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Corresponding author.
E-mail address: lanszki@mail.atk.u-kaposvar.hu (J. Lanszki).
Please cite this article as: Lanszki, J., et al., Relative spraint density and genetic structure of otter (Lutra lutra) along the Drava River in....
Mamm. Biol. (2007), doi:10.1016/j.mambio.2007.08.005
population densities, between years and areas, using the
same sampling methods over time (Jefferies 1986;
Mason and Macdonald 1987; Reuther et al. 2000).
Most recently, molecular genetic methods offer a
promising alternative for population assessment. The
application of highly variable DNA markers, drawn from
non-invasive sampling of faecal material (Kohn and Wayne
1997; Taberlet and Luikart 1999) can yield estimates of
population size, genetic structure, mating patterns and sex
ratio (Dallas et al. 1999; Pertoldi et al. 2001; Dallas et al.
2002, 2003; Randi et al. 2003; Arrendal et al. 2004).
The objective of this 2-year study was to compare
traditional methods, i.e. different spraint density indices
with genetic-based estimates and to examine the genetic
structure of the otter population along the Drava River.
Along the River Drava in Hungary, the otter shows a
continuous presence (Lanszki 2005), but at low densities
across all habitats studied. Relative density changes
were monitored continuously from 2000 until 2005 using
spraint surveys. Collection of otter DNA from fresh
spraints and anal jellies (Dallas and Piertney 1998;
Dallas et al. 1999) was initiated in 2002.
Material and methods
Study area
The study was carried out in SW Hungary, at six sites along
the River Drava, three directly on the river, and three in
backwater habitats (from D1: 46118
0
N, 16152
0
E to B3:
45157
0
N, 17130
0
E, Fig. 1a). The main stem of the Drava River
has a steep riparian region, characterised by oodplain forests.
Backwaters areas are vegetated by willows, poplar, and ash-
alder woods. More information on the habitats is given in
Juha sz and De nes (2005) and Lanszki and Sallai (2006).
Sample collection and DNA analysis
Fresh spraint and anal jelly samples (Coxon et al. 1999)
were collected monthly between June 2002 and May 2004
(between 05.00 and 10.00 h) along standard routes (Table 1).
Spraints were also collected for diet analysis, but for DNA
extraction 1 ml of fresh spraint and anal jelly were placed in
plastic vials containing 8 ml of 96% ethanol, carried in a cooler
and stored at 20 1C until analysis.
DNA was extracted using a modied hexadecyltrimethy-
lammonium bromide (CTAB)-based extraction protocol
(Coxon et al. 1999). To reduce costs and work investment an
initial control step was included in the protocol, whereby the
lysate of the spraint samples was loaded on 2% agarose gel
(stained with ethidium bromide (0.5 mg/ml) to ensure DNA
presence. In this process, samples not containing DNA were
excluded from further procedures. This step cannot exclude
DNA from other species, such as from prey, so sucessful PCR
is still not guarenteed. After diatomaceous earth binding of the
DNA, simple centrifugation was used instead of high cost
separation by ltration. The DNA was nally stored in water
(MilliQ) at 20 1C.
Nine microsatellite loci were selected Lut-435, Lut-604, Lut-
615, Lut-701, Lut-715, Lut-717, Lut-733, Lut-832 and Lut-833
described in Dallas and Piertney (1998). Annealing tempera-
ture was either 60 1C (Lut-615, Lut-833, Lut-701, Lut-715,
Lut-717, Lut-733) or 58 1C (Lut-435, Lut-604, Lut-832). The
locus Lut-SRY was used for sex identication (Dallas et al.
2000). Duplex PCR were used to identify sex from spraint
samples, where Lut-SRY was combined with Lut-701 or Lut-
615 (annealing: 60 1C). PCR reactions were performed in a
15 ml containing 50 ng of L. lutra genomic DNA, 0.01 units/ml
Dynazyme (Finnzyme) polymerase, 2.5 mM MgCl
2
, 200 mM of
each dNTP, 0.25 mM of each forward and reverse primers and
sterile ultrapure MilliQ water. The PCR programme used was:
95 1C/4 min (95 1C/15 s., annealing 30 s, 72 1C/60 s) 30 cycles,
72 1C/9 min, 20 1C/1 min. For spraint samples the number of
amplication cycles had to be increased (to 33) in order to
produce detectable PCR products (Ramekers et al. 1997). To
measure the quality and quantity of PCR products samples
were loaded on agarose gel (2% agarose gel stained with
ethidium bromide, 0.5 mg/ml). If amplication failed (no target
product detected) the PCR reaction was repeated three more
times. PCR products were genotyped (Pertoldi et al. 2001) on
an ALF Express II DNA sequencer (Amersham-Biosciences).
Internal and external molecular mass standards (100, 200,
300 bp) were used for sizing microsatellite allels and allelic
ARTICLE IN PRESS
Fig. 1. (a) Locality of the study areas in the Drava region. River sections: D1 O
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J. Lanszki et al. / Mamm. biol. ] (]]]]) ]]]]]] 5
Please cite this article as: Lanszki, J., et al., Relative spraint density and genetic structure of otter (Lutra lutra) along the Drava River in....
Mamm. Biol. (2007), doi:10.1016/j.mambio.2007.08.005
alleles (5.0) were well within the ranges of those reported
in both European and Asian studies (e.g., Randi et al.
2003; Hung et al. 2004). However, our result of nine
alleles at the Lut-717 locus was higher than that
reported across studies screening 1400 indviduals
(summarising data can be found in Table 4).
In conclusion, the spraint survey method yielded
results in close relation with abundance of otters, as
revealed by non-invasive molecular techniques. These
results may be universal for the monitoring in otter
populations in primarily natural freshwater habitats. In
this rst study of otters in Hungary, based on non-
invasive sampling, high genetic diversity, no migration
of individuals and only low densities were found.
Further investigations, with increased DNA extraction
efciency, carried out on a larger catchment area needed
to better estimate patterns of metapopulation structure
and dynamics.
Acknowledgements
We express sincere thanks to Drs. Jim Conroy and
Rob Marrs for advice and comments on early draft of
the manuscript. This work was supported by the
Hungarian Fund for Scientic Research (OTKA F
037557 and K 62216), the Directorship of the Danube-
Drava National Park and the Bolyai Scholarship (JL).
Zusammenfassung
Relative Losungsdichte und genetische Struktur von
Fischottern (Lutra lutra) entlang der Drau in Ungarn
In dieser Untersuchung verwendeten wir genetische
Methoden, um die Populationsgro e und -struktur des
eurasischen Fischotters am Flu Drau in Ungarn
abzuscha tzen. Die Ergebnisse wurden mit jenen aus
traditionellen Untersuchungen verglichen. Die relative
Losungsdichte wurde berechnet auf Grund der Anzahl
von frischem (D
f
) und der Anzahl von gesamter Losung
(D
t
), die entlang von Standardrouten in einem zwei-
ja hrigen Zeitraum gesammelt wurden. Neun Mikrosa-
telliten wurden verwendet, die insgesamt 45 Allele
aufwiesen. Dadurch konnten 17 individuelle Genotypen
unterschieden werden. Die erwartete Heterozygotie
reichte von 0.53 bis 0.89, und die beobachtete Hetero-
zygotie reichte von 0.25 bis 0.92. Die mittlere Dichte
(D
g
) u ber sechs Stellen war 0.17 Individuen/km. Es
gab eine signikante Korrelation zwischen der Anzahl
von Genotypen und der Anzahl gefundener Losung
ARTICLE IN PRESS
Table 4. Summary data of molecular genetic analyses of Eurasian otter (Lutra lutra) populations
Location Sample
type
Number of H
e
/H
o
Mean A Source
samples loci
Great Britain, Ireland,
Germany
T 32 13 n/0.55 6.69 Dallas and Piertney
(1998)
Denmark T, M 124 9 0.46/0.43 3.89 Pertoldi et al. (2001)
England, Wales,
Scotland, N. Isles
T 618 12 0.55/n 7.08 Dallas et al. (2002)
England T, F 122 7-9 0.54/n 5.11 Dallas et al. (2003)
Great Britain, Ireland,
Spain, Lithuania,
Denmark, Germany,
Sweden, France
T, O 102 11 0.74/0.55 8.09 Randi et al. (2003)
Germany F, T 59 6 0.65/0.63 5.30 Kalz et al. (2006)
Norway, Sweden T, M 114 6 0.65/0.65 8.50 Arrendal et al. (2004)
Taiwan F 38 7 0.61/0.76 3.86 Hung et al. (2004)
Czech Republic T, F 132 10 0.53/0.51 4.50 Ha jkova et al. (2007)
Slovak Republic T, F 65 10 0.59/0.55 4.70 Ha jkova et al. (2007)
Hungary F 17 9 0.68/0.53 5.00 Present study
Sample type: T: tissue, M: museum tissue (e.g., skull), F: faecal sample, O: other (e.g., hair, blood); loci: number of examined microsatellite loci; H
e
/
H
o
: expected and observed heterozygosity, n: no calculation available; mean A: mean number of alleles.
Table 3. Matrix of Nei minimum genetic distances (D
m
)
among ve otter sub-populations (on the basis of n 15
individuals) along the Drava River, Hungary
Habitats D2 D3 B1 B2
D1 0.107 0.122 0.177 0.155
D2 0.133 0.181 0.081
D3 0.212 0.117
B1 0.187
In the B3 habitat only one individual was genotyped, and only three
microsatellites amplied (for abbreviations see Table 1).
J. Lanszki et al. / Mamm. biol. ] (]]]]) ]]]]]] 6
Please cite this article as: Lanszki, J., et al., Relative spraint density and genetic structure of otter (Lutra lutra) along the Drava River in....
Mamm. Biol. (2007), doi:10.1016/j.mambio.2007.08.005
entlang von Standardrouten (frische Losung: r
P
0.85,
Po0.01; gesamte Losung r
P
0.76, Po0.05). Alle
Genotypen, die innerhalb des 50 km langen Untersu-
chungsgebietes gefunden wurden, waren nah mit einan-
der verwandt (D
m
reichte von 0.08 bis 0.21).
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Please cite this article as: Lanszki, J., et al., Relative spraint density and genetic structure of otter (Lutra lutra) along the Drava River in....
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Please cite this article as: Lanszki, J., et al., Relative spraint density and genetic structure of otter (Lutra lutra) along the Drava River in....
Mamm. Biol. (2007), doi:10.1016/j.mambio.2007.08.005