Natural Climatic Variations Induce Cohort Effect in Lacerta vivipara

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Cohort variation, climate effects and population dynamics in a short-lived lizard
1
APPENDIX A – CLIMATE VARIATION
2
Temperature and rainfall show marked variations among cohorts (Figure A1). Only temperature
3
during June showed a significant temporal increase (F1,15 = 16.94, p = 0.0009, + 0.29 °C ± 0.07 s.e. per
4
year, see Figure A1). The temporal autocorrelation coefficients of the time series for temperature and
5
rainfall were weak and non significant (p > 0.05). Furthermore, correlations of temperature and
6
rainfall data between life stages within a given birth cohort were usually small (Table A1).
7
8
24
Figure A1. Climate conditions were
10
calculated for each birth cohort at
11
distinct life stages of the common lizard
12
as defined in Figure 1. For the ease of
13
visualization, we represent here only
14
the data for mid-gestation and early
15
juvenile stages. (A) Means of maximum
16
daily
17
exceptionally warm summer for the
18
cohort born in 2003 and the warming
19
trend
20
Cumulative rainfall for birth cohorts
(A)
22
Mean temperature (°C)
9
20
18
16
14
12
Mid-gestation stage
Early juvenile stage
10
8
1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006
Birth cohort
temperatures;
note
the
350
(B)
during
mid-gestation.
(B)
21
from 1988 to 2004; note the large
22
temporal variation in rainfall during
23
mid-gestation.
Cumulative rainfall (mm)
300
250
200
150
100
50
0
1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006
Birth cohort
24
25
1
Cohort variation, climate effects and population dynamics in a short-lived lizard
26
Table A1. Pairwise correlations of temperature (TP) and rainfall (RF) data between life stages within
27
birth cohorts (upper triangular matrix: Pearson’s product moment correlation coefficient, lower
28
triangular matrix: sample size). Significant correlations are bolded. Life stages are numbered
29
according to notations defined in Figure 1: 1 = conception, 2 = mid-gestation, 3 = early juvenile, 4=
30
late juvenile, 5 = sub adult and 6 = adult. Bolded correlations are significant without correction for
31
multiple testing (p < 0.05).
32
TP1
TP1
TP2
TP3
TP4
TP5
TP6
RF1
RF2
RF3
RF4
RF5
RF6
0.153
0.377
-0.042
-0.258
0.327
0.015
0.065
-0.293
-0.194
0.052
0.290
0.354
0.458
0.462
0.181
-0.270
-0.636
0.114
-0.376
-0.380
0.207
-0.075
-0.204
-0.273
-0.279
-0.075
-0.480
-0.092
0.085
-0.156
0.679
0.176
0.302
-0.108
0.404
-0.315
-0.667
-0.240
0.337
0.085
0.197
0.464
-0.336
-0.370
-0.614
-0.105
0.276
0.041
0.195
-0.107
-0.372
0.208
-0.231
-0.275
-0.048
-0.251
-0.067
-0.104
0.078
-0.451
-0.066
-0.120
0.190
-0.036
-0.001
TP2
16
TP3
16
16
TP4
15
15
15
TP5
14
14
14
14
TP6
13
13
13
13
13
RF1
16
16
16
15
14
13
RF2
16
16
16
15
14
13
16
RF3
16
16
16
15
14
13
16
16
RF4
15
15
15
15
14
13
15
15
15
RF5
14
14
14
14
14
13
14
14
14
14
RF6
13
13
13
13
13
13
13
13
13
13
2
-0.070
13
NA
Cohort variation, climate effects and population dynamics in a short-lived lizard
33
APPENDIX B – ADULT GROWTH AND REPRODUCTIVE TRAITS
34
Table B1. Cohort variation in adult body size (55 individuals from 13 cohorts). Results are
35
from the minimum adequate mixed effects linear model. F- indicates the low-density site and
36
M indicates males.
Fixed effects
Estimate ± s.e.
F ndf,ddf
P value
Age (years)
5.36 ± 0.87
37.91,124
<.0001
Age^2
-0.35 ± 0.09
15.61,124
.0001
Sex
M: -6.78 ± 1.36
24.71,41
<.0001
Random effects
Estimate [95% CI]
LRT test
P value
Cohort identity
1.34 [0.72, 2.49]
25.09
<.0001
Female identity in cohort
1.70 [1.21, 2.39]
19.65
<.0001
Residuals
2.33 [2.06, 2.64]
37
38
39
Table B2. Cohort variation in parturition dates (453 females from 13 cohorts). Results are
40
from the minimum adequate mixed effects linear model.
Fixed effects
Estimate ± s.e.
F ndf,ddf
P value
Female SVL
-0.50 ± 0.07
48.671,128
<.0001
Current temperature
-1.33 ± 0.32
17.641,128
<.0001
Current rainfall
0.013 ± 0.005
6.151,128
0.0145
Random effects
Estimate [95% CI]
LRT test
P value
Cohort identity
2.91 [1.78, 4.76]
45.65
<.0001
Female identity in cohort
3.04 [2.27, 4.09]
11.47
<.0001
Residuals
4.67 [4.16, 5.26]
41
3
Cohort variation, climate effects and population dynamics in a short-lived lizard
42
Table B3. Cohort variation in reproductive failures (proportion of unviables eggs within a
43
clutch, 314 females from 14 cohorts). Results are from the minimum adequate mixed effects
44
generalized linear model with a logit link and binomial distribution of error terms.
Fixed effects
Estimate ± s.e.
z test
P value
Current mid-gestation rainfall
0.004 ± 0.001
2.73
0.006
Random effects
Estimate [95% CI]
LRT test
P value
Cohort identity
0
-
-
Female identity in cohort
2.33
Not available
-
Residuals
0.82
45
46
47
Table B4. Cohort variation in post-parturition body condition (body mass controlled for body
48
length, 314 females from 14 cohorts). F- indicates the low-density site.
Fixed effects
Estimate ± s.e.
F ndf,ddf
P value
Female SVL
0.066 ± 0.008
68.561,124
<.0001
Age
0.227 ± 0.006
15.451,124
.0001
Age^2
-0.025 ± 0.007
12.391,124
.0006
Habitat
F-: -1.169 ± 0.546
4.581,124
.03
SVL × Habitat
F-: 0.043 ± 0.010
5.381,124
.02
Current mid-gestation temperature 0.043 ± 0.010
18.71,124
<0.0001
Random effects
Estimate [95% CI]
LRT test
P value
Cohort identity
0.017 [0.00003, 8.08]
0.11
.74
Female identity in cohort
0.193 [0.151, 0.248]
18.47
<.0001
Residuals
0.258 [0.229, 0.291]
49
4
Cohort variation, climate effects and population dynamics in a short-lived lizard
50
APPENDIX C – STATISTICAL MODELS FOR MARK-RECAPTURE ANALYSES
51
Statistical models for juveniles
52
The recapture history file encompassed 4370 individuals and 12 birth cohorts (1989-1995, 1998-
53
2002). We excluded the 1996 and 1997 birth cohorts since sub-adults were not searched in June-July
54
1998, and included one dummy variable (habitat F+), capture effort, two individual covariate (size at
55
birth and birth date) and six climate covariates (rainfall and temperature during conception, past mid-
56
gestation and early juvenile life). Individual, climate, and capture effort covariates were standardized
57
prior to analysis, and time intervals were provided to obtain estimates of monthly survival
58
probabilities. The effect of sex was not included in this analysis since ca. 21% of hatchling born in the
59
laboratory could not be sexed at birth due to logistic failures. This should not be a major problem
60
because male and female juveniles have similar survival probabilities and sensitivities to
61
environmental conditions (Le Galliard et al. 2005, 2006).
62
63
Table C1. Mark-recapture models for age-dependence and cohort variation in capture probabilities (a)
64
and survival probabilities (b and c) of juvenile common lizards. The most parsimonious models
65
according to the AICc are indicated in bold using ΔAICc<2 as a threshold criterion. Notations used in
66
the models: Φ = survival probability, p = capture probability, a = age, a01 = early juvenile stage, a02 =
67
late juvenile stage, cohort = birth cohort, effort = capture effort (number of capture days), ct =
68
constant, linear = linear trend of birth cohort effect, climate = climate covariates (rainfall [rain] and
69
temperature [tp]) during conception [c], gestation [g] and early juvenile life [j], size = snout-vent
70
length at hatching, zone = F+ habitat (dummy variable). Notations “*” and “+” indicate additive and
71
interaction terms, respectively, while “:” indicates that the effect is restricted to a given age class.
Model number and name
AICc score
ΔAICc
AICc weight
Model likelihood
Rank
Deviance
(a) Selection of a cohort model describing recapture probabilities
1. Φa*cohort pa*cohort #
2. Φa*cohort pa+cohort
3. Φa*cohort pa*effort
6961.25
0.00
1
1
60
6839.85
6992.92
31.66
0
0
50
6891.94
7010.92
49.66
0
0
40
6930.29
5
Cohort variation, climate effects and population dynamics in a short-lived lizard
4. Φa*cohort pcohort
5. Φa*cohort pa
6. Φa*cohort pct
7024.37
63.12
0
0
48
6927.47
7028.62
67.37
0
0
38
6952.05
7051.94
90.68
0
0
37
6977.40
(b) Selection of a cohort model describing survival probabilities
7. Φa01:cohort pa*cohort
8. Φa01:linear pa*cohort
1. Φa*cohort pa*cohort
9. Φa+cohort pa*cohort
10. Φa pa*cohort
11. Φct pa*cohort
12. Φcohort pa*cohort
6955.71
0.00
0.7935
1
49
6856.77
6959.25
3.54
0.1452
0.1703
39
6880.65
6961.25
5.54
0.0496
0.0625
60
6839.85
6962.91
7.20
0.0217
0.0274
49
6863.97
6975.36
19.65
0.0000
0.0001
38
6898.80
7042.21
86.50
0.0000
0.0000
37
6967.67
7048.96
93.25
0
0
48
6952.06
(c) Selection of a covariate model describing survival probabilities
13. Φa01:(raing+rainj+size+birthd) pa*cohort
14. Φa01:(raing+rainj+birthd) pa*cohort
15. Φa01:(size+birthd) pa*cohort
16. Φa01:birthd pa*cohort
17. Φa01:(birthd+zone) pa*cohort
18. Φa01:(raing+rainj+size) pa*cohort
19. Φa01:(raing+rainj) pa*cohort
7. Φa01:cohort pa*cohort
20. Φa01:rainj pa*cohort
21. Φa01:(rainj+rainc) pa*cohort
22. Φa01:(rainj+tpc) pa*cohort
23. Φa01:(rainj+tpj) pa*cohort
24. Φa01:(rainj+tpg) pa*cohort
25. Φa01:tpj pa*cohort
26. Φa01:size pa*cohort
27. Φa01:rainc pa*cohort
28. Φa01:raing pa*cohort
6880.82
0.00
1.0000
1.0000
42
6796.13
6900.63
19.81
0.0001
0.0001
41
6817.97
6906.49
25.67
0.0000
0.0000
40
6825.86
6923.91
43.09
0.0000
0.0000
39
6845.31
6932.75
51.94
0.0000
0.0000
40
6852.13
6944.83
64.01
0.0000
0.0000
41
6862.17
6952.59
71.78
0.0000
0.0000
40
6871.97
6955.71
74.89
0.0000
0.0000
49
6856.77
6956.74
75.92
0.0000
0.0000
39
6878.14
6956.98
76.16
0.0000
0.0000
40
6876.35
6957.62
76.80
0.0000
0.0000
40
6876.99
6958.07
77.25
0.0000
0.0000
40
6877.44
6958.14
77.32
0.0000
0.0000
40
6877.51
6961.57
80.75
0.0000
0.0000
39
6882.98
6970.26
89.44
0.0000
0.0000
39
6891.66
6971.04
90.22
0.0000
0.0000
39
6892.45
6971.47
90.65
0.0000
0.0000
39
6892.88
6
Cohort variation, climate effects and population dynamics in a short-lived lizard
29. Φa01:tpg pa*cohort
30. Φa01:zone pa*cohort
31. Φa01:tpc pa*cohort
32. Φa01:raing pa*cohort
10. Φa pa*cohort
33. Φa01:rainc pa*cohort
72
6972.86
92.04
0.0000
0.0000
39
6894.26
6974.46
93.64
0.0000
0.0000
39
6895.86
6974.84
94.02
0.0000
0.0000
39
6896.24
6982.91
102.09
0.0000
0.0000
39
6904.31
6984.80
103.98
0.0000
0.0000
38
6908.23
6986.44
105.63
0.0000
0.0000
39
6907.85
# Model on which the GOF tests were performed. No evidence of lack of fit was found.
73
74
Statistical models for sub-adults and adults
75
The data set totalized 1058 histories and 11 cohorts (1997 was excluded since we did not search for
76
sub-adults in June-July 1998). Adult males, but not females, were not searched for in 1998 and we
77
therefore fixed the capture probabilities of males to zero in 1998. Fixed parameters were discounted
78
from the model selection procedure. Sexes were treated as separate groups since males and females
79
have distinct life histories (Massot et al. 1992). The analysis also included habitat (F+ or F-) and origin
80
(born in laboratory or in the field) as dummy variables, ten climate covariates (rainfall and temperature
81
during conception, past mid-gestation, early juvenile life and late juvenile life, and sub-adult life), and
82
capture effort (number of days spent in the field). Annual survival probabilities were estimated.
83
84
Table C2. Mark-recapture models for age-dependence and cohort variation in capture probabilities (a)
85
and survival probabilities (b, c and d) of sub-adult and adult common lizards. The most parsimonious
86
models according to the QAICc (over-dispersion coefficient, ĉ = 1.20) are indicated in bold using
87
ΔQAICc<2 as a threshold criterion. Notations used in the models are similar than in Table C1 with the
88
exception of age classification (a1 = sub-adult stage, a2 = age 2 years adult, a2 = age 3 years adult, a2-
89
3 = pooled age 2 and 3 years adult, jl = late juvenile life stage, s = sub-adult life stage), and origin
90
(dummy variable = born in the laboratory).
Model number and name
QAICc score
ΔQAICc
QAICc weight
Model likelihood
Rank
QDeviance
1.0000
81
1742.61
(a) Selection of a cohort model describing recapture probabilities
1. Φa*cohort*sex peffort †
1913.70
0.00
0.2176
7
Cohort variation, climate effects and population dynamics in a short-lived lizard
2. Φa*cohort*sex pa+effort †
3. Φa*cohort*sex pa*cohort †
4. Φa*cohort*sex peffort+sex †
5. Φa*cohort*sex pa †
6. Φa*cohort*sex pa*cohort+sex †
7. Φa*cohort*sex pct †
8. Φa*cohort*sex psex †
9. Φa*cohort*sex pa+sex †
10. Φa*cohort*sex pcohort+sex †
11. Φa*cohort*sex pa+cohort †
12. Φa*cohort*sex pa*sex †
13. Φa*cohort*sex pcohort †
14. Φa*cohort*sex pa+cohort+sex †
15. Φa*cohort*sex pcohort+a*sex †
16. Φa*cohort*sex pcohort*sex †
17. Φa*cohort*sex pa+cohort*sex †
18. Φa*cohort*sex pcohort*sex †
19. Φa*cohort*sex pa*cohort*sex #†
1913.80
0.10
0.2074
0.9532
83
1738.24
1914.34
0.64
0.1579
0.7253
108
1681.92
1914.59
0.89
0.1397
0.6420
82
1741.26
1916.47
2.77
0.0546
0.2509
82
1743.14
1916.48
2.78
0.0543
0.2493
109
1681.74
1916.65
2.95
0.0497
0.2285
80
1747.79
1917.42
3.71
0.0340
0.1561
81
1746.32
1917.67
3.97
0.0299
0.1372
83
1742.12
1919.25
5.55
0.0136
0.0624
90
1727.97
1919.36
5.66
0.0128
0.0590
92
1723.56
1919.69
5.99
0.0109
0.0500
85
1739.66
1919.73
6.02
0.0107
0.0492
90
1728.44
1921.33
7.63
0.0048
0.0220
93
1723.27
1923.86
10.16
0.0014
0.0062
95
1721.25
1927.11
13.41
0.0003
0.0012
100
1713.10
1927.13
13.43
0.0003
0.0012
102
1708.54
1927.24
13.54
0.0003
0.0011
100
1713.24
1940.82
27.12
0.0000
0.0000
128
1661.47
(b) Selection of a cohort model describing survival probabilities
20. Φa*cohort+sex peffort
1875.10
0.00
0.8773
1.0000
47
1778.08
21. Φa+cohort+sex peffort
1879.50
4.40
0.0973
0.1109
17
1845.10
22. Φcohort+a*sex peffort
1882.39
7.29
0.0229
0.0261
20
1841.84
23. Φa+cohort*sex peffort
1886.92
11.82
0.0024
0.0027
27
1831.92
24. Φa+sex peffort
1892.79
17.69
0.0001
0.0001
7
1878.72
25. Φa*sex peffort
1895.68
20.58
0.0000
0.0000
10
1875.54
26. Φcohort+sex peffort
1907.64
32.53
0.0000
0.0000
14
1879.36
27. Φsex peffort
1913.40
38.30
0.0000
0.0000
4
1905.37
1. Φa*cohort*sex peffort †
1913.70
38.60
0.0000
0.0000
81
1742.61
28. Φcohort*sex peffort
1917.23
42.12
0.0000
0.0000
24
1868.44
8
Cohort variation, climate effects and population dynamics in a short-lived lizard
29. Φcohort*a peffort
1976.05
100.94
0.0000
0.0000
46
1881.16
30. Φcohort+a peffort
1976.12
101.02
0.0000
0.0000
16
1943.77
31. Φcohort peffort
1981.13
106.03
0.0000
0.0000
13
1954.89
32. Φa peffort
1985.95
110.84
0.0000
0.0000
6
1973.89
33. Φct peffort
1987.10
112.00
0.0000
0.0000
3
1981.09
(c) Selection of a model describing age and cohort-dependent survival probabilities
34. Φa1:cohort+a2-3+sex peffort
35. Φa1:cohort+a2+a3+sex peffort
36. Φa1:cohort+a2:cohort+a3+sex peffort
37. Φa1:cohort*sex+a2-3+sex peffort
20. Φa:cohort+sex peffort
38. Φa1+a2-3+sex peffort
1862.41
0.00
0.6251
1.0000
16
1830.05
1864.45
2.04
0.2249
0.3598
17
1830.05
1865.36
2.95
0.1427
0.2283
27
1810.36
1871.63
9.22
0.0062
0.0100
27
1816.63
1875.10
12.70
0.0011
0.0017
47
1778.08
1890.77
28.37
0.0000
0.0000
6
1878.72
(d) Selection of a covariate model describing survival probabilities
34. Φa1:cohort+a2-3+sex peffort
1862.41
0.0000
0.3614
1
16
1830.05
35. Φa1:(cohort+zone)+a2-3+sex peffort
1863.26
0.8546
0.2357
0.6523
17
1828.86
36. Φa1:cohort+a2-3+a:zone+sex peffort
1864.16
1.7481
0.1508
0.4173
18
1827.71
37. Φa1:cohort+a2-3+zone+sex peffort
1864.41
2.0038
0.1327
0.3672
17
1830.01
38. Φa1:cohort+a2-3+a:origin+sex peffort
1864.62
2.2168
0.1193
0.3301
18
1828.18
39. Φa1:tpc+a2-3+sex peffort
1881.70
19.2904
0.0000
0.0001
7
1867.62
40. Φa1:precs+a2-3+sex peffort
1882.19
19.7781
0.0000
0.0001
7
1868.11
41. Φa1:tpj+a2-3+sex peffort
1887.48
25.0745
0.0000
0
7
1873.41
42. Φa1:rainc+a2-3+sex peffort
1890.05
27.6418
0.0000
0
7
1875.98
38. Φa1+a2-3+sex peffort
1890.77
28.3674
0.0000
0
6
1878.72
43. Φa1:raing+a2-3+sex peffort
1891.09
28.6814
0.0000
0
7
1877.02
44. Φa1:rainj+a2-3+sex peffort
1891.17
28.7604
0.0000
0
7
1877.09
45. Φa1:tpjl+a2-3+sex peffort
1891.62
29.2155
0.0000
0
7
1877.55
46. Φa1:tpg+a2-3+sex peffort
1891.96
29.5550
0.0000
0
7
1877.89
47. Φa1:rainjl+a2-3+sex peffort
1892.36
29.9538
0.0000
0
7
1878.29
48. Φa1:tps+a2-3+sex peffort
1892.73
30.3269
0.0000
0
7
1878.66
9
Cohort variation, climate effects and population dynamics in a short-lived lizard
91
# Model on which the GOF tests were performed. Evidence of a slight lack of fit was found and an over-
92
dispersion coefficient (ĉ = 1.20) was included.
93
† Models for which some of the parameters could not be estimated due to limited data; the number of identifiable
94
parameters was estimated with M-SURGE using a numerical approach (Choquet et al. 2005)
95
96
Table C3. Estimates of the mean, cohort variance and coefficient of variation for sub-adult and adult
97
survival in each sex and age class. Estimates were obtained from the best selected mark-recapture
98
model in Table C2b, which controlled for variation in capture probabilities due to capture effort.
99
Survival was calculated from age 1 to age 2 (Φ1), from age 2 to age 3 (Φ2) and from age 3 to age 4
100
(Φ3) for male and female lizards.
Φ1
Φ2
Φ3
Males
Females
Males
Females
Males
Females
0.30 ± 0.06
0.67 ± 0.06
0.15 ± 0.03
0.57 ± 0.07
0.09 ± 0.02
0.53 ± 0.08
Variance
0.023
0.026
0.002
0.023
0.0008
0.006
[95% CL]
[0.006, 0.10]
[0.009, 0.10]
[-0.006, 0.06]
[0.006, 0.10]
[-0.001, 0.015]
[-0.009, 0.09]
0.50
0.24
0.30
0.27
< 0.01
0.15
Mean (± se)
CV
101
102
References cited
103
Choquet, R., A. M. Reboulet, R. Pradel, O. Gimenez, and J. D. Lebreton. 2005. M-SURGE 1.8 User's manual.
104
105
106
107
108
109
110
CEFE, Montpellier, France.
Le Galliard, J.-F., R. Ferrière, and J. Clobert. 2005. Juvenile growth and survival under dietary restriction: are
males and females equal? Oikos 111:368-376.
Le Galliard, J.-F., M. Massot, S. Meylan, M. Landys, and J. Clobert. 2006. Ontogenic sources of variation in
sexual size dimorphism in a lizard. Journal of Evolutionary Ecology 19:690-704.
Massot, M., J. Clobert, T. Pilorge, J. Lecomte, and R. Barbault. 1992. Density dependence in the common lizard:
demographic consequences of a density manipulation. Ecology 73:1742-1756.
111
10
Cohort variation, climate effects and population dynamics in a short-lived lizard
112
APPENDIX D – COHORT COVARIATION BETWEEN LIFE HISTORY TRAITS
113
We calculated the mean and variance of the life history traits for each cohort. We included in this
114
analysis the life history traits that exhibited significant cohort variation: birth date (Date_B, time
115
interval since June 1, normal distribution), size at birth (SVL_B, mm, normal distribution), juvenile
116
growth rate (Growth_J, mm.day-1, normal distribution), early juvenile survival (Surv_J, monthly
117
survival probability, binomial distribution), sub-adult growth rate (Growth_S, mm.day-1, normal
118
distribution), sub-adult survival (Surv_S, annual survival probability, binomial distribution), total
119
clutch size (TCS, log-normal distribution) and adult body size (SVL_A, mm, normal distribution).
120
121
Table D1. Pairwise correlations between life history traits (upper triangular matrix: Pearson’s product
122
moment correlations and p values, diagonal: number of cohorts sampled for each life history trait,
123
lower triangular matrix: Spearman’s rho statistics and p values). Significant correlations are bolded
124
without correction for multiple testing.
125
Date_B
Date_B
SVL_B
Surv_J
Growth_J
Surv_S
Growth_S
TCS
SVL_A
n = 15
0.021
p = 0.939
SVL_B
Surv_J
Growth_J
Surv_S
Growth_S
TCS
SVL_A
-0.071
-0.286
0.331
-0.239
0.102
-0.329
-0.464
p = 0.800
p = 0.344
p = 0.248
p = 0.479
p = 0.740
p = 0.297
p = 0.80
0.025
-0.071
-0.035
0.076
0.361
0.414
p = 0.936
p = 0.808
p = 0.918
p = 0.805
p = 0.249
p = 0.180
0.081
0.346
-0.01
-0.19
0.213
p = 0.793
p = 0.297
p = 0.974
p = 0.576
p = 0.530
-0.058
-0.687
0.155
0.051
p = 0.867
p = 0.009
p = 0.649
p = 0.882
-0.001
0.377
-0.143
p = 0.875
p = 0.227
p = 0.659
-0.358
0.076
p = 0.254
p = 0.815
n = 15
-0.286
0.170
p = 0.344
p = 0.572
0.235
-0.288
0.120
p = 0.413
p = 0.318
p = 0.689
-0.264
0.136
0.445
-0.064
p = 0.435
p = 0.683
p = 0.168
p = 0.860
0.143
0.242
-0.093
-0.659
0.035
p = 0.636
p = 0.420
p = 0.765
p = 0.017
p = 0.912
-0.476
0.329
-0.009
0.155
0.322
-0.294
p = 0.121
p = 0.292
p = 0.989
p = 0.644
p = 0.303
p = 0.354
-0.168
0.266
-0.400
0.136
-0.266
0.021
0.626
p = 0.604
p = 0.398
p = 0.225
p = 0.683
p = 0.404
p = 0.957
p = 0.024
n = 13
n = 14
126
11
n = 12
n = 14
n = 13
0.634
p = 0.02
n = 13
Cohort variation, climate effects and population dynamics in a short-lived lizard
127
APPENDIX E – MATRIX PROJECTION MODEL
128
Table E1. Long-term mean (± s.e.) estimates from the field, growth sensitivity and growth elasticity
129
of each vital rate. Sensitivity and elasticity were calculated from an age-structured transition matrix.
130
Estimates are given on the logit scale for proportional data and on the log-normal scale for total clutch
131
size. Bold estimates give values on the natural scale.
Vital rate
Annual juvenile survival
Field estimates
27.9%
Sensitivity
0.845
Monthly early survival 0.129 (± 0.126)
0.443
Monthly late survival 2.601 (± 0.141)
2.282
Annual sub-adult survival
65.4%
Elasticity
0.244
0.361
0.244
0.512
0.268
0.099
0.244
0.635 (± 0.162)
Annual adult survival
50.6%
0.024 (± 0.108)
Fecundity
2.361
Total clutch size 1.619 (± 0.363)
0.044
Hatching success 2.380 (± 0.073)
0.258
Sex ratio (% females) -0.087 (± 0.028)
132
12
0.494
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