Page 1
RESEARCH PAPER
Trends in chromosome evolution in the genus HypostomusLacepede, 1803 (Osteichthyes, Loricariidae): a newperspective about the correlation between diploid numberand chromosomes types
Vanessa Bueno • Claudio Henrique Zawadzki •
Vladimir Pavan Margarido
Received: 17 February 2011 / Accepted: 6 April 2011 / Published online: 19 April 2011
� Springer Science+Business Media B.V. 2011
Abstract Phylogenetic relationships and identifica-
tion of species of the genus Hypostomus is still
unclear. Considering this, cytogenetics may prove
itself as an important tool in understanding the
systematic of this genus. Reviews in Hypostomus
indicate that the diploid number ranges from 54 to 84
chromosomes, and the increase in diploid number has
been associated to higher percentages of subtelocen-
tric and acrocentric chromosomes. Although there is a
high number of species in the genus, there are
relatively few papers concerning Hypostomus cyto-
genetics, and most of the data is published as grey
literature. With the aim to understand the chromo-
somal evolution in the genus (correlation between
diploid number x chromosomes types), H. ancistro-
ides and H. topavae from the Piquiri River, Upper
Parana River basin, were cytogenetically analyzed,
and the diploid number observed was 68 and 80
chromosomes, respectively. Additional data on the
diploid number and chromosome formulae was com-
piled from papers (27 analyses) and abstracts from
grey literature (77 analyses). Our analysis shows no
correlation between chromosome numbers and per-
centages of subtelocentric and acrocentric chromo-
somes for most of the species, since there is
considerable variation between these percentages
even between species with the same diploid number,
indicating that the proportion of chromosome types is
not always associated to diploid numbers.
Keywords Hypostomus ancistroides � Hypostomus
topavae � Karyotypic diversity � Fish cytogenetics
Introduction
The family Loricariidae is one of the most species rich
taxon among the Siluriformes, with about 700 species
grouped in seven subfamilies: Delturinae, Hypoptom-
atinae, Hypostominae, Lithogeneinae, Loricariinae,
Neoplecostominae and Otothyrinae (Reis et al. 2006;
Ferraris 2007; Chiachio et al. 2008). The phylogenetic
relationships among the Hypostominae are not very
clear, especially in the genus Hypostomus. The most
recent study regarding the relationships within Hypos-
tominae was published by Armbruster (2004). The
author presented a detailed analysis about the relation-
ships within Loricariidae, and synonymized the former
genera contained in the tribe Hypostomini (Aphano-
torulus, Cochliodon, Isorineloricaria, Squaliforma
and Watawata) as Hypostomus. Montoya-Burgos
V. Bueno � V. P. Margarido (&)
Centro de Ciencias Biologicas e da Saude, Universidade
Estadual do Oeste do Parana, Rua Universitaria 2069,
Cascavel, Parana State 85819-110, Brazil
e-mail: [email protected]
C. H. Zawadzki
Departamento de Biologia/Nupelia, Universidade
Estadual de Maringa, Av. Colombo, 5790, Maringa,
Parana State 87020-900, Brazil
123
Rev Fish Biol Fisheries (2012) 22:241–250
DOI 10.1007/s11160-011-9215-9
Page 2
(2003) through D-loop sequences analysis found
Aphanotorulus, Isorineloricaria and Hypostomus
emarginatus out of the main clade of Hypostomus.
The author recognized the genera Aphanotolurus and
Isorineloricaria, and considered Hypostomus as a
paraphyletic assemblage due to not include H. emar-
ginatus. Despite these studies, a broader revision of the
genus is still needed.
Accumulated cytogenetic data can be used for the
establishment of evolutionary trends, identification of
new species and distinction of cryptic species (Artoni
et al. 2000; Bellafronte et al. 2010; Blanco et al.
2010; Perazzo et al. 2010). Considering the difficul-
ties in identifying several of the Hypostomus species
and its unclear phylogeny, cytogenetics may prove
itself as an important tool in understanding the
systematics of the genus.
Reviews on Hypostomus cytogenetics data were
presented by Artoni and Bertollo (1999; 2001),
Kavalco et al. (2005) and Alves et al. (2006). The
diploid numbers in Hypostomus range from 54 chro-
mossomes in Hypostomus plecostomus (Muramoto
et al. 1968) to 84 chromosomes in Hypostomus sp. 2
(Cereali et al. 2008). Artoni and Bertollo (2001)
consider the diploid number of 54 chromosomes basal
for Hypostominae, using Trichomycteridae as out-
group. The authors observed a higher proportion of
subtelocentric and acrocentric chromosomes in spe-
cies with higher diploid numbers, suggesting that the
chromosome evolution on Hypostomus occurred
through centric fissions. Also, a considerable structural
variation among species sharing the same diploid
number was observed, indicating karyotypic differen-
tiation through structural changes like Robertsonian
rearrangements and pericentric inversions (Kavalco
et al. 2005).
The majority of the data concerning Hypostomus
cytogenetics seems to be published as grey literature,
probably because of the complications related to the
identification of the specimens, the substantial num-
ber of new species and the high amount of cytoge-
netic variation between populations. The publication
of data on meetings and the large number of species
that are unidentified or putative new to science
hampers the compilation and analysis of these data,
preventing a wider analysis of the genus. Thus, a
broader review of the cytogenetic data on Hyposto-
mus, covering also the abstracts published on the
main meetings of fish cytogenetics, would allow a
more accurate view on the results obtained for this
fish group.
Materials and methods
Two species of the genus Hypostomus (Fig. 1) from
the Piquiri River were cytogenetically analyzed:
H. ancistroides (4 males and 11 females) from
Formosa do Oeste (Parana State, Brazil) and
H. topavae (9 males and 6 females), from Nova
Laranjeiras (Parana State, Brazil). The individuals
were sacrificed with overdoses of clove oil (Griffiths
2000). Voucher specimens were deposited in the
Colecao Ictiologica do Nucleo de Pesquisas em
Limnologia, Ictiologia e Aquicultura—Nupelia—
Universidade Estadual de Maringa, Brazil (NUP
3902—H. ancistroides and NUP 11430—H. topa-
vae). Metaphasic cells were obtained from the kidney
(Bertollo et al. 1978; Foresti et al. 1993). Chromo-
somes were classified in metacentric (m), submeta-
centric (sm), subtelocentric (st) and acrocentric (a),
according to Levan et al. (1964).
The review of the available cytogenetic data for
Hypostomus included data from published papers and
from abstracts presented on the Brazilian Sympo-
siums on Fish Cytogenetics and Genetics, from its
first edition in 1986 to the latest edition in 2009.
Inconsistent and repeated data were not included, and
analyses from different populations of the same
species were taken into account when this data was
available. The linear regression and a one-way
analysis of variance were carried out using the
software Statistica 7, considering the data in Tables 1
and 2 that had information about the chromosome
formulae. The analysis of variance was performed
considering the diploid number as the grouping
factor, and each diploid number was considered as
an individual group. The analyses were performed (1)
considering only the published data and (2) consid-
ering the published data together with the grey
literature.
Results
Hypostomus ancistroides presented 2n = 68 chromo-
somes (14 m ? 14 sm ? 8 st ? 32 a) for both sexes
242 Rev Fish Biol Fisheries (2012) 22:241–250
123
Page 3
(Fig. 2a). Hypostomus topavae presented 2n = 80
chromosomes (14 m ? 10 sm ? 26 st ? 30 a) for both
sexes (Fig. 2b). Tables 1 and 2 were made consider-
ing 27 analyses compiled from manuscripts and 77
analyses from grey literature, respectively. The linear
regression for chromosome number and percentage of
subtelocentric and acrocentric chromosomes showed
correlation values of r = 0.6048, P = 0.0008, con-
sidering only the published data (Fig. 3), and
r = 0.3215, P = 0.002 considering both the pub-
lished data and the grey literature (Fig. 4). The
analysis of variance showed that the difference
between the percentage of subtelocentric and acro-
centric chromosomes is not significant between
populations with different diploid numbers, with
P-values equal to 0.05343, considering only the
published data, and 0.16714 considering both classes
of data.
Discussion
Considering the high number of species in Hypostomus,
more than 120 according to Zawadzki et al. (2010),
there is relatively few cytogenetic data available in the
literature (Table 1), and most of the data available for
the genus is presented in grey literature (Table 2). The
review presented here showed that the diploid number
for Hypostomus ranges from 54 to 84 chromosomes.
Some previous reviews consider 2n = 52 chromo-
somes in Hypostomus emarginatus the smallest diploid
number (Artoni and Bertollo 2001; Alves et al. 2006).
Although the genus Squaliforma has been considered a
synonym for Hypostomus by Armbruster (2004), his
conclusions are not consensual, and there are a number
of publications that consider Squaliforma a valid genus
(Weber 2003; Nelson 2006; Ferraris 2007; Froese and
Pauly 2010). The cytogenetic data presented herein
support the latter conclusion, once S. emarginata has a
reduced diploid number compared to all other Hypo-
stomus species.
Artoni and Bertollo (1996; 2001) observed that
species with higher diploid numbers also showed a
higher proportion of subtelocentric and acrocentric
chromosomes, when compared to species with diploid
numbers closer to 54 chromosomes. They analyzed
the linear regression between the diploid numbers and
percentage of subtelocentric and acrocentric chromo-
somes, and found significant correlation (r = 0.8122,
P \ 0.0001), proposing that centric fissions may have
been an important part of the chromosome evolution
of the genus. Since their studies, there have been no
Fig. 1 Lateral view of a Hypostomus ancistroides (255 mm SL) and b Hypostomus topavae (125 mm SL)
Rev Fish Biol Fisheries (2012) 22:241–250 243
123
Page 4
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244 Rev Fish Biol Fisheries (2012) 22:241–250
123
Page 5
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58
72
.50
Pir
acic
aba
riv
erS
PR
ub
ert
etal
.(2
00
9a)
9
Rev Fish Biol Fisheries (2012) 22:241–250 245
123
Page 6
Ta
ble
2co
nti
nu
ed
Sp
ecie
s2
nC
hro
mo
som
efo
rmu
lae
st-a
%L
oca
lity
UF
Ref
eren
ces
Mee
tin
gs
msm
sta
Hyp
ost
om
us
ma
rga
riti
fer
72
10
16
14
32
63
.89
Piq
uir
iri
ver
PR
Lo
rsch
eid
eret
al.
(20
09
9
Hyp
ost
om
us
ma
rga
riti
fer
72
32
40
55
.56
Par
do
riv
er–
Pen
tead
oet
al.
(20
09
)9
Hyp
ost
om
us
mye
rsi
74
12
14
18
30
64
.86
Igu
acu
riv
erP
RC
asal
eet
al.
(20
02
)5
Hyp
ost
om
us
mye
rsi
74
12
14
10
38
64
.86
Igu
acu
riv
erP
RL
ui
and
Mar
gar
ido
(20
06
)7
Hyp
ost
om
us
nig
rom
acu
latu
s7
66
20
50
65
.79
Tre
sB
oca
sst
ream
PR
Ru
ber
tan
dG
iuli
ano
-Cae
tan
o(2
00
6a)
7
Hyp
ost
om
us
nig
rom
acu
latu
s7
68
20
48
63
.16
Em
asw
ater
fall
SP
Ru
ber
tan
dG
iuli
ano
-Cae
tan
o(2
00
6a)
7
Hyp
ost
om
us
nig
rom
acu
latu
s7
61
22
23
01
25
5.2
6P
assa
-cin
cori
ver
SP
Tra
ldi
etal
.(2
00
9b
)9
Hyp
ost
om
us
pa
uli
nu
s7
66
16
54
71
.05
Pir
acic
aba
riv
erS
PR
ub
ert
etal
.(2
00
9a)
9
Hyp
ost
om
us
ple
cost
om
us
68
––
––
––
–P
ileg
gi
etal
.(1
98
6)
1
Hyp
ost
om
us
reg
an
i7
21
41
28
38
63
.89
––
Lar
a-K
amei
and
Juli
o-J
un
ior
(20
00
)4
Hyp
ost
om
us
reg
an
i7
21
82
01
22
24
7.2
2P
ard
ori
ver
–P
ente
ado
etal
.(2
00
9)
9
Hyp
ost
om
us
cf.
reg
an
i7
21
21
82
61
65
8.3
3A
raq
ua
stre
amS
PIs
hid
aet
al.
(20
02
)5
Hyp
ost
om
us
stri
ga
tice
ps
72
10
16
46
63
.89
Tre
sB
oca
sst
ream
–R
ub
ert
and
Giu
lian
o-C
aeta
no
(20
06
b)
7
Hyp
ost
om
us
stri
ga
tice
ps
72
10
16
46
63
.89
Jacu
tin
ga
riv
er–
Ru
ber
tan
dG
iuli
ano
-Cae
tan
o(2
00
6b
)7
Hyp
ost
om
us
stri
ga
tice
ps
72
10
16
46
63
.89
Taq
uar
iri
ver
–R
ub
ert
and
Giu
lian
o-C
aeta
no
(20
06
b)
7
Hyp
ost
om
us
stri
ga
tice
ps
72
86
30
28
80
.56
Car
rap
ato
stre
amS
PM
arti
nez
etal
.(2
00
8)
8
Hyp
ost
om
us
aff.
stri
ga
tice
ps
72
10
12
20
30
69
.44
Itai
pu
rese
rvo
irP
RB
aum
gar
tner
etal
.(2
00
9)
9
Hyp
ost
om
us
cf.
tiet
ensi
s6
81
81
01
22
85
8.8
2A
raq
ua
stre
amS
PIs
hid
aet
al.
(20
02
)5
Hyp
ost
om
us
cf.
top
ava
e8
06
84
22
48
2.5
0C
arra
pat
ost
ream
SP
Mar
tin
ezet
al.
(20
08
)8
Hyp
ost
om
us
un
ae
76
––
––
–C
on
tas
riv
erB
AB
iten
cou
rtet
al.
(20
08
)8
Hyp
ost
om
us
cf.
wu
cher
eri
76
10
18
48
63
.16
Un
ari
ver
BA
Bit
enco
urt
etal
.(2
00
9)
9
Hyp
ost
om
us
sp.
74
14
16
63
85
9.4
6–
–L
ara-
Kam
eian
dJu
lio
-Ju
nio
r(2
00
0)
4
Hyp
ost
om
us
sp.
72
14
20
38
52
.78
Ap
erta
do
sst
ream
and
Jata
izin
ho
riv
er
PR
Su
aki
etal
.(2
00
2)
5
Hyp
ost
om
us
sp.
68
––
––
–A
per
tad
os
stre
aman
d
Jata
izin
ho
riv
er
PR
Su
aki
etal
.(2
00
2)
5
Hyp
ost
om
us
sp.
74
81
02
82
87
5.6
8C
aval
ost
ream
SC
Mar
tin
ezet
al.
(20
06
)7
Hyp
ost
om
us
sp.
74
10
61
64
27
8.3
8A
rara
sst
ream
MG
Men
des
-Net
oet
al.
(20
06
)7
Hyp
ost
om
us
sp.
66
12
16
12
26
57
.58
Par
anap
anem
ab
asin
SP
Bra
nd
aoet
al.
(20
06
a)7
Hyp
ost
om
us
sp.
68
20
16
62
64
7.0
6M
og
i-G
uac
uri
ver
SP
Bra
nd
aoet
al.
(20
06
b)
7
Hyp
ost
om
us
sp.
80
––
––
–P
assa
-cin
cori
ver
SP
Tra
ldi
etal
.(2
00
8)
8
246 Rev Fish Biol Fisheries (2012) 22:241–250
123
Page 7
Ta
ble
2co
nti
nu
ed
Sp
ecie
s2
nC
hro
mo
som
efo
rmu
lae
st-a
%L
oca
lity
UF
Ref
eren
ces
Mee
tin
gs
msm
sta
Hyp
ost
om
us
sp.
68
12
12
83
66
4.7
1S
alto
Seg
red
oP
RM
auru
tto
etal
.(2
00
9)
9
Hyp
ost
om
us
sp.
76
82
04
86
3.1
6P
aran
apan
ema
riv
er–
Pen
tead
oet
al.
(20
09
)9
Hyp
ost
om
us
sp.
72
––
––
–A
rag
uar
iri
ver
MG
Co
rrei
aet
al.
(20
09
)9
Hyp
ost
om
us
sp.
16
8–
––
––
Juru
mim
rese
rvo
irS
PF
on
tan
ari
etal
.(1
99
6)
3
Hyp
ost
om
us
sp.
16
41
42
42
64
0.6
3A
rag
uai
ari
ver
MT
Oli
vei
raan
dV
ener
e(2
00
0),
Oli
vei
raet
al.
(20
02
)
4,
5
Hyp
ost
om
us
sp.
17
68
16
84
46
8.4
2K
elle
rst
ream
PR
Lar
a-K
amei
and
Juli
o-J
un
ior
(20
02
)5
Hyp
ost
om
us
sp.
27
2–
––
––
Juru
mim
rese
rvo
irS
PF
on
tan
ari
etal
.(1
99
6)
3
Hyp
ost
om
us
sp.
27
23
83
44
7.2
2A
rag
uai
ari
ver
MT
Oli
vei
raan
dV
ener
e(2
00
0)
4
Hyp
ost
om
us
sp.
27
42
64
86
4.8
6A
rag
uai
ari
ver
MT
Oli
vei
raan
dV
ener
e(2
00
0)
4
Hyp
ost
om
us
sp.
27
2–
––
––
Ara
gu
aia
riv
erM
TO
liv
eira
etal
.(2
00
2)
5
Hyp
ost
om
us
sp.
28
46
16
62
73
.81
Per
did
ori
ver
MS
Cer
eali
etal
.(2
00
6,
20
08
)7
,8
Hyp
ost
om
us
sp.
36
41
61
41
81
65
3.1
3A
rag
uai
ari
ver
MT
Oli
vei
raet
al.
(20
02
)5
Hyp
ost
om
us
sp.
37
68
61
64
68
1.5
8K
elle
rst
ream
PR
Lar
a-K
amei
and
Juli
o-J
un
ior
(20
02
)5
Hyp
ost
om
us
sp.
38
26
14
62
75
.61
Sal
ob
rari
ver
and
Sal
ob
rin
ha
stre
am
MS
Cer
eali
etal
.(2
00
6,
20
08
)7
,8
Hyp
ost
om
us
sp.
A6
8–
––
––
––
Ber
toll
oan
dS
ilv
a(1
99
0)
2
Hyp
ost
om
us
sp.
B7
4–
––
––
––
Ber
toll
oan
dS
ilv
a(1
99
0)
2
Hyp
ost
om
us
sp.
C7
2–
––
––
––
Ber
toll
oan
dS
ilv
a(1
99
0)
2
Hyp
ost
om
us
sp.
D7
2–
––
––
––
Ber
toll
oan
dS
ilv
a(1
99
0)
2
Hyp
ost
om
us
sp.
E8
0–
––
––
––
Ber
toll
oan
dS
ilv
a(1
99
0)
2
Mee
tin
gs:
1S
imp
osi
od
eci
tog
enet
ica
evo
luti
va
eap
lica
da
ap
eix
esn
eotr
op
icai
s(1
98
6),
2II
IS
imp
osi
od
eci
tog
enet
ica
evo
luti
va
eap
lica
da
ap
eix
esn
eotr
op
icai
s(1
99
0),
3V
I
Sim
po
sio
de
cito
gen
etic
aev
olu
tiv
ae
apli
cad
aa
pei
xes
neo
tro
pic
ais
(19
96
),4
VII
IS
imp
osi
od
eci
tog
enet
ica
eg
enet
ica
de
pei
xes
(20
00
),5
IXS
imp
osi
od
eci
tog
enet
ica
eg
enet
ica
de
pei
xes
(20
02
),6
XS
imp
osi
od
eci
tog
enet
ica
eg
enet
ica
de
pei
xes
(20
04
),7
XI
Bra
zili
ansy
mp
osi
um
on
fish
cyto
gen
etic
san
dg
enet
ics
(20
06
),8
XII
Sim
po
sio
de
cito
gen
etic
ae
gen
etic
ad
ep
eix
es(2
00
8),
9X
III
Sim
po
sio
de
cito
gen
etic
ae
gen
etic
ad
ep
eix
es(2
00
9)
a(c
ited
asH
ypo
sto
mu
slu
etke
ni)
Rev Fish Biol Fisheries (2012) 22:241–250 247
123
Page 8
further analyses on this matter. Our analysis consid-
ering only published data includes a relatively small
amount of new data compared to the analysis
performed by Artoni and Bertollo (2001), even so,
these new data cause the correlation coefficient to
decrease to r = 0.6048, P = 0.0008. The analysis
performed considering also the grey literature shows
an even smaller correlation coefficient (r = 0.3215,
P = 0.002), suggesting that it is not possible to
associate the increase of chromosome number to
higher percentages of subtelocentric and acrocentric
chromosomes for most of the species. The restriction
to higher proportions of subtelocentric and acrocen-
tric chromosomes seems to be characteristic to the
group of species with 2n = 80 chromosomes or
higher (including H. topavae), where subtelocentric
and acrocentric chromosomes represent from 70.00 to
82.50% of the karyotype (Fig. 4), while the groups
with lower diploid numbers have a great variation on
this proportion. Moreover, the analysis of variance
performed on these data shows that these differences
are not significant in both situations (with only the
published data and with both classes of data com-
piled), indicating that percentage of chromosome
types and diploid numbers are not always associated
(P-values equal to 0.05343 and 0.16714,
respectively).Fig. 2 Giemsa stained karyotype of a Hypostomus ancistro-ides and b Hypostomus topavae. The bar represents 5 lm
Fig. 3 Linear regression
between percentage of
subtelocentric-acrocentric
chromosomes (st-a %) and
diploid numbers (2n) in
species of Hypostomus,
considering only published
data. The size of the mark
(circles) represents the
number of populations/
species that share both the
same diploid number and
percentage of subtelocentric
and acrocentric
chromosomes
248 Rev Fish Biol Fisheries (2012) 22:241–250
123
Page 9
Amongst species with lower diploid numbers, the
mean percentage of subtelocentric and acrocentric
chromosomes is around 61.00%, but this percentage
varies considerably even between species that share
the same diploid number; therefore it is not possible
to correlate diploid numbers with the predominance
of certain chromosome types for most species of
Hypostomus (Fig. 4). The species with the lower
diploid number of the genus (H. plecostomus, with 54
chromosomes) has 55.56% of subtelocentric and
acrocentric chromosomes, even more than some
species like Hypostomus sp. G (40.63%), Hypostomus
ancistroides (44.12%) and Hypostomus sp. D1
(50.00%) with higher diploid numbers (64, 68 and
72 chromosomes, respectively, Table 1). These data
show that it is not possible to associate diploid
numbers to the percentage of subtelocentric and
acrocentric chromosomes in Hypostomus. The exis-
tence of variation among the karyotypes of species
with the same diploid number indicates that Robert-
sonian rearrangements and pericentric inversions
played an important part in the karyotypic differen-
tiation in Hypostomus (Kavalco et al. 2005).
Besides, the variation of the proportion of subtelocen-
tric and acrocentric chromosomes between species with
the same diploid number, such variation is also present
between populations of the same species, like in H. anci-
stroides, one of the most studied species in the genus.
Although all the analyzed populations show 2n = 68
chromosomes, the karyotypic formulae varies (Tables 1
and 2). Because of such variation, the percentage of
subtelocentric and acrocentric chromosomes in this
species ranges from 44.00% in the population analyzed
by Michele et al. (1977) to 79.00%, from a population
from Carrapato stream (Table 2).
Considering Hypostomus as a species rich genus
and their genetic and morphological variation, the
number of cytogenetically studied species is still
relatively small. Also, many of the analyzed species
are either new to science or unidentified.
Acknowledgments The authors are grateful to Instituto
Brasileiro do Meio Ambiente e Recursos Naturais
Renovaveis (MMA/IBAMA) for authorizing the capture of
the fishes. The authors thank to Unioeste and to the Nucleo de
Pesquisas em Limnologia, Ictiologia e Aquicultura (Nupelia)
for logistical support. This study was financed by Fundacao
Araucaria (Fundacao Araucaria de Apoio ao Desenvolvimento
Cientıfico e Tecnologico do Estado do Parana), CAPES
(Coordenadoria de Aperfeicoamento de Ensino Superior) and
CNPq (Conselho Nacional de Desenvolvimento Cientıfico e
Tecnologico).
Fig. 4 Linear regression
between percentage of
subtelocentric-acrocentric
chromosomes (st-a %) and
diploid numbers (2n) in
species of Hypostomus,
considering the published
data and grey literature. The
size of the mark (circles)
represents the number of
populations/species that
share both the same diploid
number and percentage of
subtelocentric and
acrocentric chromosomes
Rev Fish Biol Fisheries (2012) 22:241–250 249
123
Page 10
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