Population dynamics of the periwinkle Littorina littorea (Linnaeus, 1758) in the East Frisian Wadden Sea

Introduction
Objectives
Methods
Results
Conclusions
Population dynamics of the periwinkle Littorina
littorea (Linnaeus, 1758) in the East Frisian
Wadden Sea
Diplomarbeit
Alexander Jüterbock
Animal Biodiversity and Evolutionary Biology
Carl von Ossietzky University Oldenburg
Supervisors: Prof. Dr. Gabriele Gerlach, Dr. Thomas Friedl
April 13, 2010
Alexander Jüterbock Population dynamics of the periwinkle

Introduction
Objectives
Methods
Results
Conclusions
Outline
1 Introduction
2 Objectives
3 Methods
4 Results
5 Conclusions

Introduction
Objectives
Methods
Results
Conclusions
Populations get Connected by Dispersing Larvae
BENTHOSPLANKTON
Population 1 Population 3Population 2
adults adults adults

Introduction
Objectives
Methods
Results
Conclusions
BENTHOSPLANKTON
larvae

Introduction
Objectives
Methods
Results
Conclusions
BENTHOSPLANKTON
larvae
Reef ﬁsh
Photo: Randall, J.E.
Study: Gerlach et al., 2007

Introduction
Objectives
Methods
Results
Conclusions
BENTHOSPLANKTON
larvae
Corals
Photo: Ronald L. Shimek
Study: Whitaker, 2004

Introduction
Objectives
Methods
Results
Conclusions
BENTHOSPLANKTON
larvae
Gastropods
Photo: Keith Hiscock
Study: Dupont et al., 2007

Introduction
Objectives
Methods
Results
Conclusions
BENTHOSPLANKTON
larvae
Divergent selection

Introduction
Objectives
Methods
Results
Conclusions
BENTHOSPLANKTON
larvae larvae larvae
Divergent selection

Introduction
Objectives
Methods
Results
Conclusions
BENTHOSPLANKTON
larvae larvae larvae
Divergent selection
Larval recruitment

Introduction
Objectives
Methods
Results
Conclusions
Hydrodynamics of the East Frisian Wadden Sea
a
15
10
5
0
km
0 5 10 15 20 25 30 35 40 45 50 55 60
km
0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 45 50 55 60
(cm/s)
b
15
Norderney Langeoog Spiekeroog
(Staneva et al., 2009)

Introduction
Objectives
Methods
Results
Conclusions
The Periwinkle’s Life Cycle Includes a Planktotrophic
Larva
(Fretter and Graham, 1962)
(Reid, 1996)
4–7 weeks

Introduction
Objectives
Methods
Results
Conclusions
Main Objectives of the Study
Objectives
Larval recruitment?
Population preference?
Morphological differentiation?
Genetic differentiation?

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Population Preference
Shell Morphology
Population Genetics
Outline
1 Introduction
2 Objectives
3 Methods
4 Results
5 Conclusions

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Outline
3 Methods
Larval Recruitment
Shell Morphology
Population Genetics

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Larval Dispersal in the Wadden Sea was Simulated
2000
1500
1000
500
0
a
b
L1
Baltrum
Langeoog
Spiekeroog
Wangerooge
Programmed by: Gräwe, U.

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Sampling Sites
0 500 1000
kilometersN
EFWS

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Sampling Sites
0 50 100
kilometers N
Borkum
Norderney
Langeoog
Wangerooge
B2
B2.2
B1
N3
N2N1
L1
W1
W2

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Sampling Sites
0 1000 2000
kilometers N
Woods HoleNew York
USA
CANADA

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Conspeciﬁcs Aggregate on Hard Bottom Substrate
(Geller–Grimm, 2000)

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Snails of 2 Populations Aggregated in Basins
10 cm
60 cm60 cm
Population 1 Population 2

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Snails of 2 Populations Aggregated in Basins
10 cm
60 cm60 cm
60 cm
60cm
Individual from population 1

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Analysis of Aggregation Experiments
. . . one hour later
10 cm
60 cm60 cm
1
2
3
4
5
- Population Preference Index PPI -

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Testing Snails in Olfactory Choice Flumes
22 cm
5 cm
Snail at startposition
Stimulus water
from population 1
Stimulus water
from population 2

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
22 cm
5 cm
Stimulus water
from population 1
Stimulus water
from population 2
. . . %
. . . %

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
22 cm
5 cm
Stimulus water
from population 1
Stimulus water
from population 2
. . . %
. . . %
Wilcoxon signed-rank test

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Measurements of the Shell
CL
SP
SW
LA
Linear discriminant analysis

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Genetic Differentiation at Microsatellite Markers
48–55 individuals/population
5 microsatellite loci
Differentiation index Dest,
R package DEMEtics (Jüterbock et al., 2010)

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Outline
4 Results
Larval Recruitment
Shell Morphology
Population Genetics

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
No Local Recruitment by Passive Drift in the EFWS
2000
1500
1000
500
0
a
b
L1
Baltrum
Langeoog
Spiekeroog
Wangerooge

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
No Local Recruitment by Passive Drift in the EFWS
. . . one week later
300
250
200
150
100
50
0
Baltrum
Langeoog
Spiekeroog
Wangerooge
a

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Snails Prefer Conspecifics of the Own Population
Population-wise aggregation
PPI significant (p ≤ 0.1) in 6 of 17 tests
Attraction to volatile chemicals
No significant preference in any of 18 tests

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Populations Differ Morphologically
Shell morphology differs
between . . .
Woods Hole – EFWS
North coasts – South
coasts
North coasts
South coasts
Woods Hole
EFWS
Wilks’ Lambda = 0.867, p < 0.001 ***

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
between . . .
Woods Hole – EFWS
coasts
North coasts
South coasts
Borkum
Norderney
Langeoog
Wangerooge
Wilks’ Lambda = 0.802, p < 0.001 ***

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
between . . .
Woods Hole – EFWS
coasts
North coasts
South coasts
Borkum
Norderney
Langeoog
Wangerooge
Wilks’ Lambda = 0.135, p < 0.001 ***

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
between . . .
Woods Hole – EFWS
coasts
North coasts
South coasts
Borkum
Norderney
Langeoog
Wangerooge
Wilks’ Lambda = 0.731, p < 0.001 ***

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Genetic Distance . . .
. . . between Woods Hole and East Frisian populations
Dest = 0.033–0.057 ***
p ≤ 0.05
Woods Hole
EFWS

Introduction
Objectives
Methods
Results
Conclusions
Larval Recruitment
Shell Morphology
Population Genetics
Genetic Distance . . .
. . . between islands
Dest = -0.007–0.001
p > 0.05 Borkum
Norderney
Langeoog
Wangerooge

Introduction
Objectives
Methods
Results
Conclusions
Do the Results Indicate Reproductive Isolation?
Objectives
Larval recruitment?
Population preference?
Morphological
differentiation?
Genetic differentiation?
Unanswered questions
Larval behavior?
Intrinsic differences
Recognition cues?
Assortative mating?
Expressed loci?
Phenotypic plasticity?

Introduction
Objectives
Methods
Results
Conclusions
Do the Results Indicate Reproductive Isolation?
Overall conclusion

Introduction
Objectives
Methods
Results
Conclusions
Acknowledgements
Gabriele Gerlach
Thomas Glatzel
Thomas Friedl
Ulf Gräwe
Peter Harmand
Achim Wehrmann
Anke Müller
Andreas Sommer
Philipp Krämer
Jana Deppermann
Cornelia Hinz
Jelle Atema
René Spierling
Elke Frahmann
Marén Bökamp
Funded by the
Niedersächsische
Wattenmeerstiftung

Appendix
Further Reading
Introduction
Methods
Results
For Further Reading I
Reid, D.G. (1996)
Systematics and Evolution of Littorina.
The Ray Society, London.
Cowen, R.K. & Sponaugle, S. (2009)
Larval dispersal and marine population connectivity
Annual Review of Marine Science 1:443–466.

Appendix
Further Reading
Introduction
Methods
Results
For Further Reading II
Staneva, J.; Stanev, E.V.; Wolff, J.-O.; Badewien, T.H.;
Reuter, R.; Flemming, B.; Bartholomä, A. & Bolding, K.
(2009)
Hydrodynamics and sediment dynamics in the German
Bight. A focus on observations and numerical modelling in
the East Frisian Wadden Sea.
Continental Shelf Research 29(1):302–319.
Jost, L. (2008)
Gst and its relatives do not measure differentiation.
Molecular Ecology 17(18):4015–4026.

Appendix
Further Reading
Introduction
Methods
Results
Residual Current in the German Bight
55.4N
55.2N
55.0N
54.8N
54.6N
54.4N
54.2N
54.0N
53.8N
53.6N
53.4N
6.3E
6.6E
6.9E
7.2E
7.5E
7.8E
8.1E
8.4E
8.7E
9.0E
(m/s)
0.16
0.14
0.12
0.10
0.08
0.06
0.04
0.02
0.25
(Staneva et al., 2009)

Appendix
Further Reading
Introduction
Methods
Results
L. littorea Occurs on Both Sides of the Atlantic
(Reid, 1996)

Appendix
Further Reading
Introduction
Methods
Results
Larva
a b| ~
(Reid, 1996)
5–6 days
4–7 weeks
12–18 months
1–12 hours
after fertilization

Appendix
Further Reading
Introduction
Methods
Results
Larva
a b| ~
12–18 mon

Appendix
Further Reading
Introduction
Methods
Results
Sampled Snails Were Kept in Aquaria

Appendix
Further Reading
Introduction
Methods
Results
Snails Were Individually Characterized
CL
Shell height Barnacle fouling Sex

Appendix
Further Reading
Introduction
Methods
Results
Snails Were Individually Characterized
Shell height Barnacle fouling Sex

Appendix
Further Reading
Introduction
Methods
Results
Aggregation Experiments Were Performed in Special
Basins
10 cm
60 cm60 cm
60 cm
60cm

Appendix
Further Reading
Introduction
Methods
Results
Detailed Analysis of Aggregation experiments
. . . one hour later
10 cm
60 cm60 cm
. . . 15 tests at least

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5
Are certain types of
conspeciﬁcs preferred?
Snails of the same
population
Snails of the same sex
Snails of the same size

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5
- PPI -
Snails of the same
population

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
|||
|
|
1
|
||
2
|
|
3
~~
~ 4
~~~
~
5
~
~
- Sex-PI -
Snails of the same
population

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5
- Size-PI -
Snails of the same
population

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5
PPI and Size-PI, both high
Snails of the same
population

Appendix
Further Reading
Introduction
Methods
Results
Deﬁning the Population Preference Index
Population mixture
p = var =
Real population preference
p = var =
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =
p = var =
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p = var =
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =0.5 var =
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =0.5 var =0.263
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =0.5 var =0.263
Ambiguous
p =1.0 var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =0.5 var =0.263
Ambiguous
p =1.0 var =0.000

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 PPI =0.000
p =0.5 PPI =0.263
Ambiguous
p =1.0 PPI =0.000

Appendix
Further Reading
Introduction
Methods
Results
Deﬁning the Population Preference Index in Detail
Population mixture
p = var =
p = var =
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =
5
10 ∗10+ 5
10 ∗10
20
var =
p = var =
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =
( 5
10 −0.5)2∗10+( 5
10 −0.5)2∗10
19
p = var =
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p = var =
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =
10
10 ∗10+ 0
10 ∗10
20
var =
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =0.5 var =
( 10
10 −0.5)2∗10+( 0
10 −0.5)2∗10
19
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =0.5 var =0.263
Ambiguous
p = var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =0.5 var =0.263
Ambiguous
p =
10
10 ∗10
10
var =

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =0.5 var =0.263
Ambiguous
p =1.0 var =
( 10
10 −1)2∗10
9

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 var =0.000
p =0.5 var =0.263
Ambiguous
p =1.0 var =0.000

Appendix
Further Reading
Introduction
Methods
Results
Population mixture
p =0.5 PPI =0.000
p =0.5 PPI =0.263
Ambiguous
p =1.0 PPI =0.000

Appendix
Further Reading
Introduction
Methods
Results
Snails Aggregate Assortatively
Borkum
Norderney
Langeoog
Wangerooge
B1 B2 N3 N1 N2 L1 W1 W2 WH
B1
B2
N3
N1
N2
L1
W1
W2
****
·
**
·
·
· **
Population preference
signiﬁcance
Size preference
signiﬁcance
· p ≤ 0.10,* p ≤ 0.05, ** p ≤ 0.01,*** p ≤ 0.001

Appendix
Further Reading
Introduction
Methods
Results
Simpliﬁed Monte Carlo Simulation
10 cm
60 cm60 cm
1
2
3
4
5

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5
Random allocationRandom allocation

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5
Calculation of PPI

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5
A histogram of PPI values
Mean PPI
Frequency
0.00 0.02 0.04 0.06 0.08 0.10
0
20
40
60
80
100100
80
60
40
20
0
Frequency
0.00 0.02 0.04 0.06 0.08 0.10
Mean PPI
Critical value
90% < 0.068
signiﬁcantnot signiﬁcant

Appendix
Further Reading
Introduction
Methods
Results
Detailed Monte Carlo Simulation
10 cm
60 cm60 cm
1
2
3
4
5
~~|
~|||~~| ~|~|~||~|~

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5
~~|
~|||~~| ~|~|~||~|~
Random allocationRandom allocation

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
~||
~| 1
||~
2
|
| 3
~|~
4
~~
|
~
5
~
|
~

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
~||
~| 1
||~
2
|
| 3
~|~
4
~~
|
~
5
~
|
~
Calculation of PPI, Sex-PI and Size-PI

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
1
2
3
4
5
~~|
~|||~~| ~|~|~||~|~
1000-fold repetition

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
|
~|
~
~ 1
|~|
2
|
| 3
~|~
4
~~
|
~5
|~
|
1000-fold calculation of PPI, Sex-PI and Size-PI

Appendix
Further Reading
Introduction
Methods
Results
10 cm
60 cm60 cm
|
~|
~
~ 1
|~|
2
|
| 3
~|~
4
~~
|
~5
|~
|
A histogram of PPI values
Mean PPI
Frequency
0.00 0.02 0.04 0.06 0.08 0.10
0
20
40
60
80
100100
80
60
40
20
0
Frequency
0.00 0.02 0.04 0.06 0.08 0.10
Mean PPI
Critical value
90% < 0.068
signiﬁcantnot signiﬁcant

Appendix
Further Reading
Introduction
Methods
Results
Testing Snails in Olfactory Flumes
22 cm
5 cm
Stimulus water
from population 1
Stimulus water
from population 2

Appendix
Further Reading
Introduction
Methods
Results
Testing Snails in Olfactory Flumes in Detail
22 cm
5 cm
Stimulus water B
Stimulus water A

Appendix
Further Reading
Introduction
Methods
Results
22 cm
5 cm
Stimulus water A
Stimulus water B1

Appendix
Further Reading
Introduction
Methods
Results
22 cm
5 cm
Stimulus water B
Stimulus water A1

Appendix
Further Reading
Introduction
Methods
Results
22 cm
5 cm
Stimulus water A
Stimulus water B1
1

Appendix
Further Reading
Introduction
Methods
Results
22 cm
5 cm
Stimulus water B
Stimulus water A1
1

Appendix
Further Reading
Introduction
Methods
Results
22 cm
5 cm
Stimulus water B
Stimulus water A
1
2

Appendix
Further Reading
Introduction
Methods
Results
22 cm
5 cm
Stimulus water B
Stimulus water A
1
3

Appendix
Further Reading
Introduction
Methods
Results
22 cm
5 cm
Stimulus water B
Stimulus water A75%
25%

Appendix
Further Reading
Introduction
Methods
Results
22 cm
5 cm
Stimulus water B
Stimulus water A
Difference
75%−25% = 50%

Appendix
Further Reading
Introduction
Methods
Results
15 Differences
50%
25%
.
.
.
0%
Wilcoxon signed-rank
test

Appendix
Further Reading
Introduction
Methods
Results
Allometric correction
Arithmetic plot
0 2 4 6 8 10 12 14
0
2
4
6
8
10
12
14
x = shell length (cm)
y=shellwidth(cm)
y = 0.13 ∗ x1.32

Appendix
Further Reading
Introduction
Methods
Results
Logarithmic plot
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
log10(x) = log10(shell length (cm))
log10(y)=log10(shellwidth(cm))
log10(y) = log10(0.13) + 1.32 ∗ log10(x)

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
log10(y) = log10(0.13)+1.32∗log10(x)
1
1.32 1.32
1 = 1.32
b: slope = allometric coefﬁcient

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
log10(y) = log10(0.13)+1.32∗log10(x)
1
1.32 1.32
1 = 1.32
a: intersection of y-axis when x = 0

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
log10(y) = log10(0.13)+1.32∗log10(x)
1
1.32 1.32
1 = 1.32
log(y) = log(a)+b∗log(x)

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
log10(y) = log10(0.13)+1.32∗log10(x)
1
1.32 1.32
1 = 1.32
log(y) = log(a)+b∗log(x)
log(a) = log(y)−b∗log(x)

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
spread of x

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
spread of x
x mean of x

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
spread of x
x mean of x
log(x)

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
spread of x
x mean of x
log(x)
Correction
log(a) = log(y)−b∗(log(x)−log(x))

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
spread of x
x mean of x
log(x)
Correction
log(a) = log(y)−b∗(log(x)−log(x))log(a) = log(y)−b∗(log(x)−log(x))

Appendix
Further Reading
Introduction
Methods
Results
−3 −2 −1 0 1 2 3
−3
−2
−1
0
1
2
3
spread of x
x mean of x
log(x)
Correction

Appendix
Further Reading
Introduction
Methods
Results
Genetic Differentiation at Microsatellite Markers 1
Pop1 Pop2

Appendix
Further Reading
Introduction
Methods
Results
Pop1 Pop2
DNA extraction

Appendix
Further Reading
Introduction
Methods
Results
Pop1 Pop2
DNA extraction
PCR

Appendix
Further Reading
Introduction
Methods
Results
Pop1 Pop2
-
+
DNA extraction
PCR
gel electrophoresis

Appendix
Further Reading
Introduction
Methods
Results
Pop1 Pop2
-
+
DNA extraction
PCR
gel electrophoresis
178
212
162

Appendix
Further Reading
Introduction
Methods
Results
Pop1 Pop2
-
+
DNA extraction
PCR
gel electrophoresis
178
212
162
Pop1 Pop2
allele1 allele2 allele1 allele2
178 178 212 162

Appendix
Further Reading
Introduction
Methods
Results
Pop1 Pop2
-
+
DNA extraction
PCR
gel electrophoresis
178
212
162
Pop1 Pop2
178 178
178 234
234 234
234 178
... ...
... ...
... ...
212 162
162 162
212 212
212 162
... ...
... ...
... ...

Appendix
Further Reading
Introduction
Methods
Results
Pop1 Pop2
-
+
DNA extraction
PCR
gel electrophoresis
178
212
162
Pop1 Pop2
178 178
178 234
234 234
234 178
... ...
... ...
... ...
212 162
162 162
212 212
212 162
... ...
... ...
... ...
Differentiation index Dest

Appendix
Further Reading
Introduction
Methods
Results
CAT
GTA

Appendix
Further Reading
Introduction
Methods
Results
CATCATCATCATCATCAT
GTAGTAGTAGTAGTAGTA

Appendix
Further Reading
Introduction
Methods
Results
GTCTCAGC CATCATCATCATCATCATACATCGAC
CAGAGTCG GTAGTAGTAGTAGTAGTATGTAGCTG

Appendix
Further Reading
Introduction
Methods
Results
Repeat no.
12

Appendix
Further Reading
Introduction
Methods
Results
Repeat no.
12
9

Appendix
Further Reading
Introduction
Methods
Results
Repeat no.
12
9
15
15

Appendix
Further Reading
Introduction
Methods
Results
Repeat no.
12
9
15
15
ampliﬁcation by
PCR

Appendix
Further Reading
Introduction
Methods
Results
Repeat no.
12
9
15
15
15
12
9
gel
electrophoresis

Appendix
Further Reading
Introduction
Methods
Results
Repeat no.
12
9
15
15
15
12
9

Appendix
Further Reading
Introduction
Methods
Results
Repeat no.
12
9
15
15
15
12
9
15
9 12

Appendix
Further Reading
Introduction
Methods
Results
Repeat no.
12
9
15
15
15
12
9
15
9 12
Pop1
165 165
230 165
230 230
165 230
230 230
165 165
165 165
230 165
Pop2
170 170
198 170
198 198
170 198
198 198
170 170
170 170
198 170

Appendix
Further Reading
Introduction
Methods
Results
Repeat no.
12
9
15
15
15
12
9
15
9 12
Pop1
165 165
230 165
230 230
165 230
230 230
165 165
165 165
230 165
Pop2
170 170
198 170
198 198
170 198
198 198
170 170
170 170
198 170
Difference Dest

Appendix
Further Reading
Introduction
Methods
Results
Snails Prefer Conspecifics of the own Population
Population-wise aggregation
PPI significant (p ≤ 0.1) in 6 of 17 tests
Sex-bias aggregation
~~~~~
~
~~~~ ||
||
|||||
| Sex-PI significant (p ≤ 0.1) in 0 of 17 tests
Size-wise aggregation
Size-PI significant (p ≤ 0.1) in 2 of 17 tests

Appendix
Further Reading
Introduction
Methods
Results
Populations
differ
intrinsically

Appendix
Further Reading
Introduction
Methods
Results
. . . aggregation frequency
increases
during the
mating season

Appendix
Further Reading
Introduction
Methods
Results
Snails do not Prefer any Volatile ChemicalsPreference(%)
−0.6
−0.4
−0.2
0.0
0.2
0.4
0.6
B2
−
L1L1
−
B2B2
−
N
1N
1
−
B2N
1
−
L1L1
−
N
1N
2
−
N
3N
3
−
N
2W
1
−
B2B2
−
W
1W
1
−
L1L1
−
W
1W
1
−
N
1N
1
−
W
1W
2
−
N
2N
2
−
W
2
W
H
−
B2.2
B2.2
−
W
H
N
1m
a
−
N
1fe
N
1fe
−
N
1m
a
B1u
−
B1m
B1m
−
B1uN
1
−
Sw
0.93 0.32 0.54 0.47 0.59 0.62 1 0.22 0.12 0.19 0.81 0.58 0.91 1 0.99 0.22 1 0.43 0.19 0.76 0.52 0.1 0.59
own
other
.
0.6
0.4
0.2
0.0
-0.2
-0.4
-0.6
Preference(%)
ownother
B2–L1L1–B2B2–N
1N
1–B2N
1–L1L1–N
1N
2–N
3N
3–N
2W
1–B2B2–W
1W
1–L1L1–W
1W
1–N
1N
1-W
1W
2–N
2N
2–W
2
W
H
–B2.2
B2.2–W
H
N
1m
a–N
1fe
N
1fe–N
1m
aB1u–B1tB1t–B1uN
1–Sw

Appendix
Further Reading
Introduction
Methods
Results
−0.6
−0.4
−0.2
0.0
0.2
0.4
0.6
B2
−
L1L1
−
B2B2
−
N
1N
1
−
B2N
1
−
L1L1
−
N
1N
2
−
N
3N
3
−
N
2W
1
−
B2B2
−
W
1W
1
−
L1L1
−
W
1W
1
−
N
1N
1
−
W
1W
2
−
N
2N
2
−
W
2
W
H
−
B2.2
B2.2
−
W
H
N
1m
a
−
N
1fe
N
1fe
−
N
1m
a
B1u
−
B1m
B1m
−
B1uN
1
−
Sw
0.93 0.32 0.54 0.47 0.59 0.62 1 0.22 0.12 0.19 0.81 0.58 0.91 1 0.99 0.22 1 0.43 0.19 0.76 0.52 0.1 0.59
own
other
.
0.6
0.4
0.2
0.0
-0.2
-0.4
-0.6
Preference(%)
ownother
1N
1–B2N
1–L1L1–N
1N
2–N
3N
3–N
2W
1–B2B2–W
1W
1–L1L1–W
1W
1–N
1N
1-W
1W
2–N
2N
2–W
2
W
H
–B2.2
B2.2–W
H
N
1m
a–N
1fe
N
1fe–N
1m
1–Sw
Population preference

Appendix
Further Reading
Introduction
Methods
Results
−0.6
−0.4
−0.2
0.0
0.2
0.4
0.6
B2
−
L1L1
−
B2B2
−
N
1N
1
−
B2N
1
−
L1L1
−
N
1N
2
−
N
3N
3
−
N
2W
1
−
B2B2
−
W
1W
1
−
L1L1
−
W
1W
1
−
N
1N
1
−
W
1W
2
−
N
2N
2
−
W
2
W
H
−
B2.2
B2.2
−
W
H
N
1m
a
−
N
1fe
N
1fe
−
N
1m
a
B1u
−
B1m
B1m
−
B1uN
1
−
Sw
0.93 0.32 0.54 0.47 0.59 0.62 1 0.22 0.12 0.19 0.81 0.58 0.91 1 0.99 0.22 1 0.43 0.19 0.76 0.52 0.1 0.59
own
other
.
0.6
0.4
0.2
0.0
-0.2
-0.4
-0.6
Preference(%)
ownother
1N
1–B2N
1–L1L1–N
1N
2–N
3N
3–N
2W
1–B2B2–W
1W
1–L1L1–W
1W
1–N
1N
1-W
1W
2–N
2N
2–W
2
W
H
–B2.2
B2.2–W
H
N
1m
a–N
1fe
N
1fe–N
1m
1–Sw
Sex preference

Appendix
Further Reading
Introduction
Methods
Results
−0.6
−0.4
−0.2
0.0
0.2
0.4
0.6
B2
−
L1L1
−
B2B2
−
N
1N
1
−
B2N
1
−
L1L1
−
N
1N
2
−
N
3N
3
−
N
2W
1
−
B2B2
−
W
1W
1
−
L1L1
−
W
1W
1
−
N
1N
1
−
W
1W
2
−
N
2N
2
−
W
2
W
H
−
B2.2
B2.2
−
W
H
N
1m
a
−
N
1fe
N
1fe
−
N
1m
a
B1u
−
B1m
B1m
−
B1uN
1
−
Sw
0.93 0.32 0.54 0.47 0.59 0.62 1 0.22 0.12 0.19 0.81 0.58 0.91 1 0.99 0.22 1 0.43 0.19 0.76 0.52 0.1 0.59
own
other
.
0.6
0.4
0.2
0.0
-0.2
-0.4
-0.6
Preference(%)
ownother
1N
1–B2N
1–L1L1–N
1N
2–N
3N
3–N
2W
1–B2B2–W
1W
1–L1L1–W
1W
1–N
1N
1-W
1W
2–N
2N
2–W
2
W
H
–B2.2
B2.2–W
H
N
1m
a–N
1fe
N
1fe–N
1m
1–Sw
Control for effect of
label color

Appendix
Further Reading
Introduction
Methods
Results
−0.6
−0.4
−0.2
0.0
0.2
0.4
0.6
B2
−
L1L1
−
B2B2
−
N
1N
1
−
B2N
1
−
L1L1
−
N
1N
2
−
N
3N
3
−
N
2W
1
−
B2B2
−
W
1W
1
−
L1L1
−
W
1W
1
−
N
1N
1
−
W
1W
2
−
N
2N
2
−
W
2
W
H
−
B2.2
B2.2
−
W
H
N
1m
a
−
N
1fe
N
1fe
−
N
1m
a
B1u
−
B1m
B1m
−
B1uN
1
−
Sw
0.93 0.32 0.54 0.47 0.59 0.62 1 0.22 0.12 0.19 0.81 0.58 0.91 1 0.99 0.22 1 0.43 0.19 0.76 0.52 0.1 0.59
own
other
.
0.6
0.4
0.2
0.0
-0.2
-0.4
-0.6
Preference(%)
ownother
1N
1–B2N
1–L1L1–N
1N
2–N
3N
3–N
2W
1–B2B2–W
1W
1–L1L1–W
1W
1–N
1N
1-W
1W
2–N
2N
2–W
2
W
H
–B2.2
B2.2–W
H
N
1m
a–N
1fe
N
1fe–N
1m
1–Sw
Own population –
seawater

Appendix
Further Reading
Introduction
Methods
Results
Snails do not Prefer any Volatile Chemicals
Populations
are
not discriminated
by
volatile chemicals

Appendix
Further Reading
Introduction
Methods
Results
Results of Discriminant analyzes
EFWS – Woods Hole
9
8
7
6
5
4
3
2
-2 -1 0 1 2 3 4 5 6 7
Canonical2
Canonical 1
-SP (Spire height)
NorthAmerica
Europe
Continent
North America
Europe
Wilks’ Lambda: 0.867
p < 0.001 ***
signiﬁcant inﬂuence: SP

Appendix
Further Reading
Introduction
Methods
Results
North coasts – South coasts
10
9
8
7
6
5
4
3
2
Canonical2
-7 -6 -5 -4 -3 -2 -1 0 1 2 3 4 5
Canonical 1
+CL (Columellar length)
-LA (Length of aperture
south
north
Side
north
south
p < 0.001 ***
signiﬁcant inﬂuence: CL > LA

Appendix
Further Reading
Introduction
Methods
Results
Within north coasts
12
11
10
9
8
7
6
5
4
3
Canonical2
-SP(Spireheight)
-CL(Columellarlength)
-7 -6 -5 -4 -3 -2 -1 0
Canonical 1
+SP (Spire height)
-CL (Columellar length)
Wangerooge
Borkum
Norderney
Place
Borkum
Wangerooge
p < 0.001 ***
signiﬁcant inﬂuence: SP > CL
Norderney

Appendix
Further Reading
Introduction
Methods
Results
Within south coasts
6
5
4
3
2
1
0
-1
-2
Canonical2
-CL(Columellarlength)
+SW(Shellwidth)
-LA(Lengthofaperture)
-SP(Spireheight)
7 8 9 10 11 12 13 14 15 16 17 18 19
Canonical 1
+CL (Spire height)
-SW (Shell width)
+LA (Length of aperture)
+SP (Spire height)
Norderney
Langeoog
Wangerooge
Borkum
Place
Borkum
Langeoog
Norderney
Wangerooge
p < 0.001 ***
signiﬁcant inﬂuence: CL > SW > LA > SP

Appendix
Further Reading
Introduction
Methods
Results
Sexes
4
3
2
1
0
-1
-2
-3
-4
-5
Canonical2
3 4 5 6 7 8 9 10 11 12
Canonical 1
male
female
Sex
male
female
p < 0.277

Appendix
Further Reading
Introduction
Methods
Results
Genetic (Dest) and Geographic (km) Distance
B2 W1 N2 WH
B2 . . . 0.011 -0.007 0.033 *
W1 82.5 . . . 0.006 0.056 **
N2 38.0 39.9 . . . 0.057 *
WH 5662.8 5730.7 5692.3 . . .

Appendix
Further Reading
Introduction
Methods
Results
B2 W1 N2 WH
B2 . . . 0.011 -0.007 0.033 *
W1 82.5 . . . 0.006 0.056 **
N2 38.0 39.9 . . . 0.057 *
WH 5662.8 5730.7 5692.3 . . .
Borkum
Norderney
Langeoog
Wangerooge
Within the EFWS

Appendix
Further Reading
Introduction
Methods
Results
B2 W1 N2 WH
B2 . . . 0.011 -0.007 0.033 *
W1 82.5 . . . 0.006 0.056 **
N2 38.0 39.9 . . . 0.057 *
WH 5662.8 5730.7 5692.3 . . .
Woods Hole
EFWS
Between the EFWS and Woods Hole

Appendix
Further Reading
Introduction
Methods
Results
Wadden Sea
Populations
are
not differentiated
at
neutral loci

Population dynamics of the periwinkle Littorina littorea (Linnaeus, 1758) in the East Frisian Wadden Sea

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Population dynamics of the periwinkle Littorina littorea (Linnaeus, 1758) in the East Frisian Wadden Sea