Article:Eumelanin-Based Coloration and Fitness Parameters in Birds: a Meta-Analysis

Article:Eumelanin-based coloration and fitness parameters in birds: a meta-analysis

Journal:Behavioural Ecology and Sociobiology

Authors: Joël Meunier,Susana Figueiredo Pinto,Reto Burri, and Alexandre Roulin. Correspondence:

Online Resource 1: Studies used in the meta-analyses. ‘r’ is the correlation coefficient between reproductive parameters and eumelanin-based coloration in birds; ‘n’ is the sample size of the study. Positive correlations indicate that the darkest individuals lay eggs earlier in the season, produce more eggs or more offspring, and have a higher survival rate, and vice versa for negatives correlations. We present in bold and italics the mean values used in our data set when several effect sizes were available for a single species. We added in the table four studies for which we could not calculate effect sizes for the analysis of survival. In these studies, the absence of covariation between eumelanin-based coloration and survival was indicated by ‘no’, and positive covariation by ‘positive’. * indicates species that are sexually dimorphic with respect to melanin-based coloration, + that species are colour polymorphic and § that species belong to the non-passeriforme order, and a versus b that the colour traits varies between individuals in its size versus colour intensity. Significant relationships are written in bold.

Reported or estimated r
Species / Males and / Males / Females / Reference
females
r / n / r / n / r / n
LAYING DATE
Anser caerulescens caerulescens+§a / 0.003 / 127851 / Cooke et al. 19951
Apus melba+§a / 0.026 / 132 / 0.010 / 129 / 0.042 / 135 / Bize pers. com.2
Charadrius alexandrinus (mean)*§a / -0.283 / 66
Charadrius alexandrinus*§a / -0.167 / 6 / Amat 20053
Charadrius alexandrinus*§a / -0.295 / 60 / Lendvai et al. 20044
Falco naumanni*§b / 0.025 / 163 / Tella et al. 19975
Ficedula hypoleuca (mean)*b / 0.134 / 321
Ficedula hypoleuca*b / 0.170 / 158 / Järvi et al. 19876
Ficedula hypoleuca*b / 0.099 / 163 / Potti and Montalvo 19917
Geospiza fortis*b / -0.060 / 50 / Grant 19908
Parus major*a / 0.207 / 86 / Norris, 19909
Passer domesticus (mean)*a / -0.325 / 59
Passer domesticus*a / -0.610 / 26 / Vaclav and Hoi 200210
Passer domesticus*a / -0.100 / 33 / Griffith et al. 199911
Phalacrocorax carbo§a / 0.160 / 100 / 0.160 / 100 / 0.160 / 100 / Childress and Bennum 200212
Stercorarius parasiticus+§a / 0.180 / 50 / Davis 197613
Sterna paradisaea§b / 0.202 / 2924 / Møller et al. 200714
Strix aluco+§b / -0.028 / 160 / -0.070 / 152 / 0.010 / 167 / Gasparini et al. unpubl.15
Tyto alba*+§a / 0.174 / 264 / 0.084 / 266 / 0.264 / 262 / Roulin unpubl.16
Vermivora chrysoptera/pinus*b / -0.220 / 11 / McKinnon and Robertson 200817
CLUTCH SIZE
Alectoris rufa§a / -0.487 / 23 / Bortolotti et al. 200618
Anser caerulescens caerulescens+§a / 0.001 / 1354 / Cooke et al. 198519
Apus melba+§a / -0.006 / 131 / -0.023 / 128 / 0.010 / 135 / Bize pers. comm.20
Charadrius alexandrinus*§a / 0.393 / 60 / Lendvai et al. 200421
Falco naumanni*§b / 0.050 / 118 / Tella et al. 199722
Falco tinnunculus*§b / 0.224 / 83 / Vergara et al. 200923
Ficedula hypoleuca*b / 0.084 / 163 / Potti and Montalvo 199124
Lanius minor*a / -0.014 / 56 / -0.171 / 60 / -0.128 / 51 / Kristin et al. 200725
Parus major*a / 0.375 / 88 / Norris, 199026
Passer domesticus (mean)*a / 0.268 / 69
Passer domesticus*a / -0.460 / 30 / Vaclav and Hoi 200227
Passer domesticus*a / -0.283 / 39 / Griffith et al. 199928
Remiz pendulinus*a / -0.006 / 24 / Kingma et al. 200829
Setophaga ruticilla*a / 0.401 / 39 / Lemon et al. 199230
Sterna paradisaea§b / 0.056 / 2404 / Møller et al. 200731
Strix aluco+§b / -0.053 / 160 / -0.200 / 152 / 0.080 / 167 / Gasparini et al. unpubl.32
Tyto alba*+§a / 0.166 / 246 / 0.100 / 249 / 0.042 / 242 / Roulin unpubl.33
BROOD SIZE
Anser caerulescens caerulescens+§a / 0.005 / 190 / Cooke et al. 198534
Apus melba+§a / 0.014 / 111 / 0.033 / 109 / -0.004 / 113 / Bize unpubl.35
Buteo buteo+§a / -0.125 / 116 / -0.178 / 121 / -0.068 / 111 / Chakarov et al. 200836
Falco naumanni*§b / 0.050 / 122 / Tella et al. 199737
Ficedula hypoleuca (mean)*b / 0.039 / 250
Ficedula hypoleuca*b / 0.044 / 160 / Potti and Montalvo 199138
Ficedula hypoleuca*b / 0.030 / 90 / Järvi et al. 198739
Geospiza conirostris*b / 0.472 / 24.3 / Grant and Grant 198740
Geospiza fortis*b / 0.233 / 50 / Grant 199041
Geothypis trichas*a / 0.075 / 41 / Thusius et al. 200142
Gymnorhina tibicen*a / 0.272 / 93 / Hughes et al. 200243
Lanius minor*a / 0.114 / 56 / 0.079 / 60 / 0.155 / 51 / Kristin et al. 200744
Passer domesticus (mean)*a / 0.016 / 154
Passer domesticus*a / 0.226 / 84 / Jensen et al. 200445
Passer domesticus*a / -0.412 / 40 / Griffith et al. 199946
Passer domesticus*a / 0.001 / 30 / Vaclav and Hoi 200247
Phalacrocorax carbo§a / 0.228 / 104 / Childress and Bennum 200248
Poecile atricaphillus*a / 0.770 / 10 / Doucet et al. 200549
Quelea quelea*+a / -0.023 / 731 / Dale 200050
Remiz pendulinus*a / -0.050 / 57 / Kingma et al. 200851
Setophaga ruticilla*a / -0.265 / 39 / Lemon et al. 199252
Stercorarius parasiticus+§a / 0.063 / 72 / Davis 197653
Strix aluco (mean)+§b / 0.048 / 295 / -0.040 / 284 / 0.130 / 305
Strix aluco+§b / -0.087 / 160 / -0.260 / 152 / 0.070 / 167 / Gasparini et al. unpubl.54
Strix aluco+§b / 0.207 / 135 / 0.212 / 132 / 0.202 / 138 / Brommer et al. 200555
Tyto alba*+§a / 0.026 / 261 / 0.029 / 262 / 0.021 / 259 / Roulin unpubl.56
SURVIVAL
Anser caerulescens caerulescens (mean)+§a / -0.012 / 15434
Anser caerulescens caerulescens+§a / -0.206 / 784 / Cooch 196157
Anser caerulescens caerulescens+§a / -0.002 / 14650 / Rockwell et al. 198558
Anser caerulescens caerulescens+§a / No / ? / Cooke et al. 199559
Falco naumanni*§b / -0.029 / 236 / Tella et al. 199760
Ficedula hypoleuca (mean)*b / 0.018 / 87
Ficedula hypoleuca*b / -0.340 / 41 / Järvi et al. 198761
Ficedula hypoleuca*b / 0.392 / 46 / Potti and Montalvo 199162
Geospiza fortis*b / -0.183 / 241 / Grant 199063
Geospiza scandens*b / -0.206 / 64 / Grant 199064
Lanius minor*a / -0.039 / 169 / Kristin et al. 200765
Passer domesticus*a / 0.308 / 46 / Griffith 200066
Passer domesticus*a / No / 184 / Bókony et al. 200867
Stercorarius parasiticus (mean)+§a / -0.024 / 2157
Stercorarius parasiticus+§a / -0.030 / 1671 / O’Donald 198368
Stercorarius parasiticus+§a / -0.004 / 486 / Philips and Furness 199869
Strix aluco+§b / -0.110 / 270 / Brommer et al. 200570
Strix aluco+§b / No / 464 / Roulin et al. 200371
Tyto alba*+a / Positive / 1742 / Roulin and Altwegg 200772

1 Mean laying date in blue and white mother snow geese. Correlation was obtained using P=0.17 from Table9.4. 2 Relationship between the degree with which the rachis of breast feathers is melanized and laying date corrected by year (2002-2006) in Alpine swifts. We calculated an average correlation based on correlations obtained for males and females. We averaged sample sizes of males and females of each year. 3Average laying date of female Kentish plovers with and without black frontal bars (Wilcoxon matched-pairs test: z=0.41, P=0.686). 4 Relationship between badge size and laying date ( = -0.393, P = 0.004) in male Kentish plovers. 5 Partial correlation between the differential amount of grey on the rump of female lesser kestrels and laying date when controlling for age (r =0.025, n = 161, P0.50). 6Relationship between the degree of blackness and laying date in male pied flycatchers (P0.05). 7 Relationship between the degree of blackness and laying date in male pied flycatchers (F1,161=1.60, P=0.21). 8 Relationship between the degree of blackness and number of fledglings in male Darwin’s finches (rs=0.06, n=50, P0.10). 9 Relationship between stripe size and breeding date (F1,85 = 3.79, P < 0.06) in male great tits. 10 Relationship between laying date and the size of the patch of black feathers on the chest of male house sparrows (rs=0.61, n=26, P =0.001). 11 Relationship between laying date and the size of the patch of black feathers on the chest of male house sparrows (linear regression: r2= 0.33, n=33 P = 0.52). 12 Relationship between laying date and cheek, neck and upper breast plumage in male and female great cormorants from Table 2 (rs=-0.16, n=100 for both males and females). 13 Average hatching date for dark and pale male arctic skuas (respectively 28 June and 3 July; data from Table 4). Separate Student’s t-tests (pale versus dark) were calculated for experienced (t19=-0.17) and inexperienced males (t27 = 1.52), and then a weighted average of r derived from these two t-tests and the average sample size was computed. Intermediately colored individuals were not considered because available data did not allow the use of an ANOVA. 14 Average laying date of arctic terns displaying different levels of blackness on the tip of their beak (F1,2922=123.60, P0.0001). 15 Relationship between the degree of melanin-based coloration and laying date in a Swiss tawny owl population, corrected by year (2005, 2006, 2007). A pooled correlation coefficient was given by the weighted mean of the correlations obtained for males and females, while sample size is the average of male’s and female’s sample sizes. Dark reddish-brown feathers contain more eumelanin pigments than pale reddish-brown feathers (Gasparini et al. 2009). 16 Relationship between the size of black spots located on the ventral body side of barn owls and laying date in breeding males (F1,264=1.89, P=0.17) and females (F1,260=19.51, P0.0001). We calculated an average correlation coefficient based on correlations obtained for males and females. We averaged sample sizes of males and females. 17 Relationship between laying date and blackness of throat patch in male golden-winged warblers (r9=0.22, P=0.51). 18 Relationship between the size of ventral black area and clutch size in male red-legged partridges (F1,21=6.52, P=0.02). 19 Mean clutch size at hatching in blue and white mother snow geese (4.18 vs. 4.23 eggs). Overall correlation was the weighted mean of correlations extracted from Student’s t-tests calculated on the raw data collected from 1973 to 1981. Sample size was averaged across the nine years (1973: 1140 mothers; 1974: 1079; 1975: 1199; 1976: 1358; 1977: 1649; 1978: 1179; 1979: 1414; 1980: 1573: 1981: 1590). 20 Relationship between the degree with which the rachis of breast feathers is melanized in Alpine swifts and clutch size corrected by year (2002-2006). We calculated an average correlation coefficient based on correlations obtained for males and females. We averaged sample sizes of males and females. 21 Relationship between badge size and clutch volume ( = 0.295, P = 0.025) in male Kentish plovers. 22Spearman’s rank correlation between the differential amount of grey on the rump of female lesser kestrels and clutch size (rs=0.05, n=119, P=0.58). 23 Relationship between rump coloration and clutch size in European kestrel females (GLMM: F1,81=4.29, n=83, P=0.041). Females showing greyer rumps laid more eggs. 24 Relationship between the degree of male blackness and clutch size in the pied flycatcher (F1,161=1.13, P=0.29). 25 Average clutch size of lesser grey shrike males and females with and without black spots on the tail (males: 5.70.7 vs. 5.80.4 eggs; females: 5.61.6 vs. 5.70.5 eggs). We calculated an average correlation coefficient based on correlations obtained from t-tests provided by the authors for males and females (males: t=-0.13, P0.9; females: t=0.9, P0.9). We averaged sample sizes of males and females. 26 Relationship between stripe size and clutch size (F1,87 = 14.25, P < 0.001) in male great tits. 27 Relationship between clutch size and the size of the patch of black feathers on the chest of male house sparrows (rs=-0.46, n=30, P=0.01). 28Relationship between clutch size and the size of the patch of black feathers on the chest of male house sparrows (linear regression: r2=-0.08, n=39 P=0.09). 29 Relationship between clutch size and the size of the black mask in male penduline tits in nests tended either by only the male or only the female (data of Table4). 30 Number of eggs laid in relation to the amount of black on the bib of adult male American redstarts (F1,37=7.07, P0.02). 31 Average clutch size of arctic terns displaying different levels of blackness on the tip of their beak. Effect size was derived from an analysis of clutch size that controlled for the independent effects of date, body mass, wing length, and year (F1,2402=7.43, P=0.0065). 32 Relationship between the degree of reddish-brown coloration and clutch size in a Swiss population of tawny owls, corrected by year (2005, 2006, 2007). A pooled correlation coefficient was given by the weighted mean of the correlations obtained for males and females, while sample size is the average of male’s and female’s sample sizes. Dark reddish-brown feathers contain more eumelanin pigments than pale reddish-brown feathers (Gasparini et al., 2009). 33Relationship between the size of black spots located on the ventral body side of barn owls and number of fledglings in breeding males (F1,247=2.49, P=0.12) and females (F1,240=0.43, P=0.52). We calculated an average correlation coefficient based on correlations obtained for males and females. We averaged sample sizes of males and females. 34 Mean brood size at fledging in blue and white mother snow geese (3.05 vs. 2.95 fledglings). Overall correlation was the weighted mean of correlations extracted from Student’s t-tests calculated on the basis of raw data collected from 1973 to 1981. Sample size was averaged across the nine years (1973: 132 mothers; 1974: 101; 1975: 205; 1976: 166; 1977: 274; 1978: 114; 1979: 226; 1980: 279: 1981: 211). 35 Relationship between the degree with which the rachis of breast feathers is melanized in Alpine swifts and brood size at fledging corrected by year (2002-2006). We calculated an average correlation coefficient based on correlations obtained for males and females. We averaged sample sizes of males and females. 36 Lifetime production of nestlings produced by dark and light of males and females common buzzards. Effect size was obtained using P-values found in the paper (males: P=0.975; females: P=0.762). 37Partial correlation between the differential amount of grey on the rump of female lesser kestrels and brood size at fledging, when controlled for year, colony size and female age (r120= 0.05, P > 0.50). 38Relationship between the degree of male blackness and brood size at fledging in the pied flycatcher. The relationship was significantly positive in 1988 only (F1,158=0.30, n=160 P=0.59). Therefore, we assumed that the non-significant relationships in 1987 and 1989 were also positive. 39Relationship between the degree of male blackness and brood size in the pied flycatcher. Spearman’s correlation (rs=-0.03, n=90, P=0.39) provided by the authors. 40Relationship between plumage blackness and number of young fledged in male darwin’s finches. Pearson’s correlation was derived from the P-values for three years (P0.01 for 1980, 1981 and 1982) found in Table 4. The correlation coefficient was the weighted mean of these three correlations and sample size was the averaged value of the three years. 41Relationship between plumage blackness and number of young fledged in male darwin’s finches. Pearson’s correlation was derived from the P-value found in Table 4 (P0.05). 42Relationship between the size of the black facial mask and number of young sired by male common yellowthroats (F1,39=0.22, P=0.64). Number of young is the sum of the young sired by a male in his own nest plus the number of extrapair young he sired in nests of females other than the social mate. 43Average brood size at fledging for black and white male australian magpies in 1998 (1.13 vs. 0.78 fledglings) and 1999 (1.00 vs. 1.12 fledglings). Correlations were calculated using a Student’s t-test found in the paper (t=2.691, P=0.008). 44Average brood size for male and female lesser grey shrikes with and without black spots on the tail (males: 5.2 vs. 4.9 nestlings; females: 5.7 vs. 5.5. nestlings). Correlation coefficients were obtained using Student’s t-tests provided in the paper for both males and females (males: t58=0.6, P0.5; females: t49=1.1, P0.2). We averaged sample sizes of males and females. 45Relationship between lifetime production of recruits and the size of the patch of black feathers on the chest of male house sparrows. Reproductive success was based on molecular paternity analyses (z=2.07, n=84, P=0.038). 46Relationship between annual number of fledglings and the size of the patch of black feathers on the chest of male house sparrows (linear regression: r2=0.17, n=40, P0.01). 47Relationship between annual number of fledglings and the size of the patch of black feathers on the chest of male house sparrows (rs=0.001, n=30, P=0.99). 48Average brood size for male great cormorants displaying a dark vs. pale cheek, neck and upper breast plumage (2.9 vs. 1.9 fledglings; Mann-Whitney U-test, P = 0.01). 49Relationship between apparent reproductive success (estimated with paternity analysis of birds captured in winter flocks) and the size of the black bib displayed by male black-capped chickadees (r8=0.77, P=0.008). 50Average number of nestlings for male red-billed queleas displaying a white or black mask (2.50 vs. 2.45 nestlings; Student’s t-test: t729=0.63, P=0.53). 51Relationship between annual number of nestlings and the size of the black mask in male penduline tits. Correlation coefficient was calculated using data provided in the Appendix 1. 52Number of young fledged in relation to the amount of black on the bibs of adult male American redstarts (F1,37=2.8, P=0.10). 53Average brood size at fledging of dark and pale male arctic skuas (respectively 1.04 and 0.73 fledglings; data from Table 4). Separate Student’s t-tests were calculated for experienced (t25=0.14) and inexperienced males (t43=0.55), and then a weighted average of r derived from these t-tests and the average sample size were computed. Intermediately colored individuals were not considered because we could not perform an ANOVA using the data found in Table 4. 54Relationship between the degree of reddish-brown coloration and brood size at fledging in a Swiss population of tawny owls, corrected by year (2005, 2006, 2007). A pooled correlation coefficient was given by the weighted mean of the correlations obtained for males and females, while sample size is the average of male’s and female’s sample sizes. Dark reddish-brown feathers contain more eumelanin pigments than pale reddish-brown feathers (Gasparini et al. 2009). 55Relationship between the degree of reddish-brown coloration and lifetime production of fledglings in a Finish population of tawny owls. A pooled correlation coefficient was given by the weighted mean of the correlations obtained for males and females, while sample size is the average of male’s and female’s sample sizes. Dark reddish-brown feathers contain more eumelanin pigments than pale reddish-brown feathers (Gasparini et al. 2009). 56Relationship between the size of black spots located on the ventral body size of barn owls and number of fledglings in breeding males (F1,260=0.23, P=0.63) and females (F1,257=0.12, P=0.73). Separate analyses were carried out for males and females. We calculated an average correlation coefficient based on correlations obtained for males and females. We averaged sample sizes of males and females. 57Mean survival probability (percentage of bands recovered) between 1952 and 1958 of blue and white snow geese. Correlation was derived from a chi-square test performed using data found in Table5. 58Mean survival probability (capture-recapture model) between 1976 and 1981, calculated using Tanner estimate, of blue and white snow geese. Correlation was derived from a chi-square test performed using data found in Table4. 59Survival rates of snow geese calculated from band recovery data. 60Relationship between the differential amount of grey on the rump of female lesser kestrels and their survival from one year to the next (percentage of birds resighted). Correlation was derived from the P-value (P=0.33) of a chi-square test. 61Relationship between the degree of blackness and survival probability (percentage of birds resighted) in male pied flycatchers. The correlation is an average of relationship between the degree of male blackness and survival controlled by body weight (r=-0.35, P0.05) or physical condition (r=-0.33, P0.05). Sample size was averaged. 62Relationship between the degree of blackness and survival probability (percentage of birds resigned) in male pied flycatchers (2=2.49, P0.10). 63Relationship between plumage blackness and survival probability (selection analysis) of male Darwin finches (Geospiza fortis) in two different years. We averaged the Pearson’s correlation derived from the approximate P-values found in Table 2 (1983: P0.05, 1987: P0.01) and summed sample sizes. 64Relationship between plumage blackness and survival probability (selection analysis) of male Darwin finches (Geospiza scandens). Pearson’s correlation was derived from the P-value found in Table 2 (P0.05). 65Survival probability (percentage of birds resighted) of male and female lesser grey shrikes with and without black spots on the tail. We calculated an average correlation coefficient using binomial test provided by the author (males: z=0.51, P0.4; females: z=0.19, P0.3). Survival probability (percentage of birds resighted) of male and female lesser grey shrikes with and without black spots on the tail. We calculated an average correlation coefficient using binomial test provided by the author (males: z=0.51, P0.4; females: z=0.19, P0.3). 66Relationship between badge size and survival probability (percentage of birds resighted) in male house sparrows. We derived Pearson’s correlation from a Student’s t-test between mean badge size of survivors (34.65 mm) and of non-survivors (31.52 mm) (t=2.24, P=0.03). 67Mark capture-recapture analyses on 184 house sparrow males. 68Survival probability of pale and dark Arctic skuas. Correlation was calculated using data found in Table 6.3. We performed a chi-square test testing the difference of percentage of pale and dark individuals that survived in the periods 1948-62, 1973-75 and 1976-78 (2=1.480, df=2, P=0.48). 69Survival probability of pale and dark Arctic skuas. Correlation was derived by the chi-square tests provided by the authors for the three periods 1993-1994 (2=0.02), 1994-1995 (2=0.00) and 1995-1996 (2=0.00). We averaged the correlations obtained by these chi-square tests and summed sample sizes. 70Spearman’s correlation of the relationship between the degree of reddish-brown coloration and breeding lifespan in a Finish population of male and female tawny owls (rs=-0.11, P=0.07). Dark reddish-brown feathers contain more eumelanin pigments than pale reddish-brown feathers (Gasparini et al. 2009). 71Mark capture-recapture analyses on 464 tawny owl females. 72Mark capture-recapture analyses on 1742 barn owl individuals.