Heredity 59 (1987) 181—197 The Genetical Society of Great Britain Received 26 October 1986
Natural selection and the heritability of fitness components
Timothy A. Mousseau and Department of Biology, McGill University, Derek A. Roff 1205 Avenue Dr.Penfield, Montreal, Quebec, CanadaH3A 1B1.
The hypothesis that traits closely associated with fitness will generally possess lower heritabilities than traits more loosely connected with fitness is tested using 1120 narrow sense heritability estimates for wild, outbred animal populations, collected from the published record. Our results indicate that life history traits generally possess lower heritabilities than morphological traits, and that the means, medians, and cumulative frequency distributions of behavioural and physiological traits are intermediate between life history and morphological traits. These findings are consistent with popular interpretations of Fisher's (1930, 1958) Fundamental Theorem of Natural Selection, and Falconer (1960, 1981), but also indicate that high heritabilities are maintained within natural populations even for traits believed to he under strong selection. It is also found that the heritability of morphological traits is significantly lower for ectotherms than it is for endotherms which may in part be a result of the strong correlation between life history and body size for many ectotherms.
INTRODUCTION sive compilation of data in support of this view is that presented in table 10.1 of Falconer (1981). Thefundamental theorem of natural selection However, these data are rather few and derived states: "The rate of increase in fitness of any organ- primarily from domestic animals, which may be ism at any time is equal to its genetic variance in partly inbred. Extension to wild, outbred organ- fitness at that time" (Fisher, 1930). Fisher's isms may be erroneous. theorem is axiomatic to much of current evolution- It is possible that some significant amount of ary research. It has been variously interpreted but additive genetic variance can be maintained within is generally construed to imply that traits that have natural populations, even for characters tightly been closely and consistently associated with connected to fitness; possible mechanisms include fitness will exhibit low additive genetic variances mutation (Lande, 1976; Turelli, 1984), hetero- as a result of natural selection (e.g.,Hegmannand zygote advantage (Falconer, 1981), frequency Dingle, 1982; Lynch and Sulzbach, 1984; Riddel dependence (Bulmer, 1980), fluctuating environ- etcii., 1981).However, the validity of the funda- ments (Ewing, 1979) and migration (Felsenstein, mental theorem is dependent upon many assump- 1976). In addition, zero additive genetic variance tions that may not usually be met by natural in fitness is consistent with positive heritability populations (e.g.,populationequilibrium, weak estimates of fitness components when there exists selection, constancy of genotypic fitnesses over negative genetic correlations between components time, and independence of genotypic frequencies) (Rose and Charlesworth, 1981). (Charlesworth, 1987). The maintenance of low It is in response to the importance the views additive genetic variance has also been inferred to of Fisher (1930, 1958) and Falconer (1960, 1981) imply low heritability (in the narrow sense); "On have reached in the ecological literature, and the the whole, characters with the lowest heritabilities concurrent lack of rigorously examined empirical are those most closely connected with fitness, while support for these views, that we have undertaken characters with the highest heritabilities are those the compilation, analysis, and interpretation of the that might be judged on biological grounds to be extensive quantity of data concerning genetic the least important as determinants of natural variation within natural populations now available selection." (Falconer, 1960, 1981.) The most exten- from the published record. 182 T. A. MOUSSEAU AND D. A. ROFF
In this paper we address two questions: (a) For the purposes of analysis we define four What is the heritability of traits closely associated categories of traits: (a) behavioural traits, (b) life with fitness? Do traits that are directly connected history traits, (c) morphological traits, and (d) to fitness tend to have lower heritabilities than physiological traits. We have restricted the term traits more loosely associated with fitness as pro- "life history trait" to those characters that are posed by Falconer (1960, 1981) and suggested by directly and invariably connected to fitness. Some Fisher (1930, 1958)? If we find this to be not the examples of life history traits are fecundity, viabil- case a serious reevaluation of current evolutionary ity, survival, and development rate. Morphological thinking would be necessary. (b) Do patterns of traits include body size, wing size, and other metric genetic variability change from taxa to taxa? If so, characters. Behavioural and physiological traits this might suggest phylogenetically determined are much less widely reported than life history or constraints upon rates of evolution. morphological traits. Examples of behavioural traits are alarm reaction, activity level, and sensi- tivity to conditioning. Physiological traits include Thedata set oxygen consumption, resistance to heat stress, and body temperature. Thedata set comprises 1120 narrow sense heritabil- The composition of the data set, divided ity estimates collected from 140 sources, represent- according to trait type and taxonomic assignment ing 75species.Heritability estimates for the is presented in fig. 1. A total of 341 life history Drosophila genus were excluded from the present traits, 570 morphological traits, 104 physiological study as they are treated elsewhere (Rofi and traits, and 105 behavioural traits are represented Mousseau, 1987). The only prerequisite for within the data set. Alternatively, there are 414 inclusion in the data set was that the heritability heritability estimates for invertebrates, 460 esti- estimate be derived from a wild, outbred stock; so mates for ectothermic vertebrates(reptiles, as to avoid biases potentially introduced through amphibians, and fishes), and 246 estimates for our literature reviewing process, the data were in endothermic vertebrates (birds and mammals). no other way screened or selected. This compila- These taxonomic designations reflect divisions tion is the most extensive of its kind but is not between groupings of biological and ecological exhaustive. A list of the sources, divided according significance, and an attempt to maintain statisti- to species, is given in an appendix. cally sufficient sample sizes within each grouping.
300
C') w250 I- 200 I- C', w150 0LL 100 uJ 50 z 0 B L B L M P B L M p INVERTEBRATES ECTOTHERMIC ENDOTHERMIC VERTEBRATES VERTEBRATES
Figure 1The number of reported heritability estimates grouped according to trait type and taxonomic assignment. HERITABILITY OF FITNESS COMPONENTS 183
The analysis The relationship between pairs of estimates can be described by the following linear regression I. Methods of heritability estimation equation: Areestimates of additive genetic variances seriously biased by the methods of calculation? It FS=072 (±0.15)P0+0•23 (±0.09), r=065, is widely accepted that published estimates of MSE=0.064, (=4.74, p<0.0001 quantitative genetic parameters are seldom free from biases introduced by inbreeding, assortative where FS and P0 are the estimates derived from the full-sib and parent-offspring methods, respec- mating, dominance, maternal effects, non-additive tively (fig. 2). Since we are concerned with the genetic variances, and non-random sampling of functional relationship between heritability esti- genotypes from natural populations etc. (Mitchell- Giventhe mates generated using the full-sib and parent- Olds and Rutledge, 1986). offspring methods, and the errors of estimation are heterogeneous nature of our data set, we com- roughly comparable for both methods, it is mence this investigation with a comparison of appropriate that we determine the slope of the heritability estimates generated using different methods of calculation. The outcome of this functional regression using Ricker's (1973) examination will dictate whether it is necessary to formula: include the method of estimation as a factor in the v=b/r analyses to follow. The most commonly employed methods of where v is the slope of the functional regression, heritability estimation within our data set are the b is the slope obtained using an ordinary linear and the full- regression, and r is the correlation coefficient of parent-offspring regression (n =341) the relationship. Ricker's (1973) formula is used sib correlation (n =463)designs. The realised (n = because simple least-squares linear regressions will 133)and half-sib (n =176)designs are much less frequently utilized and as a result, insufficiently tend to seriously underestimate the slope (when r represented for the analyses to follow. is <1) if the abscissal (X) observations are subject to natural variability of the same magnitude as the Theory (Falconer, 1981) predicts that within a Y observations. We find that once corrected, the given population, for a given trait, estimates gener- ated using the full-sib design will usually be higher slope describing the functional relationship or equal to those produced using the parent- between full-sib and parent-offspring estimates offspring method. This results primarily from the equals 110, which is not significantly different covariance of full sibs being inflated by environ- from unity. mental variance between families and by domin- Even if the lack of statistical significance found ance variance (Falconer, 1981). between estimates generated using different The simplest and most convincing manner in methods of calculation were primarily due to which to approach this question is to compare insufficient data, the bias that might be introduced heritability estimates calculated using different is sufficiently small (<10 per cent) to justify the methods for the same character within a given pooling of full-sib and parent-offspring estimates. population. This approach avoids problems related In any case, the various estimates (full-sib, half-sib, to the sensitivity of heritability estimates to parent-offspring, and realised) were roughly evenly differences between populations in environmental represented in the different trait groupings. conditions, past and present (Falconer, 1981). Heritability estimates are frequently reported Table 1 list 33 paired heritability estimates for without standard errors; only 738 of the 1120 esti- a variety of characters, obtained from 11 indepen- mates collected for this study included standard dent studies, for 9 species. We found that on errors. We found the relationship between the stan- average, the full-sib design produced estimates that dard error and the heritability estimate to be: were greater by 0073 than those obtained using SE =0 172 (±0.014)h2+0.088 (±0.007), the parent-offspring regression method. In addi- tion, full-sib estimates were higher than parent- r=041, offspring estimates for 20 of the 33 cases. However, MSE =014, the results of a paired comparison (-test, a t=12, p
Table I Within study comparisons of heritability estimates calculated using the parent-offspring (P0) and full-sib (FS) methods of analysis. Paired comparison i-test: D = 0073, S.E. = 0046, 1 = 171, p = 010. Wilcoxon's signed rank test: sum of negative ranks= —193, sum of positive ranks 368, p = 0120. Sign test: p = 015
Species Character FS P0 FS — P0 Reference
invertebrates
Apis rnellifera chill coma 037 015 022 Hillesheim, 1984 2 consumption 041 013 028 Hillesheim, 1984 Eurytemora affinis temperature 020 Oil 009 Bradley, 1982 tolerance 080 076 003 Bradley, 1982 Euryiemora herdmani length 048 012 036 McLaren, 1976 length 065 054 011 McLaren, 1976 Gryllusfirmus wing length 064 074 —01 Roff, 1986b wing length 062 040 022 Rotf, 1986b Pseudocalanus length 092 098 —006 McLaren and Corkett,1978 Tribolium castaneum fecundity 038 036 002 Orozco, 1976 fecundity 035 030 005 Orozco, 1976
Amphibians Plethodon cinereus * of vertebrae 057 061 —004 Highton, 1960
Birds
Geospizafortis weight 078 091 —013 Boag, 1983 wing length 042 084 —042 Boag, 1983 tarsus length 032 071 —039 Boag, 1983 bill length i1i 065 046 Boag, 1983 bill depth 112 079 033 Boag, 1983 bill width 099 090 009 Boag, 1983 bill length 064 035 029 Boag, 1983 weight 080 091 —011 Boag and Grant, 1978 wing length 065 063 002 Boag and Grant, 1978 tarsus length 027 042 —015 Boag and Grant, 1978 bill length 134 085 049 Boag and Grant, 1978 bill depth 141 090 051 Boag and Grant, 1978 bill width 099 103 —004 Boag and Grant, 1978 bill length 078 041 037 Boag and Grant, 1978 Geospiza scandens weight 035 058 —023 Boag, 1983 wing length 035 012 023 Boag, 1983 tarsus length 088 092 —004 Boag, 1983 bill length 030 032 —002 Boag, 1983 bill depth 056 014 042 Boag, 1983 bill width 056 034 022 Boag, 1983 bill length 006 060 —054 Boag, 1983 associated with estimates generated using the full- will have low heritabilities while more distantly sib and parent-offspring methods of calculation related traits should have higher heritabilities. If (F =17,p > O2, n =659). this is indeed the case for natural populations we The above results suggest that there is no would expect life history traits to possess, on apparent bias introduced to the data set by the average, lower heritabilities than morphological method of heritability estimation; Roff and traits. It is more difficult to predict a priori the Mousseau (1987) obtained a similar result for relationship between fitness and behavioural and Drosophila when comparing parent-offspring esti- physiological traits. Falconer's compilation (1981, mates with half-sib and realised estimates. The table 10.1, p. 150) would suggest that physiological method of heritability estimation will not be con- traits are intermediate between life history traits sidered further in this study. and morphological traits with respect to average heritabilities. It has been proposed that many behavioural traits will be under strong stabilising II.Heritability and fitness selection (Lee and Parsons, 1968), and as a result, Classicaltheory (e.g., Falconer, 1960; Fisher, 1930) possess low heritabilities. It seems only reasonable predicts that traits closely associated with fitness to suppose that traits such as food conversion HERITABILITY OF FITNESS COMPONENTS 185
The entire data set Atthis resolution, our analysis is directed towards the illumination of gross patterns of variation. We ignore potential biases introduced via the over- representation of certain species and characters, -J and the effect of outliers. -J Figure 3 shows the cumulative distribution U- frequencies of the four trait groupings; table 2 lists descriptive statistics and the results of Kolmogorov-Smirnov tests upon paired com- parisons of each of the cumulative distributions. 0 0.3 0.6 0.9 1. 2 The results indicate that life history traits are gen- PARENT-OFFSPRING erally much lower than morphological traits, and Figure 2 Ascatterplot showing the relationship between that behavioural and physiological traits tend to paired heritability estimates generated using full-sib corre- fall in the middle. This pattern is consistent with lation and parent-offspring regression methods of estima- the observed means, medians, and cumulative dis- tion for the same population (r=O62).Theslope of the tribution frequencies, and is supported by the plotted line represents a 1: 1 correspondence. results of a one-way ANOVA (F =39.7,p < 0.0001, df= 1119) and a Kruskal-Wallis test (H = efficiency and mating propensity are more closely 106,p
>- phology:H=394,p>0l3,n=l42;F=283,p> z0 006, df= 140). w a w Randomsampling of the data set LI- Forthis analysis, as an alternative to using median Ui > values for a character, a representative estimate I- was selected at random. For example, a total of -i ten heritability estimates for age at maturity were collected for the rainbow trout Salmo gairdneri. Of 0D those 10 estimates, one was randomly chosen to represent this character, for this species. This pro- —0.2 0.15 0.5 0.85 12 HERITABILITY ESTIMATES cess was repeated for all characters and species with multiple estimates to generate a randomly Figure 4 The cumulative frequency distributions of the four selected subset of the larger data set. The entire trait categories (L =lifehistory, B behaviour, P = process was repeated to provide 25 such subsets physiology,and M =morphology)for median heritability values for each character of each species. Note that data and all tests were performed individually upon points are joined by straight lines. each subset. As with the use of medians, this approach eliminates biases due to the over- representation of particular characters and species. The results of this analysis are qualitatively similar We found that generally, the results obtained to those obtained for the larger data set; the rank- from the random sampling of the data set were ings of the means, medians, and cumulative distri- statistically similar to those found using median butions are identical. The main difference resulting values (table 4). In only one out of 150 comparisons from the use of medians is the lack of significance did the results contradict those obtained using found by the Kolmogorov-Smirnov test upon com- medians. Overall, we conclude that the pattern of parisons between the cumulative distributions of trait rankings derived from the median data set is lifehistorytraitsand physiological and robust to the variance contained within the larger behavioural traits. Given that the rankings and data set. absolute values are very similar to those obtained for the larger data set, we conclude that this lack Thepaired comparison of fitness and non- of significance is primarily due to the drastically fitness traits reduced sample sizes and the conservative nature of the Kolmogorov-Smirnov test. The results of a Theobjective of this analysis was to make com- Kruskal-Wallis test and a one-way ANOVA indi- parisons of traits independently of the species and cate that trait is indeed an important factor govern- study from which they were derived. To meet this ing the patterns of variance within the median data objective we have collected data from 30 sources set (H 474,p Table 2 Summary of descriptive statistics and Kolmogorov—Smirnov tests (D,,a,)forall data Life history Physiology Behaviour Morphology n 341 104 105 570 0262 0330 0302 0461 SE. 0012 0027 0023 0004 median 0250 0262 0280 0428 Dmax Physiology 0199 p Table 3Summary of descriptive statistics and Kolmogorov—Smirnov tests (Dma,) for character medians Life history Physiology Behaviour Morphology n 79 39 25 140 0265 0308 0374 0511 SE. 0028 O•040 0050 0021 median 0260 0270 0320 0530 Dm= Physiology 0194 p>028 Behaviour 0298 0199 p>O19 p>09 Morphology 0562 0373 0366 p Table 4 Summary of the results obtained from Kolmogorov-Smirnov tests upon comparisons of trait types within randomly generated data sets. The number given represents the number of tests (out of 25) which fell into a given significance level. For example, comparisons between life history and morphological traits yielded a Dma, which was significant at p <00001 for 25 of the 25 randomly generated data sets Life history vs. Morphology vs. Behaviour vs. Morphology Physiology Behaviour Behaviour Physiology Physiology N.S. — 25 24 — — 25 p Table 5 Within study comparison of traits. Estimates represent median values for trait. Sample sizes are given in parentheses. Paired comparison t-test: D =—015,SE. =0061,1-= —23,p =0038.Wilcoxon's signed rank test: sum of negative ranks =—88, sum of positive ranks =17;p =0012.Sign test: p =0029 Species Life history Morphology L —M Reference Crassostrea gigas 0200 (2) 0325 (14) —0125 Lannan, 1972 (oyster) Dysdercus bimaculatus 0193 (6) 0515 (1) —0372 Derr, 1980 (cotton stainer bug) Eurytemora herdmani 0115 (12) 0615 (8) —0500 McLaren, 1976 (copepod) Gambusia affinis 0285 (2) 0745 (2) —0460 Busack and Gall, 1983 (mosquitofish) Homarus americanus 0.200 (24) 0360 (20) —0160 Fairful et aL, 1981 (American lobster) Oncopeltusfasciatus 0240 (9) 0375 (2) —0135 Hegmann and Dingle, 1982 (milkweed bug) Pseudocalanus 0385 (6) 0458 (2) —0073 McLaren and Corkett, 1978 (copepod) Salmo gairdneri 0230 (1) 0405 (2) —0175 Gall and Gross, 1978a, b 0180(1) 0170(3) +0010 Gjerde and Gjedrem, 1984 0090(1) 0400 (3) —0310 Gjerde and Gjedrem, 1984 Salmo salar 0420(1) 0440 (3) —0020 Gjerde and Gjedrem, 1984 (Atlantic salmon) 0150(1) 0540 (3) —0-390 Gjerde and Gjedrem, 1984 1.000(1) 0550 (2) +0450 Naevdal et a!., 1976 0670 (1) 0-590 (2) +0080 Naevdal et al., 1976 neither the mean nor the cumulative distribution However, the estimated means for each category of life history traits were significantly different of trait differed dramatically. The mean (± 1 S.E.) between ectotherms and endotherms (fig. 5; t-test heritability values for morphological, life history, =—1-63,p >01, df= 345; Kolmogorov-Smirnov and behavioural traits in Drosophila (Roll and Dmax= 037,p>O.O5), although the medians were Mousseau 1987) were 0-32 (±0-02), 012 (±0.02), found to be significantly different (Mann-Whitney and 0-18 (±0.03) as opposed to 0-46 (±0.004), U-test: z=281,p<0005, nI =333, n2= 14). The 026 (±0.01), and 030 (±0.03) for animals in gen- statistical analysis of comparisons between inverte- eral (excluding Drosophila). This difference is con- brates and vertebrates (fig. 5) yielded insignificant sistent with the results we obtained from our results for both life history traits (t-test t = comparison of ectothermic and endothermic 0-76,p >04, df= 345; Mann-Whitney U-test z= organisms (see table 6), and may be related to the —1l6,p>O25,nI =211, n2= 136; Kolmogorov- close association between life history traits and Smirnov Dma,, =0i5,p >0.05), and morpho- body size in many ectotherms (Kusano, 1982; logical traits (t-testt =1-2,p> 02, df= 578; Peters, 1983; Roff, 1981, 1984, 1986a). Mann-Whitney U-test z= —17,p >0-i, ni = Our results show that behavioural and physio- 108,n2 =472; Kolmogorov-Smirnov max= logical traits have heritabilities more like those of 014,p >0-05). life history traits than those of morphological traits. One interpretation of this finding might be that physiological and behavioural traits are subject to DISCUSSION constraints similar to those thought to influence fitness traits. In addition, this discovery supports Thenotion that traits closely associated with fitness Falconer's (1981) suggestion that physiological will generally posses lower heritabilities than more traits generally possess heritabilities intermediate distantly related traits, is supported by all segments between life history and morphological traits, and of our analysis. Our findings extend further support Lee and Parsons' (1968) contention that to the views of Fisher (1930, 1958) and Falconer behavioural traits will usually be subject to stabilis- (1960, 1981) concerning the nature of genetic vari- ing selection. However, the magnitude of the ation within natural populations. average heritabilities of these traits suggests that The ranking of trait types that we obtained for significant genetic variance is maintained within animals in general is consistent with that reported most natural populations, even for traits closely for Drosophila (RofT and Mousseau, 1987). affiliated with fitness. HERITABILITY OF FITNESS COMPONENTS 189 LIFE HISTORY MORPHOLOGY 1.0 0.8 >- 0.6 z0 w0.4 0 w0.2 0.0 w > I— -J 0 —0.2 0.1 0.4 0.7 1.0 1.3 —0.4 0 0.4 0.8 .1.2 HERITABILITY ESTIMATES Figure 5A comparison of the cumulative frequency distributions of ectotherms versus endotherms and vertebrates versus inverte- brates for the heritability of life history and morphological traits. Note that data points are joined by straight lines. Table 6 A comparison of the relative frequencies of heritability estimates. The ectotherm and endotherm data are from this study and are based upon the median by trait data set. The Drosophila data are derived from Roff and Mousseau (1987) and is based upon the median by study data set of Heritability Estimate estimates <20% 20-40%40_60% 60-80%>80% Behaviour Drosophila n=38 684 184 79 53 0 Ectotherms n =16 13 38 25 25 0 Endotherms n =9 44 22 22 0 11 Life history Drosophila n =20 80 20 0 0 0 Ectotherms n =71 394 479 85 28 14 Endotherms n =8 0 50 375 125 0 Morphology Drosophila n=67 164 582 254 0 0 Ectotherms n =88 193 261 239 159 148 Endotherms n=52 135 173 231 288 173 190 T. A. MOUSSEAU AND D. A. ROFF There are many factors which may be respon- AYRES, F.ANDARNOLD, s.1983. Behavioural variation in sible for the maintenance of genetic variance of natural populations. IV. Mendelian models and heritability fitness characters, The rate of origin of variation of a feeding response in the garter snake, Thamnophis elegans. Heredity, 51, 405—413. by mutation alone may be sufficient to maintain BEATSON.R.1976.Environmentaland genetic correlates of substantial additive genetic variance within natural disruptive coloration in the water snake. Evolution, 30, (Lande, 1976; Turelli,1984). 241—25 2. populations BELL, G. l984a. Measuring the cost of reproduction. 1. The Heterozygote advantage(Falconer, 1981), correlation structure of the life table of a plankton rotifer. frequency dependent selection (Bulmer, 1980), Evolution, 38, 300—313. variable selection in heterogeneous environments BELL, 0. 1984b. Measuring the Cost of reproduction. TI. The (Ewing, 1979), diversifying selection (Thoday, correlation structure of the life tables of five freshwater 1972), and migration (Felsenstein, 1976) have been invertebrates. Evolution, 38, 3 14—326. BLANC, J. 54., CHEVASSLJS. B. AND BERGOT, p.1979.Deter- proposed as possible mechanisms for the susten- minisme genetique du nombre de coeea pyloriques chez Ia ance of genetic variation. Also, significant Truite faria (Salmo Irutta, Linne) et Ia truite arcenciel. Ill. heritabilities for fitness characters are not incon- Effect du genotype et de Ia taille des oeufs sur Ia realisation sistent with zero additive genetic variance in fitness du caractere chez Ia truite fario. Ann. Genet. Sel. Anim., 11, 93—103. given negative genetic correlations between fitness BOAG, P. T. 1981. Morphological Variation in the Darwin's components (the antagonistic pleiotropy Finches (Geospizinae) of Daphne Major Island, Galapagos. hypothesis). Much support has been generated for Ph.D. Dissertation, McGill University. this hypothesis (e.g., Luckinbill et al., 1984; Rose, BOACI, P. T. 1983. The heritability of external morphology in 1982, 1984; Rose and Charlesworth, 1981; Service Darwin's finches on Isla Daphne Major, Galapagos. 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Heritability of four morphological charac- Heritabilityof ecologicallyimportant traitsin the greattit. ters in honeybees (Apis mellifica me1lfica L.). Genetica Ardea, 68, 193—203. Polonica,13, 41-49. Appendix Listing of species, traits represented, and sources. B =behaviour,L =lifehistory, M =morphology,and P= physiology. Species B L M P Reference Invertebrates Aedes aegypti * Duhrkopfand Young, 1979 (mosquito) Apis mellifera Hillesheim, 1984 (honeybee)) Apis mell(fica * * Collingsci a!, 1984 (honeybee) Milneand Friars, 1984 Oldroyd and Moran, 1983 * Rinder cia!., 1983 * * Sollerand Bar-Cohen, 1967 * Zawilski, 1972 Ariantaarbustorum * Cook, 1965 (snail) Asellus aqua ticus Pashkova, 1978 (isopod) Cepaeanemoralis Clarke etal., 1978 (pulmonate) Crassostreavirginica Newkirk cia!., 1977 (oyster) C.gigas * * Lannan,1972 (Pacific oyster) Culex quinquefasciatus Ferrari eta!., 1982 (mosquito) Deloyala gut tata Rausher, 1983 (tortoise beetle) Dysdercus bimaculatus * * Derr,1980 (cottonstainer bug) Ephestriakuehniella Imura, 1980 (Mediterariean flour moth) Eurytemoraaffinis * Bradley, 1982 (copepod) E. herdmani * * McLaren,1976 (copepod) Glossina morsitan.c Gooding and Holebone, 1976 (tsetsefly) Gryllus integer Cadc, 1981 (field cricket) Heliconius era to Pansera and Aroujo, 1983 (butterfly) Heliothis zea * Holtzcrcia!., 1976 (corn earworm) Homerus a,nericarius * * Fairfullci a!., 1981 (American lobster) Finley and Haley, 1983 Hyphantria cunea Morris and Fulton, 1970 (fall webworm) HERITABILITY OF FITNESS COMPONENTS 195 Appendix continued Species B L M P Reference Lygaeus kalmii * Caldwell and Hegmann, 1969 (milkweed bug) Macrobrachium rosenbergii * Malecha et a!., 1984 (freshwater prawn) Maniolajurtina * McWhirter, 1969 (meadow brown butterfly) Musca domestica * Bryant, 1977 (house fly) * Bryant and Turner, 1978 Oncopeltusfasciatus * Dingle etaL, 1977 (milkweed bug) * Dingle et a!., 1980 * Dingle et a!., 1982 * * Hegmann and Dingle, 1982 Osmia!ignaria * Tepedina eta!.,1984 (megachilid bee) Pariula suturalis * Murray and Clarke, 1967 (snail) P. taeniata * Murray and Clarke, 1967 (snail) Pseudoca!anus * * McLaren and Corkett, 1978 (copepod) Spirorbisborealis * Doyle, 1974 (polychaete worm) Tribo!ium casteneum * Dawson, 1975 (flour beetle) * Englert and Bell, 1970 * Gall, 1971 * Halliburton and GaIl, 1981 * Lin eta!.,1979 * Minivielle and Gall, 1980 * Orozco, 1976 * Soliman, 1982 Fish Ape!tesquadracus * Hagen and Blouw, 1983 (fourspine stickleback) Coregonuslavaretus * Kirpichnikov, 1981 (whitefish) Cyprinuscarpio * Brody eta!., 1981 (common carp) * HoIm and Naevdal, 1978 * * Kirpichnikov, 1981 Moav and Wohlfarth, 1976 * * Nagy et a!., 1980 * * Smisek, 1979 Gambusiaaffinis * Busack, 1983 (mosquitofish) * Busack and Gall, 1983 Gasterosteus aculeatus * Hagen, 1973 (threespine stickleback) Icta!uruspunctatus * Bondari, 1983 (channel catfish) * Dunham and Smitherman, 1983 * * El-Ibiary and Joyce, 1978 * * Reagan, 1980 * Reagan eta!.,1976 Lebastesreticu!atus * Ryman, 1972, 1973 (guppyfish) Macropodus opercu!ar * Kirpichnikov, 1981 Oncorhynchus kisutch * Saxton et aL, 1984 (coho salmon) 0. nerka * Mcintyre and Amend, 1978 (sockeye salmon) 0. tshawytscha * Cramer and McIntyre, 1975 (chinook salmon) Oryziaslatipes * Kirpichnikov, 1981 Poeci!iareticulata * Kirpichnikov, 1981 (guppy) 196 T. A. MOUSSEAU AND D. A. ROFF Appendixcontinued Species B L M P Reference Salmo gairdneri * Aulstadet a!., 1972 (rainbow trout) * Ayles,1975 * Chevassus,1976 * Galland Gross, 1978a * Galland Gross, 1978h GaIl, 1975 Gjerde, 1982 * Gjerdeand Gjedrem, 1984 * * Gunnesand Ojedrem, 1981 * Holmand Naevdal, 1978 * Kaniscia!.,1976 * Kincaidei a!., 1977 * * Kinghorn,l983a, 1983b * Klupp,1979 * Linderei a!., 1983 * McKayet a!., 1984 * Mollerand Naevdal, 1973 * Mollerci a!., 1976 * Orska,1963, Refstie, 1980 Salmo salar * Gjedremand Aulstad, 1974 (Atlantic salmon) * * * Gjerdeand Gjedrem, 1984 * Gjerde,1981 * Gunnesand Gjedrem, 1978 * Holmand Naevdal, 1978 * Kaniset a!., 1976 * Kinghorn,1983a, l983b * Lindroth,1972 * Naevdalet a!., 1975 * * Naevdalci a!., 1976 * Refstieand Stein, 1978 * Refstieet a!., 1977 * * Riddeleta!., 1981 * * Ryman1972 Salmu irutia * Blancci a!., 1979 (brown trout) * Kanisci a!., 1976 * * Kirpichnikov,1981 Tilapia mossambila Kirpichnikov,1981 (tilapia) T. ni!ohica * Taveand Smitherman, 1980 (tilapia) Zoarce.c viviparus Kirpichnikov, 1981 (celpout) Amphibians and Reptiles Grapiemys ouachite,ns * Bullet a!., 1981 (Ouchita map turtle) Nairix sipedon * Beatson,1976 (water snake) P!eihodon cinereus * Highton,l96t) (red backed salamander) Rana pipiens Underhill, 1968 (leopard frog) R. iemporaria * Chernokozhcya,1983 (grass frog) Glushankova, 1982 Thamnophis elegans Arnold, 1981 (garter snake) * Ayresand Arnold, 1983 Rrds Anser caeru!escens * Findleyand Cooke, 1982, 1983 (lesser snow goose) HERITABILITY OF FITNESS COMPONENTS 197 Appendix continued Species B L M P Reference Branta canadensis * Lessels,1982 (Canada goose) Ficedula albicolis * Boag,1983 (collared flycatcher) F. hypoleuca * Boag,1983 (pied flycatcher) Geospizaconirostris * Grant, 1981, 1983 (Darwin'sfinch) G. fortis * Boag,1983, Boag and Grant, 1978 (Darwin's finch) G. scandens * Boag,1981, 1983 (Darwin's finch) Melospiza me!od,a * Smithand Dhondt, 1980 (song sparrow) * Smithand Zach, 1979 Parus caerulus * Dhondt,1982 (blue tit) P. major * Boag,1983, Dhondt, 1982 (great tit) * Garnett,1981 * Greenwoodet a!., 1979 * * Jones,1973 * Ojanenet a!., 1979 * Perrinsand Jones, 1974 * * * VanNoordwijk et a!., 1980, 1981a, 1981b Puffinus puffinus * Brook,1977 (manx shearwater) Sturnus vulgaris * Fluxand Flux, 1982 (starling) Mammals Macaca mulatta * Cheverud,1981 (rhesus macaque) M. radiata * Brookeret a!., 1981 (bonnett monkey) Mus musculus * Ebertand Hyde, 1976 (house mouse) * * Hydeand Sawyer, 1980 * Smith,1978 * VanOortmerssen and Bakker, 1981 Peromyscusgambeli * Sumner,1918 (deer mouse) P. rubidus * Sumner,1918 (deer mouse) P. sonoriens * Sumner,1918 (deer mouse) Raltus norvegicus * Hewittand Fulkner, 1983 (Norwegian rat)