House Finch Populations Research Articles

Stress responses and disease in three wintering house finch ( Carpodacus mexicanus) populations along a latitudinal gradient

In laboratory studies, stress hormones have been shown to impair immune functions, and increase susceptibility to diseases. However, the interactions between stress hormones and disease have rarely been studied in free-ranging populations. In this study, we measured concentrations of the avian stress hormone corticosterone across four winter months (December–March) over two years in three eastern North American house finch populations ( Carpodacus mexicanus) along a latitudinal gradient. Because Mycoplasma gallisepticum infections appear in these populations in late winter, we hypothesized that the timing of the disease outbreaks could be mediated by changes in corticosterone concentrations. We found a significant increase in baseline and stress-induced plasma corticosterone concentrations in house finches without Mycoplasma symptoms in late winter; when the prevalence of Mycoplasma infection peaks. We also found that house finches with Mycoplasma symptoms had elevated stress-induced corticosterone concentrations. High baseline concentrations were associated with a low body condition and a high fat load. We found that the relationship between corticosterone concentrations and the latitude of the study population changed between years. The first year, corticosterone concentrations were lowest in the southern latitude, but became higher in the second year when average winter temperatures were low. A causal understanding of the implications for this variation in corticosterone concentrations for Mycoplasma disease dynamics awaits further studies.

SUSCEPTIBILITY OF WILD SONGBIRDS TO THE HOUSE FINCH STRAIN OF MYCOPLASMA GALLISEPTICUM

Conjunctivitis in house finches (Carpodacus mexicanus), caused by Mycoplasma gallisepticum (MG), was first reported in 1994 and, since this time, has become endemic in house finch populations throughout eastern North America. Although the house finch is most commonly associated with MG-related conjunctivitis, MG has been reported from other wild bird species, and conjunctivitis (not confirmed as MG related) has been reported in over 30 species. To help define the host range of the house finch strain of MG and to better understand the effect of MG on other host species, we monitored a community of wild birds for exposure to MG and conducted experimental infections on nine avian species. For the field portion of our study, we conducted a 9-mo survey (August 2001 to April 2002) of wild avian species in a peri-urban environment on the campus of Auburn University. During this time 358 birds, representing 13 different families, were sampled. No clinical signs of mycoplasmosis were observed in any bird. Thirteen species from nine families had positive agglutination reactions for antibodies to MG, but all birds tested negative by polymerase chain reaction (PCR). Three mourning doves were PCR-positive for MG, but antibodies to MG were not detected. In the experimental infections, we exposed seven native avian species and two cage-bird species to MG (May 2000 to June 2002). After exposure, clinical disease was seen in all four species from the family Fringillidae and in eastern tufted titmice (Baeolophus bicolor). In addition, three other species were infected without clinical signs, suggesting that they may represent potential MG reservoirs.

Maternal Inheritance and Rapid Evolution of Sexual Size Dimorphism: Passive Effects or Active Strategies?

Adaptive evolution is often strongly influenced by maternal inheritance that transfers the parental strategies across generations. The consequences of maternal effects for the offspring generation depend on the between‐generation similarity in environments and on the evolved sensitivity of the offspring’s ontogeny to maternal effects. When these factors differ between sons and daughters, maternal effects can influence the evolution of sexual dimorphism. The establishment of house finch populations across western Montana during the last 30 years was accompanied by rapid evolutionary change in sexual size dimorphism. Here I show that traits that changed the most across generations were most influenced by maternal effects in males but not females. Maternal effects differentially affected sons' and daughters' survival; greater maternal effects were commonly associated with higher survival of sons, especially when maternal and offspring environments were similar. Stronger maternal effects extended preselection phenotypic variance in morphological traits of males, thereby producing some locally adaptive phenotypes and lessening juvenile mortality. Thus, the observed sex‐specific maternal effects and their contribution to the evolution of sexual size dimorphism are likely a passive consequence of the distinct sensitivity of sons and daughters to maternal adaptations to breeding in ecologically distinct parts of the house finch’s expanding range.

Evolution of sex-biased maternal effects in birds: I. Sex-specific resource allocation among simultaneously growing oocytes

Females in species that produce broods of multiple offspring need to partition resources among simultaneously growing ova, embryos or neonates. In birds, the duration of growth of a single egg exceeds the ovulation interval, and when maternal resources are limited, a temporal overlap among several developing follicles in the ovary might result in a trade-off of resources among them. We studied growth of oocytes in relation to their future ovulation order, sex, and overlap with other oocytes in a population of house finches (Carpodacus mexicanus) where strongly sex-biased maternal effects are favoured by natural selection. We found pronounced differences in growth patterns between oocytes that produced males and females. Male oocytes grew up to five times faster and reached their ovulation size earlier than female oocytes. Early onset and early termination of male oocytes' growth in relation to their ovulation resulted in their lesser temporal overlap with other growing ova compared with female oocytes. Consequently, ovulation mass of female but not male oocytes was strongly negatively affected by temporal overlap with other oocytes. In turn, mass of male oocytes was mostly affected by the order of ovulation and by maternal incubation strategy. These results provide a mechanism for sex-biased allocation of maternal resources during egg formation and provide insights into the timing of the sex-determining meiotic division in relation to ovulation in this species.

Mycoplasma gallisepticuminfection dynamics in a house finch population: seasonal variation in survival, encounter and transmission rate

SummaryWe considered the impact of an emerging pathogen (Mycoplasma gallisepticumEdward and Kanarek) on apparent survival, encounter and transition rates in a population of a novel host (the house finch,Carpodacus mexicanusMüller). We used a multistate analysis of mark–encounter data from individually marked birds. Individual birds were categorized to a particular disease ‘state’; transition rates among states, conditional on apparent survival, were analogous to rates of new infection and recovery from infection. We hypothesized thatM. gallisepticuminfection would reduce the apparent survival of infected individuals, and that the magnitude of this reduction would vary as a function of the physiological condition of the host (which was characterized in our analyses by including a demographic and an environmental surrogate as covariates).We found consistent support for the hypothesis thatM. gallisepticuminfection resulted in lower apparent survival among infected individuals, and that recovery rates (from infected to non‐infected) were greater than infection rates in this population. We also found strong evidence indicating that infected individuals were less likely to be encountered than were non‐infected individuals. Although we predicted that both sex and temperature (proxies for physiological condition) would explain a significant proportion of the variation in our data, only marginal influences of both factors on apparent survival, encounter and state transition rates were detected.Our analyses identified several factors that may be important to studies of disease in the wild. First, disease state assignment may be uncertain, which can complicate parameter estimation. Secondly, encounter rate for infected individuals in our study was low relative to that for non‐infected individuals, reflecting possible behavioural changes in infected individuals. Low encounter rates reduces precision of estimated parameters, especially for multistate models. Finally, our results (and mark–recapture models in general) assume independence among individual birds. However, we are aware that there is a social structuring in house finches (and in general for many bird species). Accounting for such non‐independence may be especially important for situations where the state transitions are directly related to the pattern of social contact.

Age, sex, and season affect the risk of mycoplasmal conjunctivitis in a southeastern house finch population

House finches (Carpodacus mexicanus (Muller, 1776)) in eastern North America have been affected by annual epidemics of an eye disease caused by the bacterium Mycoplasma gallisepticum since 1994. To identify factors associated with seasonal changes in prevalence and variation in host susceptibility, we monitored mycoplasmal conjunctivitis among wild house finches in a region of high prevalence in southeastern North America. We captured 888 birds between August 2001 and December 2003 and observed seasonal outbreaks characterized by rapid increases in prevalence from August to October each year. During periods of high prevalence, infection probability was significantly higher among juveniles than adults, and the severity of conjunctivitis among juvenile females was greater than for any other host category. We found no evidence linking moulting status to elevated infection risk among adult birds. Finally, house finches with conjunctivitis were in poorer condition than birds with no clinical signs of infection, particularly among those with severe infections. Results from this study are consistent with recent reports of seasonal and regional variation in mycoplasmal conjunctivitis and suggest that annual changes in host reproduction, behaviour, and age structure might be important determinants of the timing and magnitude of local epidemics.

Molecular variability of house finch Mycoplasma gallisepticum isolates as revealed by sequencing and restriction fragment length polymorphism analysis of the pvpA gene.

Mycoplasma gallisepticum, a major pathogen of chickens and turkeys, has caused significant declines in house finch (Carpodacus mexicanus) populations in the eastern United States since it was first observed in this species in 1994. There is evidence that M. gallisepticum infection is now endemic among eastern house finches, although disease prevalence has declined, suggesting an evolving host-parasite relationship. Studies based on randomly amplified polymorphic DNA (RAPD) have documented the presence of a single, unique RAPD profile in house finch M. gallisepticum isolates, suggesting a single point source of origin, which agrees with the known epidemiologic observations. In the present study, we evaluated the molecular variability of 55 house finch isolates as well as 11 chicken and turkey isolates including reference strains of M. gallisepticum. Molecular variability was evaluated by polymerase chain reaction (PCR)-restriction fragment length polymorphism (RFLP) analysis and nucleotide sequencing of the pvpA gene, which encodes for the putative cytadhesin protein PvpA. Three different RFLP groups and 16 genotypes were evident from the 55 house finch isolates evaluated. Sequence analysis of pvpA gene PCR products showed that although most house finch M. gallisepticum isolates clustered more closely to each other, others clustered more closely to either turkey or chicken field isolates. These findings suggest that house finch isolates are more polymorphic than previously recognized by RAPD studies. This feature may allow us to learn more about the molecular evolution and epidemiology of this emerging disease host-parasite relationship.

First Case of Mycoplasma Gallisepticum Infection in the Western Range of The House Finch (Carpodacus Mexicanus)

Abstract We report the first case of mycoplasmosis in the western range of the House Finch (Carpodacus mexicanus). This disease originated in the eastern United States and has been previously documented only in eastern introduced House Finch populations where it reached epizootic proportions causing extensive and widespread mortality. Documentation of this dis-ease in western Montana suggests that previously disjunct eastern and western populations of House Finches are now mixing in the northern part of their range. More importantly, as native House Finches are highly susceptible to this novel pathogen, western populations may now be at risk of high mortality, similar to that experienced by non-native eastern populations. Close monitoring of this disease in the western part of the House Finch range will provide important insight into the dynamics of the emerging disease and evolution of resistance to the pathogen.

THE EVOLUTION OF SEXUAL SIZE DIMORPHISM IN THE HOUSE FINCH. V. MATERNAL EFFECTS

The phenotype of a mother and the environment that she provides might differentially affect the phenotypes of her sons and daughters, leading to change in sexual size dimorphism. Whereas these maternal effects should evolve to accommodate the adaptations of both the maternal and offspring generations, the mechanisms by which this is accomplished are rarely known. In birds, females adjust the onset of incubation (coincident with the first egg or after all eggs are laid) in response to the environment during breeding, and thus, indirectly, determine the duration of offspring growth. In the two house finch (Carpodacus mexicanus) populations that breed at the extremes of the species' distribution (Montana and Alabama), females experience highly distinct climatic conditions during nesting. We show that in close association with these conditions, females adjusted jointly the onset of incubation and the sequence in which they produced male and female eggs and consequently modified the growth of sons and daughters. The onset of incubation in newly breeding females closely tracked ambient temperature in a pattern consistent with the maintenance of egg viability. Because of the very different climates in Montana and Alabama, females in these populations showed the opposite patterns of seasonal change in incubation onset and the opposite sex bias in egg-laying order. In females with breeding experience, incubation onset and sex bias in laying order were closely linked regardless of the climatic variation. In nests in which incubation began with the onset of egg laying, the first-laid eggs were mostly females in Montana, but mostly males in Alabama. Because in both populations, male, but not female, embryos grew faster when exposed to longer incubation, the sex-bias produced highly divergent sizes of male and female juveniles between the populations. Overall, the compensatory interaction between the onset of incubation and the sex-biased laying order achieved a compromise between maternal and offspring adaptations and contributed to rapid morphological divergence in sexual dimorphism between populations of the house finch breeding at the climatic extremes of the species range.

FIRST CASE OF MYCOPLASMA GALLISEPTICUM INFECTION IN THE WESTERN RANGE OF THE HOUSE FINCH (CARPODACUS MEXICANUS)

We report the first case of mycoplasmosis in the western range of the House Finch (Carpodacus mexicanus). This disease originated in the eastern United States and has been previously documented only in eastern introduced House Finch populations where it reached epizootic proportions causing extensive and widespread mortality. Documentation of this dis- ease in western Montana suggests that previously disjunct eastern and western populations of House Finches are now mixing in the northern part of their range. More importantly, as native House Finches are highly susceptible to this novel pathogen, western populations may now be at risk of high mortality, similar to that experienced by non-native eastern populations. Close monitoring of this disease in the western part of the House Finch range will provide important insight into the dynamics of the emerging disease and evolution of resistance to the pathogen. Received 10 May 2002, accepted 5 February 2003.

Paternal care as a conditional strategy: distinct reproductive tactics associated with elaboration of plumage ornamentation in the house finch

When individuals in a population differ in physiological condition and residual reproductive value, selection should favor phenotypic plasticity in reproductive investment such that individuals are able to adopt the reproductive tactic that results in the highest fitness under given conditions. Here we examined reproductive tactics in relation to the elaboration of conditiondependent sexual ornamentation (carotenoid breast coloration) in a Montana population of the house finches (Carpodacus mexicanus). Males used distinct reproductive tactics depending on elaboration of their sexual ornamentation. Males with red pigmentation (maximum ornament elaboration) paired with females that nested earlier, but these males did little provisioning of incubating females and nestlings. In contrast, males with yellow coloration paired with females that nested later, but these males fed female and nestlings more. Consequently, for red males offspring recruitment was primarily affected by earlier nest initiation, whereas in yellow males it was affected most by male provisioning. In males with intermediate plumage coloration, all measured components, nest initiation, provisioning of incubating female, and nestling feeding, strongly contributed to offspring recruitment. The fitness consequences of alternative reproductive tactics of males were influenced by breeding experience and fidelity of their mates. Among first-time breeders, red males achieved the highest fecundity because of the advantage gained through early nesting and pairing with more experienced females and because of compensation by their mates for low male provisioning of nestlings. Among experienced breeders, males with intermediate plumage coloration achieved the highest fecundity because of the combined benefits of relatively early pairing and high parental care. High variation in sexual ornamentation in a Montana population of house finches may favor distinct associations of sexual displays with a particular set of reproductive behaviors. Key words: Carpodacus mexicanus, conditional strategies, house finches, parental care, reproductive investment, secondary sexual traits. [Behav Ecol 13:591–597 (2002)]

Susceptibility of a naïve population of house finches to Mycoplasma gallisepticum.

Since 1994 an epidemic of mycoplasmal conjunctivitis has spread throughout the eastern house finch (Carpodacus mexicanus) population leading to a significant decline in this population. The infection has not yet been reported from house finch populations west of the Great Plains. We hypothesized that the western population, like the eastern population, is susceptible to infection, and we tested this hypothesis by experimentally infecting house finches from Missoula, Montana (USA) with the house finch strain of Mycoplasma gallisepticum (MG). We compared the response of finches from Montana infected with MG to that of finches from Auburn, Alabama (USA) (October 1999-February 2000). Fifteen house finches from Montana were shipped to Auburn and quarantined for 6 wk at the Auburn University aviary. All birds were negative for MG antibodies when tested by serum plate agglutination assay and MG could not be detected in any bird by polymerase chain reaction. We tested two methods of inoculation, ocular inoculation and contact exposure to an infected finch. Seven house finches from Montana and four house finches from Alabama were infected by bilateral ocular inoculation with 20 microliters of a culture containing 1 x 10(6) color changing units of the house finch strain of MG. The remaining eight house finches from Montana were co-housed with a house finch from Alabama exhibiting mycoplasmal conjunctivitis. After exposure to the pathogen, all house finches became infected, regardless of origin or method of exposure, and all developed conjunctivitis. All birds seroconverted, and evidence of infection could be detected in every bird at some point during the course of disease. Our results suggest that house finches from the western United States are highly susceptible to infection with the house finch strain of MG.

A condition dependent link between testosterone and disease resistance in the house finch.

Testosterone has recently been proposed as a link between male quality and health and the expression of sexual traits. We investigated the relationship between testosterone and measures of the individual condition and health of males in a natural population of house finches (Carpodacus mexicanus). We also conducted a captive experiment in order to test for the effects of testosterone on resistance to coccidia, which is a common parasite of house finches. Free-living males in better condition had higher testosterone levels and lower corticosterone levels than free-living males in poor condition. In our captive experiment, increased testosterone accelerated the rate of coccidial infection as compared with sham-implanted or gonadectomized males. Although the differences were not significant, free-living males infected with coccidia had lower levels of testosterone and higher levels of corticosterone than males that were not infected. Thus, experimentally elevating testosterone levels in captive males resulted in a higher percentage of infected males, while free-living males with coccidial infection had low testosterone levels. This apparent discrepancy between captive and free-living males in the association of testosterone and disease may be explained by the condition dependence of testosterone. These results suggest that the testosterone-dependent sexual traits reliably indicate male overall condition and health and, thus, females could benefit from assessing potential mates based on these traits.

Plumage Color as a Composite Trait: Developmental and Functional Integration of Sexual Ornamentation

Most studies of condition-dependent sexual ornaments have treated such ornaments as single traits. However, sexual ornaments are often composites of several components, each produced by partially independent developmental pathways. Depending on environmental and individual condition, components of these ornaments may reflect different behavioral or physiological properties of an individual. One of the best-known, condition-dependent ornaments is carotenoid-based plumage coloration, which has at least four distinct components: pigment elaboration, patch area, pigment symmetry, and patch area symmetry. Here we examined fitness consequences of variation in individual components of carotenoid ornamentation in male house finches (Carpodacus mexicanus). Over 5 yr and several selection episodes, we studied variation in the plumage components in a large sample (n = 498) of males from a Montana population. The ornament components were partially independent of each other and had distinct fitness consequences. Selection for higher fecundity favored an increase in redness of coloration and a decrease in pigment asymmetry and patch area asymmetry but did not act on patch area itself. In contrast, viability selection favored larger and more symmetrical ornamental patches but did not act on pigment elaboration. Developmental and functional interrelationships among individual components of ornamentation strongly differed between house finch populations. Distinct patterns of selection on individual components of condition-dependent ornaments, combined with partially independent development of components, should favor the evolution of composite sexual traits whose components reliably reflect condition across a wide array of environments.

Mycoplasma gallisepticum in House Finches (Carpodacus mexicanus) and Other Wild Birds Associated with Poultry Production Facilities

Since 1994, an epidemic of conjunctivitis caused by Mycoplasma gallisepticum (MG) has spread throughout the eastern population of house finches (Carpodacus mexicanus). The adaptation of MG to a free-flying avian species presents potential problems for the control of mycoplasmosis in commercial poultry. To evaluate risks associated with this emerging problem, a field survey was conducted to assess prevalence of MG infection in house finches and other passerine birds associated with poultry farms. Between November 1997 and March 1999, 1058 birds were captured by mist net or trap at 17 farms and at 10 feeder stations in northeast Georgia. Birds were bled and screened by serum plate agglutination (SPA) for antibodies to MG. Birds with negative or weak positive SPA results were released at capture sites, and those with strong positive SPA reactions were kept for further evaluation. Necropsies were performed on selected house finches and individuals of 11 other passerine species, and samples were collected for MG testing by culture, polymerase chain reaction (PCR), hemagglutination inhibition, and histopathology. Testing revealed 19.1% of 671 birds caught at farms and 11.6% of 387 birds caught at feeder sites were SPA positive for MG. Three house finches captured on farms were positive for MG by culture and PCR, whereas three from feeder sites were positive only by PCR. No MG isolates were made from tufted titmice (Baeolophus bicolor), but 40% were positive by PCR. Individuals from 10 additional species were SPA positive only. Results suggest that MG persists at low levels in house finches in northeast Georgia and that tufted titmice may be nonclinical carriers of MG or a related mycoplasma. Positive SPA reactions in other species may be caused by nonspecific reactions or contact exposure. Current biosecurity recommendations should be sufficient to minimize risks of transmission between wild and domestic birds.

Characterization of Mycoplasma gallisepticum Infection in Captive House Finches (Carpodacus mexicanus) in 1998

Since 1995, the epidemic of mycoplasmal conjunctivitis in eastern house finches has affected the Auburn, AL, house finch population. To better characterize the current status of this host-parasite interaction, we established a captive flock of 38 seronegative, healthy finches in fall 1998. After a minimum quarantine period of 4 wk, two Mycoplasma gallisepticum (MG)-infected house finches were introduced into this flock. Over a 12-wk period, the flock was captured every 2 wk and each bird was observed for conjunctivitis. Blood and choanal swabs were collected from each bird for serologic analysis and for the detection of MG by polymerase chain reaction. The infection spread rapidly through the flock just as it had in a similar study performed in 1996 at the height of the epidemic. Unlike the earlier study in which birds remained chronically infected, most of the birds in our study recovered rapidly, and only three of the birds died during the study. Two patterns of host response to infection with MG were observed. Twenty-seven birds (73%) experienced an acute conjunctivitis that resolved, and the birds appeared to clear the infection. Ten birds (27%) suffered prolonged clinical disease, and MG could be detected in these birds intermittently throughout the experiment. These results, in conjunction with our surveys of MG in the wild population, suggest an evolving host-parasite interaction.

THE EVOLUTION OF SEXUAL SIZE DIMORPHISM IN THE HOUSE FINCH. III. DEVELOPMENTAL BASIS

Sexual size dimorphism of adults proximately results from a combination of sexually dimorphic growth patterns and selection on growing individuals. Yet, most studies of the evolution of dimorphism have focused on correlates of only adult morphologies. Here we examined the ontogeny of sexual size dimorphism in an isolated population of the house finch (Carpodacus mexicanus). Sexes differed in growth rates and growth duration; in most traits, females grew faster than males, but males grew for a longer period. Sexual dimorphism in bill traits (bill length, width, depth) and in body traits (wing, tarsus, and tail length; mass) developed during different periods of ontogeny. Growth of bill traits was most different between sexes during the juvenile period (after leaving the nest), whereas growth of body traits was most sexually dimorphic during the first few days after hatching. Postgrowth selection on juveniles strongly influenced sexual dimorphism in all traits; in some traits, this selection canceled or reversed dimorphism patterns produced by growth differences between sexes. The net result was that adult sexual dimorphism, to a large degree, was an outcome of selection for survival during juvenile stages. We suggest that previously documented fast and extensive divergence of house finch populations in sexual size dimorphism may be partially produced by distinct environmental conditions during growth in these populations.Corresponding Editor: T. Mousseau

The evolution of sexual size dimorphism in the house finch. III. Developmental basis.

Sexual size dimorphism of adults proximately results from a combination of sexually dimorphic growth patterns and selection on growing individuals. Yet, most studies of the evolution of dimorphism have focused on correlates of only adult morphologies. Here we examined the ontogeny of sexual size dimorphism in an isolated population of the house finch (Carpodacus mexicanus). Sexes differed in growth rates and growth duration; in most traits, females grew faster than males, but males grew for a longer period. Sexual dimorphism in bill traits (bill length, width, depth) and in body traits (wing, tarsus, and tail length; mass) developed during different periods of ontogeny. Growth of bill traits was most different between sexes during the juvenile period (after leaving the nest), whereas growth of body traits was most sexually dimorphic during the first few days after hatching. Postgrowth selection on juveniles strongly influenced sexual dimorphism in all traits; in some traits, this selection canceled or reversed dimorphism patterns produced by growth differences between sexes. The net result was that adult sexual dimorphism, to a large degree, was an outcome of selection for survival during juvenile stages. We suggest that previously documented fast and extensive divergence of house finch populations in sexual size dimorphism may be partially produced by distinct environmental conditions during growth in these populations.

Nest mites (Pellonyssus reedi) and the reproductive biology of the house finch (Carpodacus mexicanus)

We investigated the effects of a hematophagous nestling mite (Pellonyssus reedi, Acari: Macronyssidae) on the reproductive biology of the house finch (Carpodacus mexicanus) in east-central Alabama, U.S.A. Mites were absent from nests for the first half of the breeding season, but after their initial appearance they increased in number and were present in almost all nests. High nest-mite levels were associated with decreased nestling mass and hematocrit, but not with decreased nestling tarsus length. Experimental elimination of mites from some nests confirmed that the effects observed were mite-induced, not seasonal. The plumage colour of breeding adult male house finches was not correlated with nest-mite levels, nor did it appear that redder males' offspring suffered less from the effects of mites. Adult house finches fed nestlings from highly parasitized nests less often than those from nests with few or no mites. It appears unlikely that mites are directly involved in the sexual selection of bright male plumage coloration in this population of house finches. However, it is known that early-nesting females preferentially pair with redder males, therefore the benefit of nesting early and avoiding mite infestations is greater for redder male house finches.

The evolution of sexual size dimorphism in the house finch. II. Population divergence in relation to local selection.

Recent colonization of ecologically distinct areas in North America by the house finch (Carpodacus mexicanus) was accompanied by strong population divergence in sexual size dimorphism. Here we examined whether this divergence was produced by population differences in local selection pressures acting on each sex. In a long-term study of recently established populations in Alabama, Michigan, and Montana, we examined three selection episodes for each sex: selection for pairing success, overwinter survival, and within-season fecundity. Populations varied in intensity of these selection episodes, the contribution of each episode to the net selection, and in the targets of selection. Direction and intensity of selection strongly differed between sexes, and different selection episodes often favored opposite changes in morphological traits. In each population, current net selection for sexual dimorphism was highly concordant with observed sexual dimorphism--in each population, selection for dimorphism was the strongest on the most dimorphic traits. Strong directional selection on sexually dimorphic traits, and similar intensities of selection in both sexes, suggest that in each of the recently established populations, both males and females are far from their local fitness optimum, and that sexual dimorphism has arisen from adaptive responses in both sexes. Population differences in patterns of selection on dimorphism, combined with both low levels of ontogenetic integration in heritable sexually dimorphic traits and sexual dimorphism in growth patterns, may account for the close correspondence between dimorphism in selection and observed dimorphism in morphology across house finch populations.