We have investigated the reduction of fitness caused by the fixation of new deleterious mutations in small populations within the framework of Fisher's geometrical model of adaptation. In Fisher's model, a population evolves in an n-dimensional character space with an adaptive optimum at the origin. The model allows us to investigate compensatory mutations, which restore fitness losses incurred by other mutations, in a context-dependent manner. We have conducted a moment analysis of the model, supplemented by the numerical results of computer simulations. The mean reduction of fitness (i.e., expected load) scaled to one is approximately n/(n 2N_e), where N_e is the effective population size. The reciprocal relationship between the load and N_e implies that the fixation of deleterious mutations is unlikely to cause extinction when there is a broad scope for compensatory mutations, except in very small populations. Furthermore, the dependence of load on n implies that pleiotropy plays a large role in determining the extinction risk of small populations. Differences and similarities between our results and those of a previous study on the effects of N_e and n are explored. That the predictions of this model are qualitatively different from studies ignoring compensatory mutations implies that we must be cautious in predicting the evolutionary fate of small populations and that additional data on the nature of mutations is of critical importance.

Corresponding Editor: H. A. Orr

Independent contrasts are widely used to incorporate phylogenetic information into studies of continuous traits, particularly analyses of evolutionary trait correlations, but the effects of taxon sampling on these analyses have received little attention. In this paper, simulations were used to investigate the effects of taxon sampling patterns and alternative branch length assignments on the statistical performance of correlation coefficients and sign tests; “full-tree” analyses based on contrasts at all nodes and “paired-comparisons” based only on contrasts of terminal taxon pairs were also compared. The simulations showed that random samples, with respect to the traits under consideration, provide statistically robust estimates of trait correlations. However, exact significance tests are highly dependent on appropriate branch length information; equal branch lengths maintain lower Type I error than alternative topological approaches, and adjusted critical values of the independent contrast correlation coefficient are provided for use with equal branch lengths. Nonrandom samples, with respect to univariate or bivariate trait distributions, introduce discrepancies between interspecific and phylogenetically structured analyses and bias estimates of underlying evolutionary correlations. Examples of nonrandom sampling processes may include community assembly processes, convergent evolution under local adaptive pressures, selection of a nonrandom sample of species from a habitat or life-history group, or investigator bias. Correlation analyses based on species pairs comparisons, while ignoring deeper relationships, entail significant loss of statistical power and as a result provide a conservative test of trait associations. Paired comparisons in which species differ by a large amount in one trait, a method introduced in comparative plant ecology, have appropriate Type I error rates and high statistical power, but do not correctly estimate the magnitude of trait correlations. Sign tests, based on full-tree or paired-comparison approaches, are highly reliable across a wide range of sampling scenarios, in terms of Type I error rates, but have very low power. These results provide guidance for selecting species and applying comparative methods to optimize the performance of statistical tests of trait associations.

Corresponding Editor: J. Losos

We develop a simple framework for modeling speciation and diversification as a continuous process of accumulation of genetic (or morphological) differences accompanied by species and subpopulation extinction and/or range expansion. This framework can be used to approach a number of questions such as species-area distribution, species–range size distribution, the rate of ecological turnover, asymmetries of range division between sister species, waiting time until speciation and extinction, the relationship between the geographic range size and the probability of speciation, the relationships between subpopulation-level parameters and metapopulation-level parameters, and the effects of taxonomic level on these rates, distributions, and parameters. We illustrate some of these applications using numerical simulations. We develop approximations describing the dependence of the number of different taxonomic units, their average range size, and the rate of their turnover on the system size, the rate of fixation of genetic (or morphological) changes in local demes, and the rate of local extinction and colonization.

Corresponding Editor: H. A. Orr

Nonadditive genetic variation and genetic disequilibrium are two important factors that influence the evolutionary trajectory of natural populations. We assayed quantitative genetic variation in a temporary-pond-dwelling population of Daphnia pulex over a full season to examine the role of nonadditive genetic variation and genetic disequilibrium in determining the short-term evolutionary trajectory of a cyclic parthenogen. Quantitative traits were influenced by three factors: (1) clonal selection significantly changed the population mean phenotype during the course of the growing season; (2) sexual reproduction and recombination led to significant changes in life-history trait means and the levels of expressed genetic variation, implying the presence of substantial nonadditive genetic variation and genetic disequilibrium; and (3) Egg-bank effects were found to be an important component of the realized year-to-year change. Additionally, we examined the impact of genetic disequilibria induced by clonal selection on the genetic (co)variance structure with a common principal components model. Clonal selection caused significant changes in the (co)variance structure that were eliminated by a single bout of random mating, suggesting that a build-up of disequilibria was the primary source of changes in the (co)variance structure. The results of this study highlight the complexity of natural selection operating on populations that undergo alternating phases of sexual and asexual reproduction.

Corresponding Editor: D. Waller

In many gynodioecous species, females produce more viable seeds than hermaphrodites. Knowledge of the relative contribution of inbreeding depression in hermaphrodites and maternal sex effects to the female fertility advantage and the genetic basis of variation in female fertility advantage is central to our understanding of the evolution of gender specialization. In this study we examine the relative contribution of inbreeding and maternal sex to the female fertility advantage in gynodioecious Thymus vulgaris and quantify whether there is genetically based variation in female fertility advantage for plants from four populations. Following controlled self and outcross (sib, within-population, and between-population) pollination, females had a more than twofold fertility advantage (based on the number of germinating seeds per fruit), regardless of the population of origin and the type of pollination. Inbreeding depression on viable seed production by hermaphrodites occurred in two populations, where inbreeding had been previously detected. Biparental inbreeding depression on viable seed production occurred in three of four populations for females, but in only one population for hermaphrodites. Whereas the maternal sex effect may consistently enhance female fertility advantage, inbreeding effects may be limited to particular population contexts where inbreeding may occur. A significant family × maternal sex interaction effect on viable seed production was observed, illustrating that the extent of female fertility advantage varies significantly among families. This result is due to greater variation in hermaphrodite (relative to female) seed fertility between families. Despite this genetic variation in female fertility advantage and the highly female biased sex ratios in populations of T. vulgaris, gynodioecy is a stable polymorphism, suggesting that strong genetic and/or ecological constraints influence the stability of this polymorphism.

Corresponding Editor: K. Holsinger

Specialists studying the genus Viola have consistently allied the Hawaiian violets comprising section Nosphinium—most of which are subshrubs or treelets—with putatively primitive subshrubs in certain South American violet groups. Hawaiian violets also possess inflorescences, a floral disposition otherwise found only in other genera of the Violaceae, thus strengthening the hypothesis of a very ancient origin for the Hawaiian species. A survey of phylogenetic relationships among infrageneric groups of Viola worldwide using nuclear rDNA internal transcribed spacer (ITS) sequences revealed a dramatically different biogeographic origin for the Hawaiian violets: A monophyletic Hawaiian clade was placed in a close sister relationship with the amphi-Beringian tundra violet, V. langsdorffii s. l., in a highly derived position. This remarkable and unforeseen relationship received strong clade support values across analyses, and monophyly of the Hawaiian lineage was further indicated by a unique 26-base-pair deletion in section Nosphinium. The high polyploid base chromosome number (n ≃ 40) in the Hawaiian violets relates them to Alaskan and eastern Siberian populations in the polyploid V. langsdorffii complex. More than 50 species of the 260 allochthonous birds wintering in the Hawaiian Islands are found to breed in the Arctic, occupying habitats in which individual birds might have encountered ancestral V. langsdorffii populations and served as dispersers to the central Pacific region. Acquisition of derived morphological traits (e.g., arborescence and inflorescences), significance of a confirmed Arctic origin for a component of the Hawaiian flora, and the likelihood of other “cryptic” Arctic elements in the Hawaiian flora deserving independent molecular phylogenetic corroboration are discussed.

Corresponding Editor: J. Conner

Trade-offs between flower size and number seem likely to influence the evolution of floral display and are an important assumption of several theoretical models. We assessed floral trade-offs by imposing two generations of selection on flower size and number in a greenhouse population of bee-pollinated Eichhornia paniculata. We established a control line and two replicate selection lines of 100 plants each for large flowers (S ), small flowers (S−), and many flowers per inflorescence (N ). We compared realized heritabilities and genetic correlations with estimates based on restricted-maximum-likelihood (REML) analysis of pedigrees. Responses to selection confirmed REML heritability estimates (flower size, h² = 0.48; daily flower number, h² = 0.10; total flower number, h² = 0.23). Differences in nectar, pollen, and ovule production between S and S− lines supported an overall divergence in investment per flower. Both realized and REML estimates of the genetic correlation between daily and total flower number were r = 1.0. However, correlated responses to selection were inconsistent in their support of a trade-off. In both S− lines, correlated increases in flower number indicated a genetic correlation of r = −0.6 between flower size and number. In contrast, correlated responses in N and S lines were not significant, although flower size decreased in one N line. In addition, REML estimates of genetic correlations between flower size and number were positive, and did not differ from zero when variation in leaf area and age at first flowering were taken into account. These results likely reflect the combined effects of variation in genes controlling the resources available for flowering and genes with opposing effects on flower size and number. Our results suggest that the short-term evolution of floral display is not necessarily constrained by trade-offs between flower size and number, as is often assumed.

Corresponding Editor: J. Conner

Although rarely tested, it is often assumed that interspecific competition results in the divergence of traits related to resource use. Using a plant-pollinator system as a model, I tested the prediction the presence of a competitor for pollination influences the strength and/or direction of pollinator-mediated selection on floral traits. I measured phenotypic selection via female fitness on five floral traits of Ipomopsis aggregata in seven populations. Four contained only conspecifics (I only) and three also contained the competitor Castilleja linariaefolia (C I). Directional selection via fruits/plant and conspecific pollen deposited/flower on corolla length was positive and significantly stronger in C I populations. This difference in selection was apparently driven by interpopulation variation in the degree to which reproduction of I. aggregata was pollen limited. Consistent with expectations of interspecific competition, I. aggregata plants in C I populations received less conspecific pollen per flower and set fewer seeds per fruit and fruits per plant than those in I only populations. Ipomopsis aggregata's corollas were also significantly longer in C I populations, suggesting that there had been a response to a similar selective regime in past generations. Phenotypic correlations between corolla length and width, which determine the variation in I. aggregata's flower shape, were significantly weaker in C I populations. These data suggest that competition for pollination can influence the strength of selection on and patterns of correlations among floral traits of I. aggregata. If I. aggregata populations with and without competitors for pollination are linked by gene flow, then measuring selection in competitive and noncompetitive environments maybe necessary to accurately predict how floral traits will evolve.

Corresponding Editor: D. Waller

Although most models of mating system evolution assign a central role to the male transmission advantage of selfing genotypes, empirical data on the male fitness consequences of increased self-pollination are still uncommon. Here, I use measures of pollen import and export by focal plants in genotyped arrays to investigate the effects of floral morphology and pollination environment on self and outcross male function. Plants from an autogamous population of Arenaria uniflora (Caryophyllaceae) exhibit complete pollen discounting relative to closely related outcrossers, as do morphologically intermediate F₁ hybrids between the two populations. However, the low cumulative male fitness of hybrids probably results from reduced pollen number or competitive ability, rather than a nonlinear relationship with floral morphology. When surrounded by selfers, plants from the outcrosser population self-fertilize at nearly the same rate as selfers (>80%), but have much lower self male fitness due to reduced fruit set. Because outcross siring success is also extremely low (<8%) in this treatment, these mate-limited outcrossers are at male fitness disadvantage to both pseudocleistogamous selfers and nonlimited outcrossers. The relative male fitness of plants with different mating systems appears dependent on the ecological context, as well as on morphological trade-offs.

Corresponding Editor: K. Holsinger

Chloroplast DNA (cpDNA) haplotype variation is compared among alpine and prairie/montane species of Packera from a region in southwestern Alberta that straddles the boundary of Pleistocene glaciation. The phylogeny of the 15 haplotypes identified reveals the presence of two groups: one generally found in coastal and northern species and the other from species in drier habitats. The presence of both groups in all four species and most populations from southwestern Alberta is evidence of past hybridization involving species or lineages that may no longer be present in the region. With the exception of the alpine P. subnuda (Φ_ST = 1.0), interpopulational subdivision of haplotype variation is low (Φ_ST < 0.350), suggesting that interpopulational gene flow is high. However, based on haplotype distribution patterns, we propose that Pleistocene hybridization and incomplete lineage sorting have resulted in reduced subdivision of interpopulational variation so that gene flow may not be as high as indicated. Drift has been more important in the alpine species populations, especially P. subnuda.

Corresponding Editor: K. Holsinger

The magnitude of fitness effects at genetic loci causing inbreeding depression at various life stages has been an important question in plant evolution. We used genetic mapping in a selfed family of loblolly pine (Pinus taeda L.) to gain insights on inbreeding depression for early growth and viability. Two quantitative trait loci (QTLs) were identified that explain much of the phenotypic variation in height growth through age 3 and may account for more than 13% inbreeding depression in this family. One of these QTLs maps to the location of cad-n1, a lignin biosynthesis mutation. Both QTLs show evidence of overdominance, although evidence for true versus pseudo-overdominance is inconclusive. Evidence of directional dominance for height growth was noted throughout the genome, suggesting that additional loci may contribute to inbreeding depression. A chlorophyll-deficiency mutation, spf, did not appear to be associated with growth effects, but had significant effects on survival through age 3. Previously identified embryonic viability loci had little or no overall effect on germination, survival, or growth. Our results challenge, at least in part, the prevailing hypothesis that inbreeding depression for growth is due to alleles of small effect. However, our data support predictions that loci affecting inbreeding depression are largely stage specific.

Corresponding Editor: O. Savolainen

Marine organisms exhibit great variation in reproductive modes, larval types, and other life-history traits that may have major evolutionary consequences. We measured local and regional patterns of genetic variation in corals along Australia's Great Barrier Reef to determine the relative contributions of sexual and asexual reproduction to recruitment and to infer levels of gene flow both locally (among adjacent sites, < 5 km apart) and regionally (among reefs separated by 500–1200 km). We selected five common brooding species (Acropora cuneata, A. palifera, Pocillopora damicornis, Seriatopora hystrix, and Stylophora pistillata) and four broadcast spawners (Acropora hyacinthus, A. cytherea, A. millepora, and A. valida), which encompassed a wide range of larval types and potential dispersal capabilities. We found substantial genotypic diversity at local scales in six of the nine species (four brooders, two spawners). For these six, each local population displayed approximately the levels of multilocus genotypic diversity (G_o) expected for outcrossed sexual reproduction (mean values of G_o:G_e ranged from 0.85 to 1.02), although consistent single-locus heterozygous deficits indicate that inbreeding occurs at the scale of whole reefs. The remaining three species, the brooder S. hystrix and the spawners A. valida and A. millepora displayed significantly less multilocus genotypic diversity (G_o) than was expected for outcrossed sexual reproduction (G_e) within each of several sites. Acropora valida and A. millepora showed evidence of extensive localized asexual replication: (1) a small number of multilocus (clonal) genotypes were numerically dominant within some sites (G_o:G_e values were as low as 0.17 and 0.20): (2) single-locus genotype frequencies were characterized by both excesses and deficits of heterozygotes (cf. Hardy-Weinberg expectations), and (3) significant linkage disequilibria occurred. For the brooding S. hystrix G_o:G_e values were also low within each of four sites (x̄ = 0.48). However, this result most likely reflects the highly restricted dispersal of gametes or larvae, because levels of genetic variation among sites within reefs were extremely high (F_SR = 0.28).

For all species, we detected considerable genetic subdivision among sites within each reef (high F_SR-values), and we infer that larval dispersal is surprisingly limited (i.e., N_em among sites ranging from 0.6 to 3.3 migrants per generation), even in species that have relatively long planktonic durations. Nevertheless, our estimates of allelic variation among reefs (F_RT) also imply that for all four broadcast spawning species and three of the brooders, larval dispersal is sufficient to maintain moderate to high levels of gene flow along the entire Great Barrier Reef (i.e., N_em among reefs ranged from 5 to 31). In contrast, widespread populations of S. hystrix and S. pistilata (the two remaining brooders) are relatively weakly connected (N_em among reefs was 1.4 and 2.5, respectively). We conclude that most recruitment by corals is very local, particularly in brooders, but that enough propagules are widely dispersed to ensure that both broadcast spawning and brooding species form vast effectively panmictic populations on the Great Barrier Reef.

Corresponding Editor: R. Burton

Allozymic variation at 21–23 loci was studied in 28 populations of Talitrus saltator, 23 populations of Orchestia montagui, 13 populations of O. stephenseni, and five populations of Platorchestia platensis from the Mediterranean Basin. Different levels of gene flow (Nm_θ) were detected within each species at the scale of the whole Mediterranean: O. montagui and P. platensis had low population structure, with levels of Nm_θ ≥ 1, whereas the T. saltator and O. stephenseni populations have values of Nm_θ < 1. The relationship between Nm_θ and geographic distance was analyzed to test for the presence of an isolation by distance pattern in the spatial genetic variation within each species. A model of isolation by distance is useful to describe the pattern of genetic structuring of study species at the scale of the whole Mediterranean: geographic distance explained from 28% to 70% of the variation in gene flow. In the Aegean area all species showed an island model of genetic structuring regardless of the levels of gene flow.

Corresponding Editor: B. Bowen

Sexual polymorphisms are model systems for analyzing the evolution of reproductive strategies. However, their plasticity and other binary traits have rarely been studied, with respect to environmental variables. A possible reason is that, although threshold models offer an adequate quantitative genetics framework for binary traits in a single environment, analyzing their plasticity requires more refined empirical and theoretical approaches. The statistical framework proposed here, based on the environmental threshold model (ETM), should partially fill this gap. This methodology is applied to an empirical dataset on a plastic sexual polymorphism, aphally, in the snail Bulinus truncatus. Aphally is characterized by the co-occurrence of regular hermaphrodites (euphallics) together with hermaphrodites deprived of the male copulatory organ (aphallics). Reaction norms were determined for 40 inbred lines, distributed at three temperatures, in a first experiment. A second experiment allowed us to rule out maternal effects. We confirmed the existence of high broad-sense heritabilities as well as a positive effect of high temperatures on aphally. However a significant genotype-by-environment interaction was detected for the first time, suggesting that sexual plasticity itself can respond to selection. A nested series of four ETM-like models was developed for estimating genetical effects on both mean aphally rate and plasticity. These models were tested using a maximum-likelihood procedure and fitted to aphally data. Although no perfect fit of models to data was observed, the refined versions of ETM models conveniently reduce the analysis of complex reaction norms of binary traits into standard quantitative genetics parameters, such as genetic values and environmental variances.

Corresponding Editor: S. Edwards

Sympatric races of pea aphids on alfalfa and red clover are highly ecologically specialized and significantly reproductively isolated. Much of the restriction of gene flow between the specialized populations is due to habitat choice behavior of the winged colonizers (Via 1999). Here, we document additional pre- and postmating reproductive isolation through selection against migrants and hybrids in the parental environments. First, a group of randomly chosen genotypes from each race that were experimentally migrated between hosts had very low survival and reproduction on the alternate host relative to genotypes originating from that host (natives). Such selection against cross-host migrants forms a premating barrier to gene flow because it is likely to reduce migrant frequencies before the sexual forms are induced in the fall. Our reciprocal transplant experiment also shows that natural selection acts directly on individual migrants between the crops to favor host choice behavior: genotypes from each host suffered large losses of fitness when forced to migrate to the alternate host plant relative to the fitness they would have enjoyed had they been able to choose their native host. In a companion field study, sequential sampling throughout the summer in newly colonized fields of both alfalfa and clover revealed a decrease in the frequency of host-specific marker alleles characteristic of the alternate crop. These field data further support the hypothesis that selection disfavors migrants that cross between crops. Second, when two sets of F₁ hybrids between the races were reciprocally tested on alfalfa and clover, both sets had significantly lower average fitness than the specialized parent in each of the two environments. This demographic selection against hybrids in the parental environments is a source of postmating reproductive isolation between the specialized races. Finally, significant genetic variation in fitness traits was seen among F₁ hybrid genotypes from both crosses between alfalfa and clover specialists. Although this variation suggests that a generalized pea aphid could evolve, such generalists are not seen in field collections of these populations.

Corresponding Editor: C. Boggs

The insect ovary is a modular structure, the functional unit of which is the ovariole. Ovariole number is positively correlated with potential reproductive output. Among drosophilids (Insecta: Diptera), ovariole number shows both phenotypic plasticity and substantial interspecific and interpopulational variation. Here we examine the mechanistic connection between phenotypic plasticity and genetically fixed variation in ovariole number within the melanogaster species group. When a laboratory population of Drosophila melanogaster was reared under reduced food conditions, differences in ovariole number were entirely due to alterations in cell differentiation during the wandering stage at the very end of larval development. Cell growth and cell death were not affected. When these same flies were reared under a variety of temperatures, ovariole number differences arose during the latter half of the third (final) larval instar. Cell differentiation was affected, although cell number was not, and ovariole number differences were established before metamorphosis. In contrast, genetically fixed, interspecific and interpopulational variability in ovariole number was caused by alterations in the dynamics of cell differentiation and by cell number differences. Furthermore, the stages affected were different in different species and populations in the melanogaster species group, ranging from the first (D. sechellia) through the middle of the third (D. simulans and D. mauritiana) larval stage. Therefore, the mechanistic bases for plasticity-based variability are largely distinct from the mechanistic bases for interspecific and interpopulational variability. Our results suggest that phenotypic plasticity indicates evolutionary flexibility in underlying ontogenetic processes.

Corresponding Editor: W. T. Starmer

Two factors that can affect genetic load, synergistic epistasis and sexual selection, were investigated in Drosophila melanogaster. A set of five chromosomal regions containing visible recessive mutations were put together in all combinations to create a full set of 32 homozygous lines fixed for different numbers of known mutations. Two measures of fitness were made for each line: productivity (a combined measure of fecundity and egg-to-adult survivorship) and competitive male mating success. Productivity, but not male mating success, showed a pattern of strong average synergistic epistasis, such that the log fitness declined nonlinearly with increasing numbers of mutations. Synergistic epistasis is known to reduce the mutation load. Both fitness components show some positive and some negative interactions between specific sets of mutations. Furthermore, alleles with deleterious effects on productivity tend to also diminish male mating success. Given that male mating success can affect relative fitness without changing the mean productivity of a population, these additional effects would lead to lower frequencies and lower fixation rates of deleterious alleles without higher costs to the mean fitness of the population.

Corresponding Editor: A. Caballero

The aim of this study is to examine the expression of cytoplasmic incompatibility and investigate the distribution and population frequencies of Wolbachia pipientis strains in Drosophila simulans. Nucleotide sequence data from 16S rDNA and a Wolbachia surface protein coding sequence and cytoplasmic incompatibility assays identify four distinct Wolbachia strains: wHa, wRi, wMa, and wAu. The levels of cytoplasmic incompatibility between six lines carrying these strains of bacteria and three control lines without bacteria are characterized. Flies infected with wHa and wRi are bidirectionally incompatible, and males that carry either strain can only successfully produce normal numbers of offspring with females carrying the same bacterial strain. Males infected with wAu do not express incompatibility. Males infected with the wMa strain express intermediate incompatibility when mated to females with no bacteria and no incompatibility with females with any other Wolbachia strain. We conduct polymerase chain reaction/restriction fragment length polymorphism assays to distinguish the strain of Wolbachia and the mitochondrial haplotype to survey populations for each type and associations between them. Drosophila simulans is known to have three major mitochondrial haplotypes (siI, siII, and siIII) and two subtypes (siIIA and siIIB). All infected lines of the siI haplotype carry wHa, wNo, or both; wMa and wNo are closely related and it is not clear whether they are distinct strains or variants of the same strain. Infected lines with the siIIA haplotype harbor wRi and the siIIB haplotype carries wAu. The wMa infection is found in siIII haplotype lines. The phenotypic expression of cytoplasmic incompatibility and its relation to between-population differences in frequencies of Wolbachia infection are discussed.

Corresponding Editor: H. A. Orr

Seasonal variation (spring, early summer, late summer, and autumn) of gametic disequilibrium between gene arrangements (O_ST and O_{3 4}) of the O chromosome and Lap, Pept-1, and Acph allozyme loci, located inside these inversions, has been recorded in a natural population of Drosophila subobscura during seven years over a 15-year period. The length of the study allowed us to investigate the temporal variation of the allozyme-inversion associations by statistical methods of time series analysis. Cyclic seasonal changes of allozyme-inversion associations for both Lap and Pept-1 are detected in the natural population. In both cases, the patterns of seasonal change are due to the seasonal change of frequency of Lap and Pept-1 allozymes occurring exclusively within the O_ST gene arrangement. In contrast, the allozyme frequencies at these loci within the O_{3 4} gene arrangement are stable along seasons. The patterns of temporal variation of allozyme-inversion associations for Lap and Pept-1 in the natural population are contrasted with those previously published that correspond to gene arrangements of the O chromosome and nucleotide polymorphism at the rp49 region located inside these inversions, suggesting that natural selection is operating on these allozyme-inversion associations.

Corresponding Editor: L. Nunney

Both female choice and male-male competition may take place during reproduction in many species. Female choice tends to be less obvious than male-male competition and consequently has received less attention from researchers. The opportunity for cryptic female choice arises after multiple insemination. Through postcopulatory processes, a female could alter the pattern of paternity among her offspring so that it does not directly reflect the different contributions of sperm made by her mates. To be able to determine if a female alters the relative sperm contributions of her mates, the behaviors and influences of the males must therefore be first taken into account. The interest of each male is to father all the offspring, and the interest of each female is to maximize paternal quality. Female yellow dung flies have complex internal reproductive tracts that may give them considerable control over the fertilization success of stored sperm from different males. In laboratory trials to date, the last male to mate has usually been most successful. In the present study, cryptic choice occurred in Scathophaga stercoraria and the pattern of choice was consistent with previously reported results. The fertilization success of a female's second mate (P2) was substantially larger if a female was kept at constant temperature and if the second male was genetically similar to her at the phosphoglucomutase (Pgm) locus. Females from the field normally have three spermathecae, but some have four. Lines were bred to have either three or four spermathecae. Flies from the different lines were crossed to generate females with similar genetic backgrounds that had either three or four spermathecae. P2 was significantly lower for high-quality females, that is, those that laid larger-than-average-clutches, with four spermathecae than for low-quality females with four spermathecae; female quality had no influence on P2 for females with three spermathecae. The results suggest that only large females may benefit from increased spermathecae number by being able to act against male interests. Females may only have three spermathecae, even though genetic variation for more is present, because selection for more spermathecae is weak.

Corresponding Editor: D. Wheeler

There are many proposed routes for the origin of premating reproductive isolation, but few systematic studies aimed at testing their relative importance. Accumulated information about the biogeographical history of the European meadow grasshopper, Chorthippus parallelus, has allowed us to make a planned series of comparisons among populations aimed at distinguishing the contributions of some of these hypotheses. We have compared the effects on assortative mating of long-term isolation in glacial refugia, founder events during postglacial colonization, and sympatry with a closely related species. A likelihood-based analysis allowed us to separate effects of variation in male and female mating propensity among populations from variation in mate choice leading to assortative mating. All three effects contributed significantly to the overall variation in mating pattern in a set of 21 pairwise comparisons among seven populations. Male cuticular composition, but not other candidate signals, was significantly associated with the level of assortative mating. Of the hypotheses for the origin of reproductive isolation, only the predictions of the founder hypothesis explained a significant amount of the variation in assortative mating. This does not rule out the possiblity that there may be some other explanation. Having established the pattern of divergence, it is possible to generate hypotheses that explain our results at least as well as the founder hypothesis. However, because many such post hoc hypotheses are possible, they cannot be tested with this dataset. On this basis, our results favor the hypothesis that some aspect of the colonization process tends to accelerate divergence in mating signals leading to premating reproductive isolation. This could be accomplished through any one of several mechanisms. Colonization involves many bottlenecks as new populations are established at the edge of the range by long-distance migrants. Genetic effects may be important, but these bottlenecks may also alter the conditions under which mates are found and chosen, as suggested by Kaneshiro. At the same time, the colonizing populations may encounter novel environmental challenges.

Corresponding Editor: J. Mallet

In island systems with diverging populations, the history of island formation and genealogical estimates of divergence dates can be mutually informative. In the “sky islands” of southeastern Arizona, climate-induced contraction of woodlands appears to have fragmented populations of woodland-dwelling species onto disjunct mountain ranges. Montane populations of the jumping spider, Habronattus pugillis, display striking amounts of phenotypic divergence among ranges. Paleoclimatic estimates date woodland fragmentation at approximately 10,000 years ago, suggesting that phenotypic divergence has been extraordinarily rapid in these spiders. This phylogeographic study of populations of H. pugillis attempts to clarify the species' history of isolation and divergence and to address the suitability of available paleoclimatic data for dating divergences among populations of the region's woodland-dwelling organisms. Mitochondrial sequence data of spiders from 13 mountain ranges was used to reconstruct genealogical relationships. Gene trees show that small mountain ranges tend to have populations whose sequences form monophyletic groups, whereas larger ranges do not. Paraphyly among genes from larger ranges could result from either recent migration or incomplete lineage sorting. I use phylogenetic and geographic information to test these alternatives, and conclude that incomplete lineage sorting best explains the observed paraphyly. Gene trees are concordant with some of the predictions of vegetation history generated by examination of topography. Dates estimated for divergence of populations vary from 30,000 years to more than 2 million years ago, suggesting multiple vicariance events that are older than would be inferred from paleoclimatic studies. These findings illustrate that use of any single paleontological dataset to calibrate molecular clocks can potentially greatly underestimate actual divergence times.

Corresponding Editor: S. Edwards

Under the indicator models of mate choice, female preferences evolve to exploit the condition-dependence or “indicator value” of male traits, which in turn may cause these traits to evolve to elaborate extremes. If the indicator value of a male trait changes, the payoff function of the female preference for that trait should change and the preference should evolve to a new optimum. I tested this prediction in the guppy, Poecilia reticulata, a species in which the indicator value of a sexually selected male trait, carotenoid coloration, varies geographically. Carotenoid coloration is thought to be an indicator of foraging ability and health because animals must obtain carotenoid pigments from their diet. The primary dietary source of carotenoids for guppies is unicellular algae, the abundance of which varies among natural streams because of variation in forest canopy cover. Carotenoid availability limits male coloration to a greater extent in streams with greater forest canopy cover. Thus, the indicator value of male coloration covaries positively with canopy cover. To test the indicator model prediction, I measured genetic divergence in the strength of female preferences for carotenoid coloration between high- and low-carotenoid availability streams in each of three river drainages. Second-generation laboratory-born females were given a choice between full-sib males raised on three different dietary levels of carotenoids. For all six populations, male attractiveness (as determined from the responses of females to male courtship displays) increased with dietary carotenoid levels. However, the strength of female preferences differed between populations in the predicted direction in only one of three river drainages. These results fail to support a crucial prediction of the indicator model. More studies taking an interpopulation approach to studying mate preference evolution are needed before the explanatory value of the indicator models can be rigorously assessed.

Corresponding Editor: B. Sullivan

Within the past two million years, more than 450 species of haplochromine cichlids have diverged from a single common ancestor in Lake Malawi. Several factors have been implicated in the diversification of this monophyletic clade, including changes in lake level and low levels of gene flow across limited geographic scales. The objectives of this study were to determine the effect of recent lake-level fluctuations on patterns of allelic diversity in the genus Metriaclima, to describe the patterns of population structure within this genus, and to identify barriers to migration. This was accomplished through an analysis of allele frequencies at four microsatellite loci. Twelve populations spanning four species within Metriaclima were surveyed. The effect of lake-level fluctuations can be seen in the reduced genetic diversity of the most recently colonized sites; however, genetic diversity is not depressed at the species level. Low levels of population structure exist among populations, yet some gene flow persists across long stretches of inhospitable habitat. No general barrier to migration was identified. The results of this study are interpreted with respect to several speciation models. Divergence via population bottlenecks is unlikely due to the large allelic diversity observed within each species. Genetic drift and microallopatric divergence are also rejected because some gene flow does occur between adjacent populations. However, the reduced levels of gene flow between populations does suggest that minor changes in the selective environment could cause the divergence of populations.

Corresponding Editor: S. Edwards

Biologists have long known that closely related species are often phenotypically different where they occur together, but are indistinguishable where they occur alone. The causes of such character displacement are controversial, however. We used polyphenic spadefoot toad tadpoles (Spea bombifrons and S. multiplicata) to test the hypothesis that character displacement evolves to minimize competition for food. We also sought to evaluate the role of phenotypic plasticity in the mediation of competitive interactions between these species. Depending on their diet, individuals of both species develop into either a small-headed omnivore morph, which feeds mostly on detritus, or a large-headed carnivore morph, which specializes on shrimp. Laboratory experiments and surveys of natural ponds revealed that the two species were more dissimilar in their tendency to produce carnivores when they occurred together than when they occurred alone. This divergence in carnivore production was expressed as both character displacement (where S. multiplicata's propensity to produce carnivores was lower in sympatry than in allopatry) and as phenotypic plasticity (where S. multiplicata facultatively enhanced carnivore production in S. bombifrons, and S. bombifrons facultatively suppressed carnivore production in S. multiplicata). In separate experiments, we established that S. bombifrons (the species for which carnivore production was enhanced) was the superior competitor for shrimp. Conversely, S. multiplicata (the species for which carnivore production was suppressed and omnivore production enhanced) was the superior competitor for detritus. These results therefore demonstrate that selection to minimize competition for food can cause character displacement. They also suggest that both character displacement and phenotypic plasticity may mediate competitive interactions between species.

Corresponding Editor: T. Mousseau

The European pool frog, Rana lessonae, is widely polymorphic for two common alleles (b, e) at the lactate dehydrogenase-B (LDH-B) locus. We compared fitness-related larval life-history traits among LDH-B genotypes, which originated from segregation in heterozygous parents, in an artificial pond experiment where tadpoles of R. lessonae from a Swiss population were raised together with tadpoles of the hemiclonal hybrid R. esculenta at two densities. In R. lessonae, LDH-B e/e homozygotes at each density had a higher proportion of metamorphs among survivors, reached metamorphosis earlier, and were heavier at metamorphosis than b/b homozygotes; b/e heterozygotes had intermediate values. That e/e individuals were superior to b/b in both time to and mass at metamorphosis is surprising because these two life-history traits are thought to reflect a performance trade-off; e/e genotypes apparently compensated for shorter time to metamorphosis by a higher growth rate. The two alleles showed the same performance ranking when combined in hybrids with a R. ridibunda allele: When R. esculenta from Swiss populations reared in the same ponds had received the e allele rather than the b allele from their R. lessonae parent, they reached metamorphosis earlier, but did not differ in mass at metamorphosis. The degree of linkage disequilibrium in the source population of the eight R. lessonae used as parents of the R. lessonae tadpoles is unknown, so we cannot exclude the possibility that the performance differences are caused by some anonymous tightly linked gene, rather than the LDH-B locus, that constitutes the genomically localized target of natural selection. A causal involvement of LDH-B is plausible, nevertheless, because this enzyme takes part in the central energy-metabolizing processes and has been reported to underlie fitness differences in other animals; also, differential performance of LDH-B genotypes has been observed in R. lessonae larvae from another population. The present results suggest strong directional selection for allele e; the sum of available data, including an independent laboratory experiment, suggests that partial environment-dependent overdominance combined with balancing selection favoring e/e homozygotes under some and b/b homozygotes under other conditions may be partially responsible for the broad maintenance of the LDH-B polymorphism in R. lessonae.

Corresponding Editor: E. Brodie III

The western terrestrial garter snake (Thamnophis elegans) varies significantly in individual growth rates and life-history traits (maturation, reproduction, and survival) among adjacent populations in nature. This study focuses on assessing the genetic and environmental components of the substantial among-population variation in growth rates. Litters of neonates from nine populations inhabiting either mountain meadow or lakeshore habitat were reared for one year in a common-garden experiment with two temperature treatments. Diet, frequency of feeding, light exposure, and daytime temperatures were identical for all individuals. The two different nighttime temperatures (20°C and 25°C) were chosen to mirror field differences in nighttime thermoregulatory constraints for mountain-meadow and lakeshore snakes, respectively. Temperature and source habitat interacted to affect first-year growth rate. Neonates from meadow dams grew fastest in the cooler treatment, whereas those from lakeshore dams grew fastest in the warmer treatment. The observation that naive neonates, which were gestated and raised under identical conditions, grew fastest in environments characteristic of their natal population is evidence both that there are genetic differences among populations for growth and that these differences reflect adaptation to local habitats at a very small geographic scale. In addition, significant directional selection for large birthweight was measured for neonates from all populations. These results are considered in the context of population colonization history, migration and selection, and competing models for growth rate variation.

Corresponding Editor: E. Brodie III

The thermoregulatory hypothesis proposes that endothermy in mammals and birds evolved as a thermoregulatory mechanism per se and that natural selection operated directly to increase body temperature and thermal stability through increments in resting metabolic rate. We experimentally tested this hypothesis by measuring the thermoregulatory consequences of increased metabolic rate in resting lizards (Varanus exanthematicus). A large metabolic increment was induced by feeding the animals and consequent changes in metabolic rate and body temperature were monitored. Although metabolic rate tripled at 32°C and quadrupled at 35°C, body temperature rose only about 0.5°C. The rate of decline of body temperature in a colder environment did not decrease as metabolic rate increased. Thus, increasing the visceral metabolic rate of this ectothermic lizard established neither consequential endothermy nor homeothermy. These results are inconsistent with a thermoregulatory explanation for the evolution of endothermy.

Corresponding Editor: M. Zelditch

There has been considerable debate in the study of hybrid zones as to whether hybrids may be superior to parental types within the area of contact (bounded hybrid superiority). In birds, naturally occurring hybridization is relatively common, and hybridization within this group always involves mate choice. If hybrids are superior, females choosing heterospecific mates should be expected to show higher fitness under the conditions prevalent in the hybrid zone. Hybrid superiority under these circumstances would reduce reinforcement and thereby help to maintain the hybrid zone. To examine this issue, we studied reproductive performances of hybrids and parental species of gulls (Larus occidentalis and Larus glaucescens) at two colonies within a linear hybrid zone along the west coast of the United States. This hybrid zone contains predominantly gulls of intermediate phenotype. Previous studies indicated that hybrids were superior to one or both parental types, but provided no data on possible mechanisms that underlie this hybrid superiority. Using a hybrid index designed specifically for these species, we identified to phenotype more than 300 individuals associated with nests, including both individual males and females within 73 pairs in the central portion of the hybrid zone and 74 pairs in the northern portion of the hybrid zone. There was little evidence of assortative mating, and what little there was resulted solely because of pairings within intergrades. In the central hybrid zone, females paired with hybrid males produced larger clutches and hatched and fledged more chicks compared with females paired to western gull males. This was a result of heavy predation on eggs in sand habitat, where male western gulls established territories. In contrast, many hybrid males established territories in vegetated cover that was less vulnerable to predation. In the northern part of the hybrid zone, clutch size did not differ among pair categories, however, there were differences in hatching and fledging success, with females paired to hybrid males showing better success compared to females paired to glaucous-winged gull males. Hybrids showed better hatching and fledging success in the north because hybrids are more like western gulls than glaucous-winged gulls in foraging behavior, taking a higher percentage of fish in their diet, which enhances chick growth and survival. This is believed to be the first documentation of bounded hybrid superiority that delineates the mechanisms that underlie hybrid superiority.

Corresponding Editor: S. Edwards

Patterns of genetic variation and covariation strongly affect the rate and direction of evolutionary change by limiting the amount and form of genetic variation available to natural selection. We studied evolution of morphological variance-covariance structure among seven populations of house finches (Carpodacus mexicanus) with a known phylogenetic history. We examined the relationship between within- and among-population covariance structure and, in particular, tested the concordance between hierarchical changes in morphological variance-covariance structure and phylogenetic history of this species. We found that among-population morphological divergence in either males or females did not follow the within-population covariance patterns. Hierarchical patterns of similarity in morphological covariance matrices were not congruent with a priori defined historical pattern of population divergence. Both of these results point to the lack of proportionality in morphological covariance structure of finch populations, suggesting that random drift alone is unlikely to account for observed divergence. Furthermore, drift alone cannot explain the sex differences in within- and among-population covariance patterns or sex-specific patterns of evolution of covariance structure. Our results suggest that extensive among-population variation in sexual dimorphism in morphological covariance structure was produced by population differences in local selection pressures acting on each sex.

Corresponding Editor: T. Mousseau

Segregation distorters are selfish genetic elements that bias Mendelian segregation in their favor. All well-known segregation distortion systems consist of one or more “distorter” loci that act upon a “responder” locus. At the t complex of the house mouse, segregation distortion is brought about by the harmful effect of t alleles at a number of distorter loci on the wild-type variant of the responder locus. The responder and distorter alleles are closely linked by a number of inversions, thus forming a coherent t haplotype. It has been conjectured that the close integration of the various components into a “complete” t haplotype has been crucial for the evolutionary success of these selfish genetic elements. By means of a population genetical metapopulation model, we show that this intuition may be unfounded. In fact, under most circumstances an “insensitive” t haplotype retaining only the responder did invade and reach a high frequency, despite the fact that this haplotype has a strong segregation disadvantage. For certain population structures, the complete t haplotype was even competitively excluded by partial t haplotypes with lower segregation ratios. Moreover, t haplotypes carrying one or more recessive lethals only prevailed over their nonlethal counterparts if the product of local population size and migration rate (Nm) was not much smaller or larger than one. These phenomena occurred for rather realistic fitness, segregation, and recombination values. It is therefore quite puzzling that partial t haplotypes are absent from natural house mousepopulations, and that t haplotypes carrying recessive lethals prevail over nonlethal t haplotypes.

Corresponding Editor: S. Edwards

The D′ coefficient is one of the most commonly used measures of the extent of gametic disequilibrium between multiallelic loci. It has been suggested that the range of the D′ measure of overall disequilibrium between pairs of multiallelic loci depends on allele frequencies, except under some very restricted conditions. Nevertheless, the problem of dependence of the range of D′ has not been characterized under a wide set of possible polymorphisms. Evaluation of the utility of D′ as a measure of the strength of overall disequilibrium between all possible pairs of alleles at two multiallelic loci requires better knowledge of its range than is currently available. In this work, the conditions of polymorphism under which the range of D′ is frequency independent are given. It is found that the range of D′ is more often independent of allelic frequencies than is commonly thought. Furthermore, the range of D′ undergoes only small fluctuations as a function of the polymorphisms at the loci. Numerical cases and microsatellite data from humans are used for illustration. These observations indicate that the D′ coefficient is a useful tool for the estimation and comparison of the extent of overall disequilibrium across pairs of multiallelic loci.

Corresponding Editor: D. Waller

We performed an allozyme survey of genetic differentiation in Pinus balfouriana, a subalpine conifer endemic to California that is comprised of two allopatric subspecies, one in the Klamath Mountains and the other in the southern Sierra Nevada. Although the two subspecies are morphologically distinct and gene flow between them is virtually nonexistent, we observed much higher levels of differentiation among populations within a subspecies than between the two subspecies. Differentiation is particularly strong in the Klamath populations (multilocus F_ST = 0.242), which are small, isolated, and ecologically marginal. We attribute this strong differentiation to the mountain island effect, in which populations restricted to high elevations become isolated from each other on different mountains separated by unsuitable intervening habitat, with consequent reduced gene flow allowing populations to evolve independently. Populations of P. balfouriana in the Klamath region only exist scattered on the few highest ridges and peaks that rise above 2000 m, which defines the lower limit of the species elevational distribution. This pattern of distribution has allowed genetic drift and allelic sorting through historical events to produce strong population-level differentiation, which was likely in place before the two subspecies were geographically separated. Because P. balfouriana occurs on both serpentine soils and nonserpentine soils in the Klamath Mountains, we tested for genetic differentiation between populations growing on serpentine versus nonserpentine soils and our results were equivocal. Our data, combined with several other studies of conifers, show that the mountain island effect can produce significant genetic differentiation in conifers whose life-history traits of widely dispersed pollen, long generation times, and high outcrossing rates would lead us to predict a more homogenous population genetic structure.

Corresponding Editor: M. Dudash

Latitudinal geographic variation in Drosophila melanogaster is pervasive. Parallel clines in traits such as body size, egg size, ovariole number, and development time have been found on several continents throughout the world. However, a cline in starvation resistance and fat content in D. melanogaster has so far been found only in India. Here we investigate starvation resistance and fat content in 10 populations from South America, in which clines in body size, egg size, and development time have previously been found. We find no evidence for a cline in starvation resistance or fat content in South America. We therefore suggest that the cline in starvation resistance in India may have evolved in response to specific climatic variation found only in India.

Corresponding Editor: W. T. Starmer

Estimates of the intensity of selection based on measurements of the living and the dead require knowledge of the fraction of the original population dying. We apply recently developed methods (Blanckenhorn et al. 1999) to estimate the intensity of selection in a population of cliff swallows. In this population the fraction of individuals dying was unknown, but certainly high. The inferred selection is very strong and impossible to achieve if the original population is assumed to have followed a normal distribution. We consider several alternative explanations for this result including measurement biases, undetected immigration, and sampling biases. Of these, sampling biases are perhaps the most likely. We conclude that the intensity of selection on the swallows was probably strong, but its absolute magnitude is unknown.

Corresponding Editor: D. Roff