana, and V. destructor is the new species
parasitizing A. mellifera. However, V. destructor is not native
to the Americas, Europe, Africa, or even to A. mellifera
itself. Varroa destructor was originated on the mainland
Asian continent, where it infests A. cerana. According to
Anderson and Trueman (2000), only two V. destructor
haplotypes (mitochondrial genotypes)—the Korean and
Japan—Thailand haplotypes—have colonized A. mellifera.
Currently, this mite causes serious damage by feeding on
hemolymph of larvae, pupae, and adults, and by transmitting
or activating viral diseases (Shen et al. 2005; Chen et al.
2006).
Studies involving relationships between phenotypes and
genotypes of both, hosts, and parasites, could help to
understand their distribution and speciation, as well as their
interaction (Lambrechts et al. 2006). Anderson and Trueman
(2000) have demonstrated that a clear correlation between
A. cerana and V. jacobsoni haplotypes exists, and they
suggested that novel variants of a host could lead to novel
variants of its parasites. With regard to phenotypes, it was
reported by Delfinado-Baker and Houck (1989) and by
Anderson and Trueman (2000) that bee size is correlated with
mite size, so V. jacobsoni which is smaller than V. destructor,
affects A. cerana which is smaller than A. mellifera.
When morphometric characters were compared between
populations of mites parasiting the European honey and the
Asiatic honey bees, no significant differences were reported
(Anderson and Trueman 2000). Nevertheless, several studies
have registered that in V. destructor species there is a wide
phenotypic plasticity within a same mite population (Akimov
and Yastrebtsov 1985; Akimov and Zaloznaya 1996;
Akimov et al. 1990, 2004). A number of publications have
been carried out with the aim to study the morphometric
variations of A. mellifera populations, and different morpho-
types of the bee were established (Buco et al. 1987; Radloff
and Hepburn 2000; Kandemir et al. 2000). However, it is
unknown if there are morphologic variations in V. destructor
populations affecting different A. mellifera populations.
The aim of this research was to study the morpho-
logical plasticity of V. destructor among populations of
mites affecting A. mellifera in Argentina, with the purpose
Parasitol Res (2009) 105:1629–1636
to expand the ecologic and epidemiologic knowledge of
the parasitism caused by the mite. The hypothesis tested
was that different morphotypes of V. destructor popula-
tions parasitizing different A. mellifera populations from
Argentina exist.
Materials and methods
The study was conducted in the Apiculture Laboratory
located on the inter-beach Rout 11 in the coastal station of
J. J. Nágera (38º10′06″S; 57º38′10″W) as well as in the
Arthropods Laboratory of the School of Exact and Natural
Sciences of the Universidad Nacional de Mar del Plata,
between April and October 2007.
Collection of mites
A total of 360 V. destructor adult female specimens
belonging to 12 populations from Argentina, were studied
(Fig. 1a). Populations were selected based on geographic
locations with an inter-locality distance higher than 200 km
and were collected from two apiaries of A. mellifera in each
place, following the sampling method of De Jong et al.
(1982). Sampling was carried out between summer 2006
and spring 2007. Each sample was labeled with the name of
the geographic location and stored in alcohol 70% v/v until
its utilization.
Processing of mites
Varroa mites were processed for its observation following
techniques described by Grandjean (Krantz 1978). Each
specimen was placed in lactic acid 50% and was cleared
during 2 hours at 100ºC. Mites were stored in alcohol 50%
v/v until their observation. Morphometric characters were
measured using a stereoscopic microscope (Lancet, Series
2000), fitted with an ocular micrometer.
Morphometry
Eight variables were measured on each of the 360 speci-
mens (Fig. 1b): width of dorsal shield (WDS), length of
dorsal shield (LDS), width of genital shield (WGS), length
of genital shield (LGS), width of metapodal shield (WMS),
large of metapodal shield (LMS), width of anal shield
(WAS), and length of anal shield (LAS).
Statistical analysis
Statistical analysis was carried out using the free R
software, following statistics proceedings described by R
Development Core Team (2009).
Parasitol Res (2009) 105:1629–1636
Fig. 1 a Geographic locations of V. destructor populations from
Argentina: 1 Jujuy, 2 Tucumán, 3 Santiago del Estero, 4 Santa Fé, 5
Misiones, 6 Corrientes, 7 Entre Ríos, 8 San Luis, 9 Buenos Aires, 10
Neuquén, 11 Río Negro, and 12 Santa Cruz. b Variables measured in
Univariate (analysis of variance (ANOVA), one way) and
multivariate (MANOVA) variance analysis were used to test if
the 12 populations of V. destructor differ in the measured
variables. To identify morphometric relationships among
populations of mites and significant differences, a posteriori
cluster test was done (Valdano and Di Rienzo 2007).
To determine the lineal combinations of morphological
characters that best separate the a priori groups established
by the cluster analysis, a discriminant analysis was done.
Results
Multivariate analysis of variance and cluster analysis
The mean of each variable measured belonging to the 12
populations of V. destructor are shown in Table 1.
1631
V. destructor mites. WDS width of dorsal shield, LDS length of dorsal
shield, WGS width of genital shield, LGS length of genital shield,
WMS width of metapodal shield, LMS length of metapodal shield,
WAS width of anal shield, LAS length of anal shield
Multivariate analysis showed that mite populations differ
significantly according to geographic locations when all
characters were analyzed simultaneously (MANOVA,
Wilk’s lambda=0.283, P<0.001). Seven morphotypes were
established by the cluster analysis (Fig. 2; cutline B,
distance=1.108, α=0.01).
Morphotype A was established in mites collected from
Santa Cruz province; morphotype B was established in
mites belonging from Santa Fe, Río Negro, Misiones, and
San Luis provinces; morphotype C was established in mites
from Entre Ríos and Neuquén provinces; morphotype D
was established in mites from Buenos Aires and Corrientes
provinces; morphotype E was established in mites from
Tucumán province; morphotype F was established in mites
from Jujuy province; and finally, morphotype G was
established in mites from Santiago del Estero province.
The mean of each variable studied belonging to the seven
1632 Parasitol Res (2009) 105:1629–1636

Table 1 Mean of the variables

studied (millimeters) belonging WDS LDS WMS LMS WGS LGS WAS LAS
to the 12 populations of V.
destructor from Argentina Buenos Aires 1.699 1.128 0.341 0.447 0.719 0.569 0.262 0.153
Corrientes 1.687 1.121 0.348 0.444 0.708 0.550 0.261 0.151
Neuquén 1.718 1.128 0.355 0.452 0.718 0.564 0.263 0.157
Tucumán 1.694 1.143 0.357 0.437 0.702 0.534 0.256 0.145
Jujuy 1.700 1.130 0.354 0.435 0.720 0.544 0.256 0.145
Misiones 1.746 1.176 0.359 0.446 0.725 0.557 0.265 0.146
WDS width of dorsal shield, San Luis 1.752 1.182 0.359 0.446 0.722 0.553 0.263 0.142
LDS length of dorsal shield, Santa Fe 1.752 1.174 0.358 0.460 0.739 0.579 0.261 0.149
WGS width of genital shield, Santa Cruz 1.756 1.183 0.354 0.447 0.725 0.566 0.243 0.147
LGS length of genital shield,
WMS width of metapodal shield, Río Negro 1.791 1.186 0.360 0.449 0.736 0.565 0.262 0.140
LMS length of metapodal shield, S. del Estero 1.672 1.131 0.340 0.430 0.694 0.538 0.251 0.143
WAS width of anal shield, LAS Entre Ríos 1.733 1.134 0.358 0.452 0.716 0.561 0.274 0.158
length of anal shield

morphotypes of V. destructor and the ANOVA results are
shown in Tables 2 and 3, respectively.
Nevertheless, in Fig. 2, it could be established three
main morphotypes (cutline A, distance=1.4). The first
morphotype was established in mites from Río Negro,
Santa Fé, Santa Cruz, Misiones, and San Luis provinces;
the second morphotype was established in mites from
Buenos Aires, Neuquén, Corrientes, and Entre Ríos
provinces; and the third morphotype was established in
mites from Santiago de Estero, Tucumán, and Jujuy
provinces. Morphotypes 2 and 3 were phenotipically more
related between them than with morphotype 1 (Fig. 2;
Table 4). The three morphotypes differed significantly in all
measured characters (ANOVA; Table 5).

Fig. 2 Cluster dendrogram for V. destructor populations based on represents the mayor level of discrimination among parasites
eight morphometric variables studied and geographic locations where populations (α=0.01). Cutline A (distance=1.4) establish the three
parasite populations were sampled. Cutline B (distance= 1.108) main morphotypes of V. destructor
Parasitol Res (2009) 105:1629–1636 1633

Table 2 Mean of the variables studied (millimeters) belonging to the seven morphotypes of V. destructor from Argentina established by the
cluster dendrogram

WDS LDS WMS LMS WGS LGS WAS LAS

Aa 1.754 1.18 0.356 0.447 0.725 0.564 0.243 0.149

Bb 1.758 1.177 0.358 0.45 0.731 0.565 0.262 0.145
Cc 1.729 1.132 0.356 0.453 0.719 0.565 0.268 0.158
Dd 1.693 1.124 0.344 0.446 0.714 0.559 0.261 0.152
Ee 1.685 1.138 0.361 0.433 0.699 0.535 0.250 0.143
Ff 1.714 1.140 0.354 0.439 0.721 0.545 0.261 0.144
Gg 1.69 1.126 0.341 0.433 0.695 0.542 0.255 0.146

Letters (A to G) correspond to morphotypes of the Provinces

WDS width of dorsal shield, LDS length of dorsal shield, WGS width of genital shield, LGS length of genital shield, WMS width of metapodal
shield, LMS length of metapodal shield, WAS width of anal shield, LAS length of anal shield
a
Santa Cruz
b
Santa Fe + Río Negro + Misiones + San Luis
c
Entre Ríos + Neuquén
d
Buenos Aires + Corrientes
e
Tucumán
f
Jujuy
g
Santiago del Estero

Discriminant analysis was LDS (0.773), following by WDS (0.665), WGS (0.368),
LGS (0.205), WAS (−0.111), and LAS (−0.283). WMS
Based on the two groups established by the cluster analysis (0.38) and LMS (0.2784) had almost no effect on the
(group of seven and three morphotypes), two discriminant discriminant function and were removed from later analysis.
analyses were performed: The discriminant function obtained was D=0.619WDS+
The first two canonical discriminant functions were used for 0.706LDS−0.312WGS+0.124LGS−0.183WAS−0.393LAS
the discriminant analysis of the seven morphotypes (test of and explained only 50.9% of the original cases.
function 1: Wilk’s lambda=0.39, χ2 =293.89, df=36, p<0.001 The first two canonical discriminant functions were used
and test of function 2: Wilk’s lambda=0.657, χ2 =131.43, for the discriminant analysis of the three morphotypes (test
df=25, p<0.001) and explained 81.8% of all variance, of function 1: Wilk’s lambda=0.547, χ2 =195.27, df=10,
contributing to 59.9% (eigenvalue, 0.682) and 21.9% p<0.001 and test of function 2: Wilk’s lambda=0.811, χ2 =
(eigenvalue, 0.249), respectively. The variable showing a 67.95, df=4, p<0.001) and explained 100% of the variance,
larger correlation coefficient with the discriminant function 1 contributing to 67.4% (eigenvalue, 0.481) and 32.3%
(eigenvalue, 0.233), respectively.
Table 3 ANOVA of the variables studied of the seven morphotypes The variable showing a larger correlation coefficient
of V. destructor from Argentina with the discriminant function 1 was LDS (0.756) followed
Wilks' lambda F df Sig. by WDS (0.598), LGS (0.195), and LMS (0.075) while
LAS (−0.393). WGS (0.409), WMS (0.296), and WAS
WDS 0.778 15.297 6 0.000 (0.095) had almost no effect on the discriminant function
LDS 0.743 18.524 6 0.000 and were removed from later analysis. The discriminant
WMS 0.881 7.247 6 0.000 function was, D = 0.396WDS + 0.766LDS − 0.321LMS +
LMS 0.899 6.012 6 0.000 0.143LGS−0.459LAS, and correctly explained 66.9% of
WGS 0.825 11.359 6 0.000 the original cases. Figure 3 shows the three groups
LGS .859 8.832 6 0.000 established by the cluster.
WAS 0.867 8.258 6 0.000
LAS 0.861 8.697 6 0.000
Discussion
WDS width of dorsal shield, LDS length of dorsal shield WGS width
of genital shield, LGS length of genital shield, WMS width of
metapodal shield, LMS length of metapodal shield, WAS width of Morphometric studies based on multivariate analysis have
anal shield, LAS length of anal shield been successful to characterize the species of the genus
1634 Parasitol Res (2009) 105:1629–1636

Table 4 Mean of each variable studied (millimeters) belonging to the three morphotypes of V. destructor from Argentina established by the
cluster dendrogram

WDS LDS WMS LMS WGS LGS WAS LAS

1a 1.757 1.178 0.358 0.449 0.729 0.565 0.258 0.146

2b 1.711 1.128 0.351 0.449 0.716 0.562 0.265 0.155
3c 1.696 1.135 0.352 0.435 0.705 0.541 0.255 0.144

Numbers (1–3) correspond to morphotypes of provinces

WDS width of dorsal shield, LDS length of dorsal shield, WGS width of genital shield, LGS length of genital shield, WMS width of metapodal
shield, LMS length of metapodal shield, WAS width of anal shield, LAS length of anal shield
a
Río Negro + Santa Fé + Santa Cruz + Misiones + San Luis
b
Buenos Aires + Neuquén + Corrientes + Entre Ríos
c
Santiago de Estero + Tucumán + Jujuy

Apis and their parasites (Buco et al. 1987; Delfinado-Baker
and Houck 1989; Anderson and Trueman 2000; Radloff
and Hepburn 2000; Kandemir et al. 2000). In this research,
when mite populations collected from different geographic
locations of Argentina were studied, different morphotypes
of mites were found, and a maximum number of seven
groups could be established. Nevertheless, when this level
of discrimination was considered, a wide overlap of the
morphometric variables among morphotypes was regis-
tered. This fact could be explained through the discriminant
analysis where the total samples were reclassified and only
a 50.9% of them were correctly assigned to the groups they
belong to. Only when three morphotypes were considered,
the separation among groups was more evident. By this
way, the existence of different morphotypes within V.
destructor found in this research represents the first report
of phenotypic plasticity among V. destructor populations.
Previous studies conducted on the genus Varroa, have
demonstrated the existence of morphological differences
when populations of V. jacobsoni and V. destructor were
compared, being the parasites of the latter species of bigger
size (Delfinado-Baker and Houck 1989; Anderson and
Trueman 2000). However, no researches have reported the
existence of intra-specific morphological phenotypes in
these species. Akimov et al. (1988, 2004) registered a wide
morphological plasticity in V. destructor male and female
mites within a same population, relating this plasticity to
different seasons of bee reproduction: in winter, selection
was directed upon elimination of individuals that worse
adapted to phoresia on bee, while in summer, selection
upholds individuals that better adapted to reproduction. In
the present research, due to the sampling was conducted
during the warm seasons, where bee brood is always
present, possible effects of seasonality determining mor-
photypes were discarded. In addition, the three morpho-
types identified here belonged to different geographic
locations of Argentina, therefore latitudinal effects were

Table 5 ANOVA of the variables studied of the three morphotypes of

V. destructor from Argentina

Wilks' lambda F Df Sig.

WDS 0.819 39.538 2 0.000

LDS 0.763 55.506 2 0.000
WMS 0.966 6.210 2 0.002
LMS 0.920 15.448 2 0.000
WGS 0.874 25.779 2 0.000
LGS 0.871 26.375 2 0.000
Fig 3 Scatterplot of the results of the discriminant analysis of the
WAS 0.969 5.736 2 0.004 three generalized groups established by the cluster dendrogram.
LAS 0.879 24.581 2 0.000 Triangles represent morphotype 1 characterized by mite populations
from Río Negro, Santa Fé, Santa Cruz, Misiones, and San Luis
WDS width of dorsal shield, LDS length of dorsal shield, WGS width provinces. Circles represent morphotype 2 characterized by mite
of genital shield, LGS length of genital shield, WMS width of populations from Buenos Aires, Neuquén, Corrientes, and Entre Ríos
metapodal shield, LMS length of metapodal shield, WAS width of provinces. Crosses represent morphotype 3 characterized by mite
anal shield, LAS length of anal shield populations from Santiago de Estero, Tucumán, and Jujuy provinces
Parasitol Res (2009) 105:1629–1636
also not considered as probable factors regulating the
morphological differences among parasite populations.
Respect to body size of parasites reported in this
research, WDS and LDS means corresponding to morpho-
types 2 and 3, were similar to values observed by Anderson
and Trueman (2000) for V. destructor (WDS=1.708 mm±
0.041; LDS = 1.167 mm ± 0.026), while morphotype 1
presented higher values of these variables with respect to
the reported by the authors mentioned above.
According to the morphotypes found, different hypoth-
esis could be postulated to be tested in future investigations:
(a) the morphotypes are the result of morphometric
correlations between coexisting populations of V. destructor
and A. mellifera. Relationships of this nature have been
observed for others host–parasite systems (Poulin 1998;
Poulin et al. 2003). Recently, parasitological studies have
interpreted these parasite–host interactions in terms of
energy flux, showing that parasite biomass is controlled
by food resources offered by the host, which metabolic rate
and the size of the host (George-Nascimento et al. 2004;
Poulin and George-Nascimento 2007). Within A. mellifera
bee populations, morphological differences have been
found (Buco et al. 1987; Radloff and Hepburn 2000;
Kandemir et al. 2000). The bee morphotypes could offer
different energetic resources to maintain the parasite
morphotypes. In this sense, future researches would analyze
if a particular morphotype of V. destructor is related to a
particular morphotype of A. mellifera. (b) The morphotypes
observed for V. destructor are the expression of the
interaction between genotypes of both, parasite and host
(“Extended Phenotype Theory”, Lambrechts et al. 2005).
The potential importance of this concept was demonstrated
by a hypothetical coevolutive model from parasite–host
systems, which consider that epidemiologic features are
controlled by the interaction between both components of
the system (Restif and Koella 2003). Several studies have
documented the existence of different genotypes in A.
mellifera in Argentina, reflecting different levels of Afri-
canized bees in the country (De Santis and Cornejo 1968;
Dietz et al. 1985). In this way, the morphotypes observed in
this study could represent the expression of the interaction
between different genotypes of the parasite and its host. (c)
The morphotypes found are the result of selective pressures
produced by different intensities of acaricide use, which are
characteristic of each geographic location. Clarke and
McKenzie (1987) reported that insecticide-resistant pheno-
types of the Australian sheep blowfly, Lucilia cuprina,
showed greater fluctuating asymmetry (random differences
between left and right sides of a normally bilaterally
symmetric organism) than susceptible phenotypes. In V.
destructor mites, relationships between mite survival rate
and body size were observed under exposition to organic
acaricides in laboratory: the parasites of bigger sizes
1635
showed higher survival rates than the ones of smaller sizes
(Maggi pers. Obs.). Consequently, it is possible that
geographic locations where the sampling was conducted,
corresponded to different intensities of acaricide use, which
selected the morphotypes encountered in V. destructor.
Finally, the morphotypes registered in this study would be
the result of local and current conditions (bee body-size
variability, selective pressures of acaricides, interactions
between genotypes of host and parasite, or another hypothesis
not discussed here) through small changes in body size,
resulting in the life-history strategy best suited to those
conditions. This phenotypic plasticity should be an immediate
response to environmental changes and not result in changes
in genotypic frequencies in the population. According to
Stearns (1992) a true evolutionary response, involving a shift
in gene frequencies in the population, can occur only when
environmental changes persist in time. The parasite system
V. destructor/A. mellifera is a “young system” because
V. destructor colonized A. mellifera host in 1960 when
beekeepers move colonies of A. mellifera from Europe to
Asia and the parasite could switch its host (Anderson and
Trueman 2000). In this sense, morphotypes observed in this
research could represent an immediate response to environ-
mental changes.
According to the results presented in this research, the
first step to characterize mite populations of Argentina was
assessed. These kinds of studies in host–parasite systems
carried out in different geographical locations allow
extending the knowledge of parasites in epidemic-ecologic
terms, helping to reformulate sanitary programs.
Acknowledgments This study was supported by Creo (Oportunidades
para Conservación, Investigación y Educación) to M. M. The authors
would like to thank Msc. Di Rienzo for the assistance in the statistical
analysis and to Nancy García, Manuel Darío Mathot, Juan Mange, Pablo
Pérez, Natalia Messina, Fernando Mariani, Pedro Bailac, José Ernesto
Nuñez Camelino, Cristiansen, Jose Luis Carlachiani, Freddy Alemán,
Santiago Plischuk, NOA program Schools (PROAPI), Apicultura Sin
Fronteras, and Campo & Abejas for their help in the sampling. The
experiments comply with the current laws of the country in which they
were performed.
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