Density-mediated responses of bark beetles to host allelochemicals: a link between individual behaviour and population dynamics

Abstract  1. Bark beetles (Coleoptera: Scolytidae) accept or reject host conifers based partly on concentrations of phloem monoterpenes. They colonise trees in aggregations, in response to pheromones that attract flying beetles to trees undergoing colonisation. A series of entry and gallery construction assays was conducted to determine whether responses by individual beetles to monoterpenes are altered by pheromones and/or the presence of other beetles.
2. Entry into the amended media by Ips pini and the length of time until entry were not influenced by the presence of aggregation pheromones.
3. Entry into amended media was influenced by the presence of other beetles on the surface of, or constructing galleries in, the substrate. The effects of alpha-pinene and limonene on host entry behaviour were mediated by the density of beetles on the surface of the assay arena, and by the density of beetles constructing galleries within the medium.
4. The percentage of beetles entering medium amended with higher concentrations of monoterpenes increased with increased density of beetles on the surface of the assay arena, until a threshold density of three or four beetles per assay arena, after which entrance rate declined.
5. The presence of other beetles constructing galleries elicited more rapid entry by the test beetles.
6. Gallery lengths were generally higher in the presence of aggregation pheromones.
7. Gallery lengths increased with increased density of beetles within the assay arena.
8. These results suggest a link between the density of bark beetles and responses of individuals. This linkage may partially explain behavioural changes observed during population eruptions.     

Sex‐specific effects of inbreeding in wild‐caught Drosophila melanogaster under benign and stressful conditions

In animal populations, sib mating is often the primary source of inbreeding depression (ID). We used recently wild‐caught Drosophila melanogaster to test whether such ID is amplified by environmental stress and, in males, by sexual selection. We also investigated whether increased ID because of stress (increased larval competition) persisted beyond the stressed stage and whether the effects of stress and sexual selection interacted. Sib mating resulted in substantial cumulative fitness losses (egg to adult reproduction) of 50% (benign) and 73% (stressed). Stress increased ID during the larval period (23% vs. 63%), but not during post‐stress reproductive stages (36% vs. 31%), indicating larval stress may have purged some adult genetic load (although ID was uncorrelated across stages). Sexual selection exacerbated inbreeding depression, with inbred male offspring suffering a higher reproductive cost than females, independent of stress (57% vs. 14% benign, 49% vs. 11% stress).     

Variation of selfing rate and inbreeding depression among individuals and across generations within an admixed Cedrus population

We investigated the variation and short-term evolution of the selfing rate and inbreeding depression (ID) across three generations within a cedar forest that was established from admixture ca 1860. The mean selfing rate was 9.5%, ranging from 0 to 48% among 20 seed trees (estimated from paternally inherited chloroplast DNA). We computed the probability of selfing for each seed and we investigated ID by comparing selfed and outcrossed seeds within progenies, thus avoiding maternal effects. In all progenies, the germination rate was high (88–100%) and seedling mortality was low (0–12%). The germination dynamics differed significantly between selfed and outcrossed seeds within progenies in the founder gene pool but not in the following generations. This transient effect of selfing could be attributed to epistatic interactions in the original admixture. Regarding the seedling growth traits, the ID was low but significant: 8 and 6% for height and diameter growth, respectively. These rates did not vary among generations, suggesting minor gene effects. At this early stage, outcrossed seedlings outcompeted their selfed relatives, but not necessarily other selfed seedlings from other progenies. Thus, purging these slightly deleterious genes may only occur through within-family selection. Processes that maintain a high level of genetic diversity for fitness-related traits among progenies also reduce the efficiency of purging this part of the genetic load.     

Invasion of sibmating genes in diploid and haplodiploid populations

Summary Existing genetic models of the evolution of sibmating behaviour in diploids incorporate inbreeding depression in terms of reduced fecundity of consanguineous mating pairs rather than reduced survival or fecundity of the progeny of such matings. Here we derive a model to correct this deficiency and extend the model to haplodiploids where differential effects of inbreeding in males and females is a crucial consideration. Our analyses indicate that sibmating can readily evolve in both diploids and haplodiploids in which male mating costs and inbreeding depression are reasonably low, provided there is some mechanism to permit sibmating such as siblings being reared in nests or other forms of aggregation. Our analyses also indicate that once sibmating invades, it typically will go to fixation, although sib-/randommating polymorphisms can persist in both diploids and haplodiploids if male mating costs are close to zero and inbreeding depression reduces survival by around one-third. The conditions favouring sibmating are slightly more restrictive in haplodiploids than in diploids. In light of this we may ask why we see intense sibmating in many haplodiploids such as parasitic wasps, fig wasps, ants, bark beetles and mites, and only rarely in diploid animals. The common factor could be certain kinds of aggregation behaviour that are a prerequisite for sibmating in the absence of kin recognition. Another possibility is that inbreeding depression is likely to be more severe in diploids than in haplodiploids because deleterious recessives are purged from haplodiploid populations when expressed by haploid males. Thus, lower levels of inbreeding depression might be one important reason why sibmating appears to arise more frequently in haplodiploids than diploids. Phylogenetic analysis of groups, such as bark beetles and mites, exhibiting both diploid and haplodiploid populations may be useful in elucidating the relative importance of gregarious behaviour and haplodiploidy in facilitating sibmating systems.     

Evolution of Inbreeding and Outcrossing in Ferns and Fern-Allies

Abstract Mating systems of 18 species of homosporous ferns follow a bimodal distribution, similar to that observed for seed plants (Schemske and Lande, 1985). Most species are highly outcrossing, a few are inbreeding, and two species examined to date have mixed mating systems. Equisetum arvense and several species of lycopods are also highly outcrossing. Several mechanisms, including inbreeding depression, antheridiogen, and ontogenetic sequences that result in effectively unisexual gametophytes, promote outcrossing in homosporous ferns and perhaps other homosporous pteridophytes as well. In some species of homosporous ferns, selection has favored the evolution of inbreeding as an adaptation for colonization. High levels of intra- and interpopulational gene flow via spore dispersal, coupled with high levels of intergametophytic crossing, generally lead to genetically homogeneous populations and species of homosporous ferns. However, rock-dwelling ferns and ferns from xeric habitats may exhibit significant population genetic structure due to physically patchy habitats. Reticulate evolution in homosporous ferns may be enhanced by high levels of intergametophytic crossing.     

Drift load in populations of small size and low density

According to theory, drift load in randomly mating populations is determined by past population size, because enhanced genetic drift in small populations causes accumulation and fixation of recessive deleterious mutations of small effect. In contrast, segregating load due to mutations of low frequency should decline in smaller populations, at least when mutations are highly recessive and strongly deleterious. Strong local selection generally reduces both types of load. We tested these predictions in 13 isolated, outcrossing populations of Arabidopsis lyrata that varied in population size and plant density. Long-term size was estimated by expected heterozygosity at 20 microsatellite loci. Segregating load was assessed by comparing performance of offspring from selfings versus within-population crosses. Drift load was the heterosis effect created by interpopulation outbreeding. Results showed that segregating load was unrelated to long-term size. However, drift load was significantly higher in populations of small effective size and low density. Drift load was mostly expressed late in development, but started as early as germination and accumulated thereafter. The study largely confirms predictions of theory and illustrates that mutation accumulation can be a threat to natural populations.     

INBREEDING AND VARIANCE EFFECTIVE SIZES FOR NONRANDOM MATING POPULATIONS

Following an inbreeding approach and assuming discrete generations and autosomal inheritance involving genes that do not affect viability or reproductive ability, I have derived expressions for the inbreeding effective size, N_eI, for a finite diploid population with variable census sizes for three cases: monoecious populations with partial selfing; dioecious populations of equal numbers of males and females with partial sib mating; and unequal numbers of males and females with random mating. For the first two cases, recurrence equations for the inbreeding coefficient are also obtained, which allow inbreeding coefficients to be predicted exactly in both early and late generations. Following the variance of change in gene frequency approach, a general expression for variance effective size, N_eV, is obtained for a population with unequal numbers of male and female individuals, arbitrary family size distribution, and nonrandom mating. All the parameters involved are allowed to change over generations. For some special cases, the equation reduces to the simple expressions approximately as derived by previous authors. Comparisons are made between equations derived by the present study and those obtained by previous authors. Some of the published equations for N_eI and N_eV are shown to be incomplete or incorrect. Stochastic simulations are run to check the results where disagreements with others are involved.     

Prolonged development time of the bark beetle predator Thanasimus formicarius (Col.: Cleridae) in relation to its prey species Tomicus piniperda (L.) and Ips typographus (L.) (Col.: Scolytidae)

1 The generation time of the bark beetle predator Thanasimus formicarius (L.) (Col.: Cleridae) was found to be predominantly two years both in the field and in rearing experiments conducted with two of its main prey species, the pine shoot beetle Tomicus piniperda (L.) and the spruce bark beetle Ips typographus (L.) (Col.: Scolytidae). 2 Emergence of T. formicarius adults in the first summer was only observed in one of the two rearing experiments, and these individuals represented only 6% of that generation. 3 All individuals not emerging as adults in the first summer remained as larvae in their pupal chambers until the second summer. Pupae were found starting around mid-June, and adults (in pupal chambers) were found from late July through to the end of August. 4 Newly emerged adults had to feed in order to survive hibernation. 5 The existence of T. formicarius races, specialized on certain bark beetle species and with phenologies matching their hosts, could not be demonstrated. After hibernation there was no difference in feeding activity, timing of egg-laying or proportion of egg-laying females between the T. formicarius adults reared as larvae on T. piniperda (flight period in April) and those reared as larvae on I. typographus (main flight period generally starting in late May or early June) when exposed to a temperature and day-length typical of the early spring conditions prevailing during the flight period of T. piniperda. 6 T. formicarius was parasitized by Enclisis vindex (Tschek) (Hym.: Ichneumonidae) in the pupal chamber. 7 The importance of these findings for the population dynamics of bark beetles is discussed.     

Inbreeding depression in self-incompatible North-American Arabidopsis lyrata: disentangling genomic and S-locus-specific genetic load

Newly formed selfing lineages may express recessive genetic load and suffer inbreeding depression. This can have a genome-wide genetic basis, or be due to loci linked to genes under balancing selection. Understanding the genetic architecture of inbreeding depression is important in the context of the maintenance of self-incompatibility and understanding the evolutionary dynamics of S-alleles. We addressed this using North-American subspecies of Arabidopsis lyrata. This species is normally self-incompatible and outcrossing, but some populations have undergone a transition to selfing. The goals of this study were to: (1) quantify the strength of inbreeding depression in North-American populations of A. lyrata; and (2) disentangle the relative contribution of S-linked genetic load compared with overall inbreeding depression. We enforced selfing in self-incompatible plants with known S-locus genotype by treatment with CO₂, and compared the performance of selfed vs outcrossed progeny. We found significant inbreeding depression for germination rate (δ=0.33), survival rate to 4 weeks (δ=0.45) and early growth (δ=0.07), but not for flowering rate. For two out of four S-alleles in our design, we detected significant S-linked load reflected by an under-representation of S-locus homozygotes in selfed progeny. The presence or absence of S-linked load could not be explained by the dominance level of S-alleles. Instead, the random nature of the mutation process may explain differences in the recessive deleterious load among lineages.     

Low genetic variation reduces cross-compatibility and offspring fitness in populations of a narrow endemic plant with a self-incompatibility system

Genetic variation was shown earlier to bereduced in smaller populations of the narrowendemic putatively self-incompatible Cochlearia bavarica. To test whether thisnegatively affects plant fitness by reducedavailability of compatible mates and byinbreeding depression, we studied effects ofpopulation size and pollination treatments oncross-compatibility and offspring fitness in 16isolated populations of this plant. After openpollination, compatibility of crosses (i.e.,whether at least one fruit developed per markedflower), fruit set of compatible crosses, andcumulative fitness (number of plants permaternal ovule) after 14 months in a commongarden were lower for plants from smallerpopulations. Throughout the study, cumulativefitness was lower after hand pollination withpollen of one donor than after open pollination(finally 73.4% lower), suggesting that severalpollen donors or single pollen donors of higherquality are involved in open pollination.Moreover, cumulative fitness was lower afterhand selfing than after hand outcrossing(finally 69.4% lower), indicating bothinbreeding depression and reduced compatibilityafter selfing. High self-compatibility(40.6%), dry stigmas, and differences in thecompatibility of 11 of 33 experimentalreciprocal crosses between plant pairsconfirmed that C. bavarica has asporophytic self-incompatibility system, as iscommon in the Brassicaceae. Our studydemonstrates, that plants in smallerpopulations of species with a sporophyticself-incompatibility system can experiencetwofold fitness reductions associated withreduced genetic variability, i.e., twofoldgenetic Allee effects: via reducedcross-compatibility and via reduced offspringfitness.