Hereditas 143: 123 129 (2006)
Unusual karyotype diversity in the European spiders of the genus Atypus
(Araneae: Atypidae)
MILAN ŘEZÁČ1, JIŘÍ KRÁL1, JANA MUSILOVÁ1 and STANO PEKÁR2
1
Laboratory of Arachnid Cytogenetics, Department of Genetics and Microbiology, Faculty of Sciences, Charles
University in Prague, Prague, Czech Republic
2
Institute of Botany and Zoology, Faculty of Science, Masaryk University, Brno, Czech Republic
Řezáč, M., Král, J., Musilová, J. and Pekár, S. 2006. Unusual karyotype diversity in the European spiders of
the genus Atypus (Araneae: Atypidae). * Hereditas 143: 123 129. Lund, Sweden. eISSN 1601-5223. Received
February 28, 2006. Accepted May 11, 2006
Compared with araneomorph spiders, karyotypes of the spider infraorder Mygalomorphae are nearly unknown. In this
study we investigated karyotypes of European species of the genus Atypus (Atypidae). The male karyotype of A. muralis and
A. piceus comprises 41 chromosomes, whereas female complements contain 42 chromosomes. On the other hand, both sexes
of A. affinis possess 14 chromosomes only. It is the lowest diploid number found in mygalomorph spiders so far.
Furthermore, obtained data suggest X0 sex chromosome system in A. piceus, A. muralis and neo-XY system in A. affinis.
Karyotypes of all three Atypus species are composed of biarmed chromosomes only. Thus they differ significantly from the
karyotype of A. karschi , the only other species of this genus studied so far. Its karyotype was reported to be composed of
acrocentric chromosomes and possesses X1X20 sex chromosome system. All this shows that unlike in most genera of
araneomorph spiders, mygalomorphs of the genus Atypus exhibit unusual diversity in the number, morphology of
chromosomes, and the sex chromosome system. Considering high number of chromosomes being plesiomorphic character in
spiders, then karyotypes of A. muralis and A. piceus represent ancestral situation and that of A. affinis being derived by
multiple fusions. Karyotype differences in Atypus correspond with morphological differences, namely the number of
segments of the posterior lateral spinnerets. Thus in contrast to published hypothesis, the 3-segmented posterior lateral
spinnerets of A. affinis should present a derived state.
Milan Řezáč, Laboratory of Arachnid Cytogenetics, Dept of Genetics and Microbiology, Faculty of Sciences, Charles University,
Viničná 5, CZ-128 44 Prague 2, Czech Republic. E-mail: [email protected]
Spiders (Araneae) appear to be the best studied order
of the class Arachnida concerning cytogenetics. Untill
now, karyotypes of approximately 500 species of
spiders have been described (GOWAN 1985; KRÁL
1994; ARAÚJO et al. 2005). Nevertheless, taking into
account the enormous diversity of the order (roughly
39 000 species; PLATNICK 2005), our knowledge about
spider karyology is still unsatisfactory.
The order Araneae is divided into three phylogenetic lineages, namely Mesothelae, Mygalomorphae
and Araneomorphae, the last one being phylogenetically most derived (CODDINGTON and LEVI 1991).
Mesothelae represents a relict group with hardly 100
species in contrast to Araneomorphae that includes
roughly 36 000 species classified into 94 families
(PLATNICK 2005). At last, mygalomorphs are represented by fifteen families comprising roughly 2500
species (PLATNICK 2005). The sum of karyological
data about the three major spider lineages conforms
with their diversity. The vast majority of karyological
data concern araneomorph spiders. Considering mygalomorphs, only fragmentary data on karyotypes of
14 representatives belonging to five families have been
published so far. This is due to limited access to
mygalomorphs that are almost restricted to tropics
and subtropics. Very few species of mygalomorphs
occur in the temperate zone, for example, purse web
spiders (Atypidae) (PLATNICK 2005).
Spiders exhibit great diversity in diploid chromosome numbers: 2n of males ranges between 7 to 94
(SUZUKI 1954). The karyotype of the majority of the
species studied is composed of acrocentric chromosomes only (TUGMON et al. 1990). Biarmed chromosomes prevail only in karyotypes of some groups,
namely the family Dictynidae (KRÁL 1995) and in
the haplogyne lineage of araneomorph spiders
(COKENDOLPHER 1989; RODRÍGUEZ GIL et al. 2002,
KRÁL et al. 2004). Primitive araneomorph spiders of
the families Dysderidae and Segestriidae have karyotypes composed of holokinetic chromosomes (DÍAZ
and SÁEZ 1966; RODRÍGUEZ GIL et al. 2002).
Spiders are also unique by predomination of multiple sex chromosome determination ß/X1X2/ à/
X1X1X2X2 (BENAVENTE et al. 1982). This system,
often assigned as X1X20 (where 0 denotes the absence
of the chromosome Y) was found in 77% of the spider
species studied so far (ARAÚJO et al. 2005). Such
determination is probably ancestral in spiders as it was
found also in the most plesiomorphic recent spider
taxon, the Mesothelae (SUZUKI 1954). In some spider
124
M. Řezáč et al.
groups, the following systems X1X2X30, X1X2X3X40
and X0 were derived from the original sex chromosome system, X1X20. The first two systems evolved
probably by nondisjunction of the X chromosome
(BRUM-ZORRILLA and POSTIGLIONI 1981), the last
one was derived mostly by centric or tandem fusion of
the original sex chromosomes X1 and X2 (KRÁL
1994). Sex chromosome systems containing neo-sex
chromosomes are rare among spiders. Untill now, they
have been described only in two genera of North
American jumping spiders, Habronattus and Evarcha
(MADDISON 1982), in an Australian social huntsman
spider, Delena cancerides (ROWELL 1985), and in an
European funnel-web spider, Tegenaria ferruginea
(KRÁL 2001).
In order to fill gap in the karyology of mygalomorph spiders, we have focused on the European
representatives of the family Atypidae. Studies of this
family is important for the understanding of karyotype evolution in spiders, since this family is supposed
to be one of the most primitive groups of mygalomorphs (CODDINGTON and LEVI 1991). Moreover,
obtained results might also reveal pathways of evolution within European atypids. At present, three genera
are distinguished within the family Atypidae;
Calommata (southeast Asia and Africa), Sphodros
(North America), and Atypus (Europe and Asia). The
majority of species of the genus Atypus occur in
southeast Asia. However, some species occur also in
the Palearctic region. Three species of this genus are
known from Europe, A. piceus (SULZER, 1776), A.
muralis (BERTKAU, 1890) and A. affinis (EICHWALD,
1830) (KRAUS and BAUR 1974). When LATREILLE
(1804) used the name Atypus for purse-web spiders, he
meant to express their peculiarity among other
European spiders; they are the only central European
mygalomorphs. Little attention has been paid to these
spiders so far, particularly due to their rareness and
anachoretic lifestyle. They occur in sunny xerothermic
slopes, where they live in relatively deep burrows
(BROEN and MORITZ 1964; HIEBSCH and KRAUSE
1976). Their presence is mainly revealed by spotting
the purse-webs that serve them for peculiar prey
capture (BRISTOWE 1958).
Hereditas 143 (2006)
stage turned out to be the most appropriate ontogenetic stages for the analysis. Gonads of pre-subadult
instars contained numerous mitoses. Testes of subadult males of A. piceus collected at the end of April
gave various stages of meiotic division. In contrast to
many other spiders, young adult males appeared to be
unsuitable for karyotype analysis as their testes
contained hardly any meiotic cells. We suggest that
meiosis is completed already before adult stage due to
short lifetime of adult males. All karyotyped specimens are deposited in the institutional collection of
the first author who also identified the species. Species
nomenclature follows PLATNICK (2005).
Chromosome preparations were obtained by a
modification of the spreading technique described by
TRAUT (1976). The gonads were dissected from the
abdomen in a hypotonic solution (0.075M KCl) and
moved to a fresh hypotonic solution so that the tissue
was hypotonized for 10 min in total. This was followed
by 1012 min fixation in freshly prepared Carnoy
fixative (ethanol: chloroform: glacial acetic acid 6:3:1)
and 20 min fixation in a new Carnoy fixative. Afterwards, the tissue was placed in a drop of 60% acetic
acid on a clean slide and quickly shredded as finely as
possible with a pair of fine tungsten needles. In the
end, the slide was quickly moved onto a warm
histological plate (surface temperature of 408C) and
the drop of dispersed tissue was allowed to evaporate
while moving it constantly using a fine tungsten
needle. Slides were air-dried at room temperature
overnight, and stained with 5% Giemsa solution in
Sörensen phosphate buffer (pH /6.8) for 2530 min.
Preparations were inspected using Jeneval microscope (Carl Zeiss Jena) by means of an immersion
lens. The best figures were photographed. To obtain
data on chromosome morphology, ten gonial metaphases were evaluated. Relative chromosome lengths
(RCL, Table 1) were calculated as a percentage of the
total chromosome length in the diploid set, including
the sex chromosome(s). Calculation of centromeric
index (CI) and the chromosome classification system
follows LEVAN et al. (1964).
RESULTS
Atypus piceus
MATERIAL AND METHODS
All specimens for the karyological analyses were
collected in Prague, or in its surroundings, specifically
A. muralis in Divoká Šárka valley, A. affinis in
Břežanské údolı́ valley and A. piceus in Karlštejn.
The spiders were dug out of their burrows. In each
species, two or three individuals of both sexes were
used for the analysis. Large nymphs and subadult
The male karyotype comprised of 41 (Fig. 1a), and the
female karyotype of 42 relatively small chromosomes.
All autosome pairs were metacentric except for a
submetacentric pair no. 2. This chromosome pair
beared a subterminal secondary constriction on its
long arm. Autosomes gradually decreased in size. In
the mitotic metaphase, the autosome lengths ranged
from 8.0 mm (RCL /3.68%) to 2.9 mm (RCL /1.40%).
Karyotype diversity in European Atypus species
Hereditas 143 (2006)
125
Table 1. European Atypus species. Chromosome relative lengths (RCL) and centromeric indexes (CI) based on
mitotic metaphases from gonads.
Pair no.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
X
Y
Atypus piceus
Atypus muralis
male
male
Atypus affinis
male
female
RCL
CI
RCL
CI
RCL
CI
RCL
CI
3.68
3.47
3.39
3.11
2.77
2.65
2.58
2.50
2.39
2.35
2.29
2.25
2.07
1.99
1.97
1.89
1.80
1.71
1.59
1.40
4.32
1.28
2.52
1.30
1.05
1.21
1.06
1.35
1.09
1.25
1.14
1.30
1.09
1.10
1.50
1.08
1.08
1.38
1.14
1.10
1.00
1.11
4.04
3.73
3.47
3.24
3.09
2.84
2.57
2.43
2.41
2.41
2.08
2.03
2.01
1.89
1.84
1.82
1.75
1.66
1.58
1.35
3.54
1.10
1.37
1.06
1.23
1.28
1.05
1.24
1.54
1.32
1.21
1.14
3.34
1.08
1.23
1.26
1.45
1.11
1.18
1.12
2.25
1.21
12.22
10.16
9.02
7.80
1.90
1.47
1.16
1.67
1.12
2.17
1.18
1.45
12.32
11.24
7.33
7.15
1.38
1.27
1.23
2.26
1.35
2.33
1.17
1.43
9.21
5.65
1.05
1.40
9.31
1.15
The X chromosome was metacentric. This chromosome was the longest in the karyotype (Table 1). In
males, the X chromosome was placed on the periphery
of the nucleus during prophase of the first meiotic
division. Furthermore, the X chromosome exhibited
positive heteropycnosis until the end of pachytene
(Fig. 2). Inspection of male meiotic division and
comparison of mitotic metaphases of both sexes
indicated the X0 sex chromosome system.
Atypus muralis
The karyotype of A. muralis was similar to that of A.
piceus. It also comprised of 41 relatively small
chromosomes in males (Fig. 1b), and of 42 chromosomes in females. All autosome pairs were metacentric
except for a submetacentric pair no. 20 and subtelocentric pair no. 12. Autosomes gradually decreased in
size. In the mitotic metaphase, the autosome lengths
ranged from 9.4 mm (RCL /4.04%) to 3.5 mm (RCL /
1.35%). Autosome pair no. 9 probably beared a
secondary constriction in the middle of its short
arm. The metacentric X chromosome was relatively
large (Table 1). Comparison of mitotic metaphases of
both sexes confirmed the X0 sex chromosome system.
Atypus affinis
Karyotypes of both sexes of A. affinis possessed only
14 chromosomes (Fig. 1c, 1d). The karyotype of this
species was characterized by a remarkable asymmetry.
The first four autosome pairs were significantly larger
than remaining two pairs. The largest autosome pair
was eight times longer (mean 21.4 mm) than the
smallest one (mean 2.7 mm). The two pairs of large
(no. 1 and 3) and tiny autosome pairs (no. 5 and 6)
were metacentric. Large pairs no. 2 and 4 were
submetacentric. Prominent secondary constriction
was placed subterminally in a long arm of the
submetacentric autosome pair no. 2. The length of
the constriction and adjacent satellite was variable.
Most of the studied specimens were heterozygous in
these markers. Two large metacentric chromosomes
were apparently odd in the male karyotype, so we
considered them to be sex chromosomes. Based on
comparison of male and female karyotype, they are X
and Y chromosome. In female karyotype, one of these
chromosomes was paired, whereas the second one was
missing. The arms of the X chromosome were
approximately of the identical length, while the
centromeric index of Y chromosome was 1.4 (Table 1).
DISCUSSION
Karyotypes of all European species of the mygalomorph genus Atypus are predominated by metacentric
chromosomes. The male karyotype of A. piceus and
that of A. muralis comprises of 41 and the female
126
M. Řezáč et al.
Hereditas 143 (2006)
Fig. 1a-d. Karyotypes of the European representatives of the genus Atypus. (a) A. piceus
(spermatogonial metaphase), (b) A. muralis (spermatogonial metaphase), (c) A. affinis
(spermatogonial metaphase), (d) A. affinis (oogonial metaphase). Scale bar/10mm.
karyotype comprises of 42 chromosomes. Karyotypes
of these species differ only in morphology of several
autosome pairs. In contrast to this, karyotype of A.
affinis consists of 14 chromosomes only. Male diploid
Fig. 2. A. piceus, male pachytene. Note positively heteropycnotic X chromosome on the periphery of the nucleus.
Arms of the X chromosome are closely aligned (arrow).
Scale bar /10mm.
chromosome numbers in mygalomorph spiders range
from 42 to 81 (Table 2). Therefore, the diploid
chromosome number found in A. affinis is the lowest
number found in mygalomorphs so far. In araneomorph spiders the lowest diploid number is even lower
(2n /7) and was found in the species of the family
Segestriidae, Ariadna lateralis, that exhibits holokinetic chromosomes (SUZUKI 1954).
Our analysis revealed the sex chromosome system
X0 in A. piceus and A. muralis. HACKMAN (1948)
supposed such system already in the theraphosid
mygalomorph Aphonopelma hentzi (Table 2). His
conclusion was based merely on Painter’s original
drawing (1914). However, PAINTER (1914) himself
mentioned two sex chromosomes in the male karyotype of the species. Moreover, his verbal description of
sex chromosome behaviour during meiosis indicated
presence of two X chromosomes. Unfortunately, the
karyotype of A. hentzi has never been revised.
Beside diploid number of chromosomes, A. affinis
differs remarkably from two other European Atypus
species in the sex chromosome system. Both sexes of
A. affinis possess 14 chromosomes, thus, the sex
chromosome system can be neither the common
X1X20 type nor X0. According to our data, the sex
Karyotype diversity in European Atypus species
Hereditas 143 (2006)
127
Table 2. List of karyotyped species of mygalomorph spiders. Abbreviations: m metacentric; smt submetacentric;
st subtelocentric; a acrocentric chromosomes, ? unknown, uncertain. Species nomenclature follows PLATNICK
(2005).
Family/species
Antrodiaetidae
Antrodiaetus unicolor
Atypidae
Atypus affinis
Atypus karschi
Atypus muralis
Atypus piceus
Cyrtaucheniidae
Cyclocosmia torreya
Myrmekiaphila torreya
Dipluridae
Ischnothele indicola
Theraphosidae
Acanthoscurria gomesiana
Aphonopelma hentzi
Brachypelma albopilosa
Vitalius dubius
Vitalius roseus
Vitalius sorocabae
Vitalius wacketi
Vitalius sp. 1
Vitalius sp. 2
Number and morphology
of chromosomes
Sex chromosome
system
Reference
ß/ 46
?
HETZLER (1979)
ß/ 14, à/: 14; m, smt
ß/ 44?; a
ß/ 41, à/: 42; m, smt, st
ß/ 41, à/: 42; m, smt
neoXneoY
X1X20
X0
X0
this study
SUZUKI (1954)
this study
this study
ß/ 42
ß/ 80?
?
?
HETZLER (1979)
HETZLER (1979)
ß/ 42
X1X20
SRIVASTAVA and SHUKLA (1986)
ß/ 43 46; m, smt, st, a
ß/ 44
ß/ 74
ß/ 74 81; m, smt, st, a
? ß/ 48; st, a
? ß/ 48; st, a
? ß/ 48; st, a
à/ 48; m, smt, st, a
juveniles 47, 48; m, smt, st, a
?
X1X20 or X0
?
?
?
?
?
?
?
OLIVEIRA (1998)
PAINTER (1914); HACKMAN (1948)
VÍTKOVÁ et al. (2005)
OLIVEIRA (1998)
LUCAS et al. (1993)
LUCAS et al. (1993)
LUCAS et al. (1993)
OLIVEIRA (1998)
OLIVEIRA (1998)
chromosome determination of A. affinis is probably
XY; this system has not been found in spiders so far.
Such system has probably evolved by rearrangements
of the sex chromosomes of the original X0 system,
found in other European Atypus species, and autosomes. Thus it represents neo-XY system.
Only in a few of mygalomorph species, the morphology of chromosomes and the sex chromosome
system has been described (Table 2). As concerns the
genus Atypus, karyotype of a single species only, A.
karschi from eastern Asia, has been studied so far.
Interestingly, karyotypes of all European species of the
genus Atypus differ markedly from the karyotype of
A. karschi both in the morphology of chromosomes
and the sex chromosome system. The karyotype of
this species is supposed to consist of 44? relatively
small acrocentric chromosomes including the sex
chromosome system X1X20 (SUZUKI 1954). We suggest that the sole metacentric X chromosome of
A. piceus and A. muralis have originated by Robertsonian translocation betwen original acrocentric sex
chromosomes, X1 and X2. The frequent changes in the
sex chromosome constitution suggest that the sex
chromosomes has been involved in the speciation
process within the genus Atypus. Rearrangements of
sex chromosomes may influence considerably the
balance between male and female sex factors breaking
realization of sexual phenotype. In this way, extensive
rearrangements of sex chromosomes can play an
important role in postzygotic reproductive isolation
of species (CHARLESWORTH 1987, KING 1993).
Our data show that karyotypes of mygalomorphs
are very variable in both number and morphology of
chromosomes. Like in haplogyne lineage of araneomorph spiders (KRÁL et al. 2004), mygalomorph
karyotypes are usually dominated by biarmed chromosomes. Furthermore, sex chromosome systems of
both groups are considerably diversified, often including an Y chromosome. More interestingly, in the genus
Atypus, diversity in the number and morphology of
chromosomes as well as in the sex chromosome
determination occurs even within a single genus. In
contrast to this, karyotypes of entelegyne lineage of
araneomorph spiders are considerably conservative
being composed usually entirely by acrocentric chromosomes. In majority of entelegyne spiders the system
X1X20 is present (KRÁL 1994).
A high number of chromosomes probably represents
a plesiomorphic state in spiders (SUZUKI 1954).
Therefore, karyotypes of A. karschi , A. muralis and
A. piceus may be ancestral to the karyotype of A.
affinis. This hypothesis is supported by a comparison
with the karyotype of the representative of the family
Antrodiaetidae, which is considered to be the sister
128
M. Řezáč et al.
family of Atypidae (CODDINGTON and LEVI 1991).
Diploid chromosome number of Antrodiaetus unicolor
is similar to that of Atypus species with higher number
of chromosomes (HETZLER 1979; Table 2). The
reduction of chromosome number in A. affinis is
probably result of multiple fusions. Two pairs of small
autosomes in A. affinis can represent relict pairs of the
original karyotype which remained intact.
The considerable karyotype differences in the European Atypus spiders correspond well with morphological differences, particularly with the number of
segments of the posterior lateral spinnerets (PLS).
The PLS of species with high number of chromosomes
are composed of four segments (A. muralis, A. karschi )
or with the fourth one hinted (A. piceus ). In contrast
to this, the PLS of A. affinis are composed of
three segments only. The 3-segmented PLS were
considered to be plesiomorphic in the genus Atypus
(SCHWENDINGER 1990). On the contrary to this, our
data indicate that karyotype of A. affinis is derived and
so the species. Therefore, we propose that 3-segmented
PLS of A. affinis represent an apomorphy; they
established by fusion of the last two segments.
Drastic reduction of chromosome number by multiple fusions in A. affinis seem to be analogous to that
discovered in some other groups of closely related
animal species, for example, in the deer genus Muntiacus (LIN et al. 1991; WANG and LAN 2000). European
Atypus species are, therefore, promising models for
analysis of speciation processes related to multiple
fusions.
Acknowledgements Collection of the spiders in protected
areas was confered by the edict of the Ministry of the
Environment of the Czech Republic No. 27068/03 /5536/03
(locality Karlštejn) and by the edict of the Municipality of
the city of Prague MHMP 221/2004/OŽP V 7/R 4/Pra
(localities Divoká Šárka valley and Břežanské údolı́ valley).
MŘ was supported by project no. 233/2004/B-BIO provided
by the Charles University, Prague. JK was supported by the
project no. 0021620828 and SP by the project no.
0021622416 provided by the Ministry of Education, Youth
and Sports of the Czech Republic.
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