Change in frequency of a rare mutant allele: A general formula and applications to partial inbreeding models

 We deduce and prove a general formula to approximate the change in frequency of a mutant allele under weak selection, when this allele is introduced in small frequency into a population which was previously at a fixation state. We apply the formula to autosomal genes in partial selfing models and to autosomal as well as sex-linked genes in partial sib mating models. It is shown that the fate of a rare mutant allele depends not only on the selection parameters, the inbreeding coefficient and the reproductive values of the sexes in sex-differentiated populations, but also on coefficients of relatedness between mates. This is interpreted as a kin selection effect caused by inbreeding per se. Received: 3 December 2001 / Revised version: 10 April 2002 / Published online: 19 November 2002 Research supported in part by NSERC of Canada and FCAR of Québec. Mathematics Subject Classification (2000): Primary 60J80, Secondary 92D10, 92D25 Keywords or phrases: Adaptive topography – Partial selfing – Partial sib mating – Kin selection     

Distribution, abundance, and seasonal ecology of Listronotus maculicollis (Coleoptera: Curculionidae) on golf courses in Québec, Canada

The weevil Listronotus maculicollis Dietz (Coleoptera: Curculionidae) is a major pest of annual bluegrass, Poa annua L., on golf courses in northeastern North America. To determine the distribution, abundance, and seasonal ecology of L. maculicollis on golf courses in Québec, Canada, we sampled 19 golf courses (GC1-GC19) from different geographic and climatic conditions during 2001, 2002, and 2003. L. maculicollis was found on all golf courses except GC19, which was located in northeastern Québec. In most sites, L. maculicollis completed two generations per year. A linear regression model comparing L. maculicollis larval densities as a function of spatial coordinates revealed that L. maculicollis were more abundant in western Québec than in eastern Québec. Pearson correlations on the probability of both L. maculicollis larval and pupal presence with soil parameters indicated significant positive correlations for fine sand (larvae) and soil pH (pupae) and negative correlations for gravel.     

Canid hybridization: contemporary evolution in human‐modified landscapes

Contemporary evolution through human‐induced hybridization occurs throughout the taxonomic range. Formerly allopatric species appear especially susceptible to hybridization. Consequently, hybridization is expected to be more common in regions with recent sympatry owing to human activity than in areas of historical range overlap. Coyotes ( Canis latrans) and gray wolves ( C. lupus) are historically sympatric in western North America. Following European settlement gray wolf range contracted, whereas coyote range expanded to include eastern North America. Furthermore, wolves with New World (NW) mitochondrial DNA (mtDNA) haplotypes now extend from Manitoba to Québec in Canada and hybridize with gray wolves and coyotes. Using mtDNA and 12 microsatellite markers, we evaluated levels of wolf‐coyote hybridization in regions where coyotes were present (the Canadian Prairies, n = 109 samples) and absent historically (Québec, n = 154). Wolves with NW mtDNA extended from central Saskatchewan (51°N, 69°W) to northeastern Québec (54°N, 108°W). On the Prairies, 6.3% of coyotes and 9.2% of wolves had genetic profiles suggesting wolf‐coyote hybridization. In contrast, 12.6% of coyotes and 37.4% of wolves in Québec had profiles indicating hybrid origin. Wolves with NW and Old World ( C. lupus) mtDNA appear to form integrated populations in both regions. Our results suggest that hybridization is more frequent in historically allopatric populations. Range shifts, now expected across taxa following climate change and other human influence on the environment, might therefore promote contemporary evolution by hybridization.     

Tree-ring analysis of white cedar (Thuja occidentalis L.) archaeological and historical wood in Québec City (Québec,Canada)

In Québec City, a segment of an old wooden palisade built for protection was found buried in situ. The palisade was excavated by the Laval University archaeology field school in 2004–2005, and 29 posts were recovered and sampled. The palisade enclosed the Intendant's Palace compound, which was the residence of the governor of New France at the end of the 17th century. Tree-ring analysis was performed on wood excavated from the Intendant's Palace archaeological site (PDI) and additional wood from two historical military buildings, the Artillery Park (ART) (early 18th century) and the Québec Citadel (CIT) (17th–19th centuries). Wood identification revealed that white cedar (Thuja occidentalis L.) was used for wood construction at the three sites. Many trees used for the construction of the PDI palisade were felled after the 1690 growing season, likely in September/October 1690. Posts probably came from trees growing close to the site on the banks of the Saint-Charles River. A white cedar ring-width chronology from the Rimouski area, approximately 300 km northeast of Québec City, along the St. Lawrence River, was used for cross-dating. Archaeological and historical wood samples from two of the three sites (PDI and ART) were first assembled in a 235-year tree-ring chronology, called the Québec chronology, extending from 1489 to 1723. The two master chronologies (Québec and Rimouski) were merged into a single 513-year tree-ring chronology (1489–2001), called the Saint-Laurent chronology.     

Conservation genetics of the threatened horned grebe (<Emphasis Type="Italic">Podiceps auritus</Emphasis> L.) population of the Magdalen Islands,Québec

The horned grebe (Podiceps auritus) population of the Magdalen Islands in the St. Lawrence Gulf (Québec, Canada) has declined sharply over the last decades. It is the only breeding population of this species in eastern North America with nearest breeding populations being >2500 km apart in western North America and Europe, We used three types of genetic markers: mitochondrial (mt) DNA ND2 sequence, α-enolase intron sequence, and 25 amplified fragment length polymorphism loci (AFLPs) to quantify the genetic diversity within the Magdalen Island population and to assess its genetic distinctiveness relative to populations from western Canada (five sites) and Iceland (one site). The Magdalen Island population retained a comparable amount of genetic diversity to the average diversity observed across all populations in all three markers. Horned grebe mtDNA sequences formed a monophyletic group and nearly all haplotypes present in Québec were found elsewhere. In the ND2 fragment, populations partitioned into two groups corresponding to subspecies (Iceland versus North American sites) and more strongly in three groups according to geographic disjunctions (Iceland versus Québec versus western Canada). In contrast, there was no evidence of structure between sites in the α-enolase intron. In the AFLPs, Iceland showed the greatest level of differentiation, followed by the Québec and British Columbia populations. For conservation purposes, we suggest that the Magdalen Islands population should be recognized as a separate unit.     

Genetic structure of the white‐footed mouse in the context of the emergence of Lyme disease in southern Québec

The white‐footed mouse (Peromyscus leucopus) has expanded its northern limit into southern Québec over the last few decades. P. leucopus is a great disperser and colonizer and is of particular interest because it is considered a primary reservoir for the spirochete bacterium that causes Lyme disease. There is no current information on the gene flow between mouse populations on the mountains and forest fragments found scattered throughout the Montérégie region in southern Québec, and whether various landscape barriers have an effect on their dispersal. We conducted a population genetics analysis on eleven P. leucopus populations using eleven microsatellite markers and showed that isolation by distance was weak, yet barriers were effective. The agricultural matrix had the least effect on gene flow, whereas highways and main rivers were effective barriers. The abundance of ticks collected from mice varied within the study area. Both ticks and mice were screened for the presence of the spirochete bacterium Borrelia burgdorferi, and we predicted areas of greater risk for Lyme disease. Merging our results with ongoing Lyme disease surveillance programs will help determine the future threat of this disease in Québec, and will contribute toward disease prevention and management strategies throughout fragmented landscapes in southern Canada.     

Do genetic drift and accumulation of deleterious mutations preclude adaptation? Empirical investigation using RADseq in a northern lacustrine fish

Understanding genomic signatures of divergent selection underlying long‐term adaptation in populations located in heterogeneous environments is a key goal in evolutionary biology. In this study, we investigated neutral, adaptive and deleterious genetic variation using 7,192 SNPs in 31 Lake Trout (Salvelinus namaycush) populations (n = 673) from Québec, Canada. Average genetic diversity was low, weakly shared among lakes, and positively correlated with lake size, indicating a major role for genetic drift subsequent to lake isolation. Putatively deleterious mutations were on average at lower frequencies than the other SNPs, and their abundance relative to the entire polymorphism in each population was positively correlated with inbreeding, suggesting that the effectiveness of purifying selection was negatively correlated with inbreeding, as predicted from theory. Despite evidence for pronounced genetic drift and inbreeding, several outlier loci were associated with temperature and found in or close to genes with biologically relevant functions notably related to heat stress and immune responses. Outcomes of gene–temperature associations were influenced by the inclusion of the most inbred populations, in which allele frequencies deviated the most from model predictions. This result illustrates challenge in identifying gene–environment associations in cases of high genetic drift and restricted gene flow and suggests limited adaptation in populations experiencing higher inbreeding. We discuss the relevance of these findings for the conservation and management, notably regarding stocking and genetic rescue, of Lake Trout populations and other species inhabiting highly fragmented habitats.     

How did common ragweed (Ambrosia artemisiifolia L.) spread in Québec? A historical analysis using herbarium records

Aim To reconstruct the spread of common ragweed (Ambrosia artemisiifolia L.; Asteraceae) using herbarium specimens to document whether the habitat preferences of this plant have shifted through time. Location Southern Québec. Methods Herbarium specimens stored in the main herbaria of southern Québec were used to reconstruct the spread of common ragweed. All data (sampling location, year of sampling, habitat characteristics, etc.) were incorporated into a geographical information system. Maps indicating the spatial distribution of common ragweed were produced for four time periods. The cumulative number of locations was plotted against time to construct invasion curves. The sequence of habitats where herbarium specimens were collected was also reconstructed. Results A data base incorporating 707 common ragweed herbarium specimens was constructed for this study. The spread of common ragweed in most regions of southern Québec was initiated at the beginning of the 20th century. Herbarium specimens suggest that common ragweed first spread along river corridors. Specimens of common ragweed were not collected in agricultural fields before the mid‐1920s, nor along roads and railways before the mid‐1930s. The colonization of a large number of agricultural fields by common ragweed probably began with seed‐contaminated crops, but was certainly accelerated by the dispersal of seeds from populations growing along nearby roads. Main conclusions Herbarium specimens suggest that common ragweed has been present in southern Québec for at least 200 years, but the species was probably restricted to the Montréal area during the 19th century. It is likely that the development of the road network in Québec since the mid‐1930s significantly contributed to the spread of common ragweed. Controlling common ragweed solely in agricultural fields would not prevent the re‐infestation of crops, because roadsides would act as refuges for the weed.     

Organizational challenges to equity in the delivery of services within a new personalized risk-based approach to breast cancer screening

Emerging evidence opens new possibilities to improve current breast cancer mammography screening programs. One promising avenue is to tailor mammography screening according to individual risk. However, some factors could challenge the implementation of such approach, specifically its potential impact on the equitable delivery of services. This study aims to identify the barriers and facilitators to the equitable delivery of services within a future integration of a personalized approach in the Québec screening program. We then propose different means to address them. We conducted 16 semi-structured interviews with stakeholders with a role in the management, implementation or assessment of the Québec screening program. The barriers and facilitators identified by respondents were regrouped in two themes: 1) Reproduction of social inequities, and 2) Amplification of regional disparities in access to services. We consider that fostering inclusion through communication strategies and relying on electronic communication technologies could help in addressing these issues.     

Are Stepparents Always Evil? Parental Death,Remarriage, and Child Survival in Demographically Saturated Krummhörn (1720–1859) and Expanding Québec (1670–1750)

Parental death precipitates a cascade of events leading to more or less detrimental exposures, from the sudden and dramatic interruption of parental care to cohabitation with stepparents and siblings in a recomposed family. This article compares the effect of early parental loss on child survival in the past in the Krummhörn region of East Frisia (Germany) and among the French Canadian settlers of the Saint Lawrence Valley (Québec, Canada). The Krummhörn region was characterized by a saturated habitat, while the opportunities for establishing a new family were virtually unlimited for the French Canadian settlers. Early parental loss had quite different consequences in these dissimilar environments. Event history analyses with time-varying specification of family structure are used on a sample of 7,077 boys and 6,906 girls born between 1720 and 1859 in the Krummhörn region and 31,490 boys and 33,109 girls whose parents married between 1670 and 1750 in Québec. Results indicate that in both populations, parental loss is associated with increased infant and child mortality. Maternal loss has a universal and consistent effect for both sexes, while the impact of paternal loss is less easy to establish and interpret. On the other hand, the effect of the remarriage of the surviving spouse is population-specific: the mother's remarriage has no effect in Krummhörn, while it is beneficial in Québec. In contrast, the father's remarriage in Krummhörn dramatically reduces the survival chances of the children born from his former marriage, while such an effect is not seen for Québec. These population-specific effects appear to be driven by the availability of resources and call into question the universality of the “Cinderella” effect.     

Impact of urban fragmentation on the genetic structure of the eastern red-backed salamander

Urban development is a major cause of habitat loss and fragmentation. Few studies, however, have dealt with fragmentation in an urban landscape. In this paper, we examine the genetic structure of isolated populations of the eastern red-backed salamander (Plethodon cinereus) in a metropolitan area. We sampled four populations located on a mountain in the heart of Montréal (Québec, Canada), which presents a mosaic of forested patches isolated by roads, graveyards and buildings. We assessed the genetic structure of these populations using microsatellite loci and compared it to the genetic structure of four populations located in a continuous habitat in southern Québec. Our results indicate that allelic richness and heterozygosity are lower in the urban populations. Exact differentiation tests and pairwise F _ST also show that the populations found in the fragmented habitat are genetically differentiated, whereas populations located in the continuous habitat are genetically homogeneous. These results raise conservation concerns for these populations as well as for rare or threatened species inhabiting urban landscapes.     

Seroprevalence of antibodies to Toxoplasma gondii in lynx (Lynx canadensis) and bobcats (Lynx rufus) from Québec, Canada.

The seroprevalence of antibodies to Toxoplasma gondii was investigated in trapped lynx (Lynx canadensis) and bobcats (Lynx rufus) from Québec, Canada. Forty-seven of 106 (44%) lynx and 4 of 10 (40%) bobcats had positive titers for T. gondii (> or = 25) by means of the modified agglutination test incorporating mercaptoethanol and formalin-fixed tachyzoites. Seroprevalence was significantly higher (P < 0.0001) in adult lynx than in juvenile lynx. The presence of antibodies to T. gondii in lynx and bobcats suggests that this organism is widespread in the wild and that exposure to wild felids and game animals from Québec may represent a potential source of infection for humans.     

Population structure and impact of supportive breeding inferred from mitochondrial and microsatellite DNA analyses in land-locked Atlantic salmon Salmo salar L.

Four tributaries of Lake St-Jean (Québec, Canada) are used for spawning and juvenile habitat by land-locked Atlantic salmon. Spawning runs have drastically declined since the mid-1980s, and consequently, a supportive-breeding programme was undertaken in 1990. In this study, we analysed seven microsatellite loci and mtDNA, and empirically estimated effective population sizes to test the hypotheses that (i) fish spawning in different tributaries form genetically distinct populations and (ii) the supportive breeding programme causes genetic perturbations on wild populations. Allele frequency distribution, molecular variance and genetic distance estimates all supported the hypothesis of genetic differentiation among salmon from different tributaries. Gene flow among some populations was much more restricted than previously reported for anadromous populations despite the small geographical scale (40 km) involved. Both mtDNA and microsatellites revealed a more pronounced differentiation between populations from two tributaries of a single river compared with their differentiation with a population from a neighbouring river. The comparison of wild and F₁-hatchery fish (produced from breeders originating from the same river) indicated significant changes in allele frequencies and losses of low-frequency alleles but no reduction in heterozygosity. Estimates of variance and inbreeding population size indicated that susceptibility to genetic drift and inbreeding in one population increased by twofold after only one generation of supplementation.     

Effective size of populations under partial inbreeding and selection on a set of linked additive genes

The prediction theory of effective population size (Ne) is extended to cover selection on a set of linked additive genes and partial inbreeding (partial selfing or partial full-sib mating). Ne under selection is generally expressed as a function of the cumulative change in frequency of a neutral gene due to the random association between the neutral and selected genes generated by finite sampling. In this study, the association under partial selfing was classified into two types, the association between the neutral and selected genes on the same gamete, and the association between the neutral and selected genes each on the different gametes in the same parent. For partial full-sib mating, an additional association, i.e., the association between the neutral and selected genes each in the different parents in the same family, was included in the model. According to this classification of the association, the coefficient accounting for the cumulative change in frequency of the neutral gene was partitioned into two or three components. A method for computing the partitioned coefficients was obtained from the transition matrix approach, in which the joint effect of linkage, selection and partial inbreeding was taken into account. To assess the joint effects of linkage, selection and partial inbreeding on Ne, numerical computations with the obtained expressions were carried out. The effect of linkage on Ne was generally small, except for an extremely small genome size, while the partial inbreeding resulted in a drastic reduction in Ne. For a given genome size, Ne was essentially independent of the length and number of chromosomes. Some of these results were verified by stochastic simulations.     

An example of phenotypic adherence to the island rule? – Anticosti gray jays are heavier but not structurally larger than mainland conspecifics

The island rule refers to the tendency of small vertebrates to become larger when isolated on islands and the frequent dwarfing of large forms. It implies genetic control, and a necessary linkage, of size and body‐mass differences between insular and mainland populations. To examine the island rule, we compared body size and mass of gray jays (Perisoreus canadensis) on Anticosti Island, Québec, located in the Gulf of St. Lawrence, with three mainland populations (2 in Québec and 1 in Ontario). Although gray jays on Anticosti Island were ca 10% heavier, they were not structurally larger, than the three mainland populations. This suggests that Anticosti jays are not necessarily genetically distinct from mainland gray jays and that they may have achieved their greater body masses solely through packing more mass onto mainland‐sized body frames. As such, they may be the first‐known example of a proposed, purely phenotypic initial step in the adherence to the island rule by an insular population. Greater jay body mass is probably advantageous in Anticosti's high‐density, intensely competitive social environment that may have resulted from the island's lack of mammalian nest predators.     

Testing isonymy with paternal and maternal lineages in the early Québec population: the impact of polyphyletism and demographic differentials

Isonymy is an ingenious and useful approach to studying kinship in human populations. However, it relies on assumptions that are difficult to verify. In this study, we provided a way to assess, in the early Québec population, the impact of factors such as polyphyletism, unbalanced sex-ratio among founders, and age differentials between spouses. All data were taken from the Population Register of Early Québec, which contains births, marriages, and deaths (>712,000) recorded in parish registers from the beginning of colonization (in 1608) to 1800. More specifically, using the 70,869 marriages recorded during that period, we compared kinship estimates given by genealogies, surnames, and paternal and maternal lineages. We also calculated a fifth coefficient of kinship by combining paternal and maternal lineage, thus providing a new way to test the isonymy method. The results show a good agreement between genealogical and isonymous estimates. However, this good correspondence is due to counterbalancing biases. Some of the implications of our results are discussed in the context of colonial America.     

Change from pre‐settlement to present‐day forest composition reconstructed from early land survey records in eastern Québec,Canada

Questions: What was the tree species composition of forests prior to European settlement at the northern hardwood range limit in eastern Québec, Canada? What role did human activities play in the changes in forest composition in this region? Location: Northern range limit of northern hardwoods in the Lower St. Lawrence region of eastern Québec, Canada. Methods: We used early land survey records (1846–1949) of public lands to reconstruct pre‐settlement forest composition. The data consist of ranked tree species enumerations at points or for segments along surveyed lines, with enumerations of forest cover types and notes concerning disturbances. An original procedure was developed to weigh and combine these differing data types (line versus point observations; taxa versus cover enumerations). Change to present‐day forest composition was evaluated by comparing survey records with forest decadal surveys conducted by the government of Québec over the last 30 years (1980–2009). Results: Pre‐settlement dominance of conifers was strong and uniform across the study area, whereas dominance of maple and birches was patchy. Cedar and spruce were less likely to dominate with increasing altitude, whereas maple displayed the reverse trend. Frequency of disturbances, especially logging and fire, increased greatly after 1900. Comparison of survey records and modern plots showed general increases for maple (mentioned frequency increased by 39%), poplar (36%) and paper birch (31%). Considering only taxa ranked first by surveyors, cedar displayed the largest decrease (19%), whereas poplar (15%) and maple (9%) increased significantly. Conclusions: These changes in forest composition can be principally attributed to clear‐cutting and colonization fire disturbances throughout the 20th century, and mostly reflected the propensity of taxa to expand (maples/aspen) or decline (cedar/spruce) with increased disturbance frequency. Québec's land survey archives provide an additional data source to reconstruct and validate our knowledge of North America's pre‐settlement temperate and sub‐boreal forests.     

Isolation and diversity of culturable rhizobacteria associated with economically important crops and uncultivated plants in Québec,Canada

Plants are chronically associated with microorganisms, residing all tissues. Holonomic analysis of diversity of established rhizobacteria in uncultivated plants is scarce. Thus, the present study was conducted to access the root-associated bacterial diversity of 6 crops (maize, canola, soybean, reed canarygrass, alfafa, and miscanthus) and 20 uncultivated plant species in the region of Sainte-Anne-de-Bellevue, Québec, Canada, using pure-culture methods. Based on 16S rRNA gene sequence analysis, 446 bacterial isolates were distributed onto four phyla (Proteobacteria, Firmicutes, Actinobacteria and Bacteroidetes), 32 families and 90 genera. Proteobacteria constituted the largest group of isolates (240), 40% of ectophytic and 61% of endophytic bacteria. Representatives of the genera Bacillus and Pseudomonas dominated in rhizosphere soil; Microbacterium and Pseudomonas were the predominant endophytes. Some genera were associated with specific plant species, such as Stenotrophomonas, Yersinia, Labrys and Luteibacter. Several endophytes were occasionally observed in the rhizosphere, and vice versa. This is the first survey of culturable endophytic bacteria associated with uncultivated plants in Québec. The culturable bacterial community studied herein are assumed to represent a portion of the entire phytomicrobiome of the evaluated plants. Results confirmed that the crops and uncultivated plants of Québec represent an extremely rich reservoir of diverse rhizobacteria.     

Effects of climate on occurrence and size of large fires in a northern hardwood landscape: historical trends,forecasts, and implications for climate change in Témiscamingue,Québec

Questions: What climate variables best explain fire occurrence and area burned in the Great Lakes‐St Lawrence forest of Canada? How will climate change influence these climate variables and thereby affect the occurrence of fire and area burned in a deciduous forest landscape in Témiscamingue, Québec, Canada? Location: West central Québec and the Great Lakes‐St Lawrence forest of Canada. Methods: We first used an information‐theoretic framework to evaluate the relative role of different weather variables in explaining occurrence and area burned of large fires (>200 ha, 1959‐1999) across the Great Lakes‐St Lawrence forest region. Second, we examined how these weather variables varied historically in Témiscamingue and, third, how they may change between the present and 2100 according to different scenarios of climate change based on two Global Circulation Models. Results: Mean monthly temperature maxima during the fire season (Apr‐Oct) and weighted sequences of dry spells best explained fire occurrence and area burned. Between 1910 and 2004, mean monthly temperature maxima in Témiscamingue showed no apparent temporal trend, while dry spell sequences decreased in frequency and length. All future scenarios show an increase in mean monthly temperature maxima, and one model scenario forecasts an increase in dry spell sequences, resulting in a slight increase in forecasted annual area burned. Conclusion: Despite the forecasted increase in fire activity, effects of climate change on fire will not likely affect forest structure and composition as much as natural succession or harvesting and other disturbances, principally because of the large relative difference in area affected by these processes.     

Invasion of American native lily populations by an alien beetle

The lily leaf beetle (Lilioceris lilii), indigenous to Eurasia, was first detected in North America on the island of Montréal, Québec, Canada in 1943. Populations remained limited to the original site of introduction but since 1978 the distribution of this species has expanded in all directions. To date L. lilii has only been recorded feeding on cultivated lilies in urban and rural areas. But as the beetle spreads its geographic range it could encounter native lily species. The objectives of this study were to determine the suitability of two indigenous North American lily species (Lilium canadense and L. philadelphicum) as host for L. lilii, and to assess the degree of infestation of wild lily populations. Measurement of fitness parameters under laboratory conditions (survival, development time, size) indicated that L. lilii has the capacity to develop on both species of native lilies. In the provinces of Québec and Ontario eight of the 20 wild populations of L. canadense sampled were infested. In contrast, L. philadelphicum was not attacked by the alien beetle. A number of native lilies are already endangered in North America, so there is an urgent necessity to address the ecological impact of L. lilii in natural ecosystems.