Male-biased mutation rates and the overestimation of extrapair paternity: problem, solution, and illustration using thick-billed murres (Uria lomvia, Alcidae)

The widespread utility of hypervariable loci in genetic studies derives from the high mutation rate, and thus the high polymorphism, of these loci. Recent evidence suggests that mutation rates can be extremely high and may be male biased (occurring in the male germ-line). These two factors combined may result in erroneous overestimates of extrapair paternity, since legitimate offspring with novel alleles will have more mismatches with respect to the biological father than the biological mother. As mutations are male driven, increasing the number of hypervariable loci screened may simply increase the number of mismatches between fathers and their legitimate offspring. Here we describe a simple statistic, the probability of resemblance (PR), to distinguish between mismatches due to parental misassignment versus mutation in either sex or null alleles. We apply this method to parentage data on thick-billed murres (Uria lomvia), and demonstrate that, without considering either mutations or male-biased mutation rates, cases of extrapair paternity (7% in this study) would be grossly overestimated (14.5%-22%). The probability of resemblance can be utilized in parentage studies of any sexually reproducing species when allele or haplotype frequency data are available for putative parents and offspring. We suggest calculating this probability to correctly categorize legitimate offspring when mutations and null alleles may cause mismatches.     

Population genetic structure in Atlantic and Pacific Ocean common murres (Uria aalge): natural replicate tests of post‐Pleistocene evolution

Understanding the factors that influence population differentiation in temperate taxa can be difficult because the signatures of both historic and contemporary demographics are often reflected in population genetic patterns. Fortunately, analyses based on coalescent theory can help untangle the relative influence of these historic and contemporary factors. Common murres (Uria aalge) are vagile seabirds that breed in the boreal and low arctic waters of the Northern Hemisphere. Previous analyses revealed that Atlantic and Pacific populations are genetically distinct; however, less is known about population genetic structure within ocean basins. We employed the mitochondrial control region, four microsatellite loci and four intron loci to investigate population genetic structure throughout the range of common murres. As in previous studies, we found that Atlantic and Pacific populations diverged during the Pleistocene and do not currently exchange migrants. Therefore, Atlantic and Pacific murre populations can be used as natural replicates to test mechanisms of population differentiation. While we found little population genetic structure within the Pacific, we detected significant east–west structuring among Atlantic colonies. The degree that population genetic structure reflected contemporary population demographics also differed between ocean basins. Specifically, while the low levels of population differentiation in the Pacific are at least partially due to high levels of contemporary gene flow, the east–west structuring of populations within the Atlantic appears to be the result of historic fragmentation of populations rather than restricted contemporary gene flow. The contrasting results in the Atlantic and Pacific Oceans highlight the necessity of carefully considering multilocus nonequilibrium population genetic approaches when reconstructing the demographic history of temperate Northern Hemisphere taxa.     

Philopatry, kin clusters, and genetic relatedness in a population of woodrats (Neotoma macrotis)

Studies of highly kin-structured mammal societies have revealedthe importance of natal philopatry in determining the distributionof genetic variation within populations. In comparison, therelationship between philopatry and genetic diversity withinpopulations of moderately kin-structured societies has receivedrelatively little attention. Previous studies of Neotoma macrotishave suggested that females form distinct kin clusters. Eachkin cluster overlaps spatially with the home range(s) of oneor more males that are not related to each other or to the femaleswith which they are spatially associated. To examine interactionsbetween philopatry and genetic structure in this apparentlymoderately kin-structured species, we characterized spatialand genetic relationships among individually marked femalesin a population of N. macrotis from central coastal California.Our field studies revealed that, contrary to expectation, femalesin this population were not strongly philopatric and spatiallyclustered females were not characterized by high levels of geneticrelatedness. Nevertheless, genetic structure was evident withinthe study population; spatial and genetic distances among femaleswere significantly correlated, suggesting that dispersal patternsinfluenced genetic structure even in the absence of marked femalephilopatry. Because females with overlapping spatial distributionswere not typically closely related to one another, opportunitiesfor the evolution of kin-selected social behavior (e.g., cooperativecare of young) appear to be limited in this population.     

Trophic specialization in Arctic charr Salvelinus alpinus (Pisces; Salmonidae): morphological divergence and ontogenetic niche shifts

Lake Thingvallavatn supports four trophic morphs of Arctic charr, Salvelinus alpinus (L.); two of the morphs are benthic (small and large benthivorous charr) one exploits pelagic waters (planktivorous charr) and the fourth is found in both habitats (piscivorous charr). The morphological variation among these morphs was analysed by use of principal component analysis and canonical discriminant analysis. The benihic morphs have a short lower jaw and long pectoral fins. The benthic fish also have fewer gillrakers than the other morphs. Small and large benthivorous charrs attain sexual maturity from 2 and 6 years of age, and at fork lengths from 7 and 22 cm, respectively. Small benthivorous charr retain their juvenile parr marks as adults, have beige ventral colours, and are frequently melanized under the lower jaw. Planktivorous and piscivorous charr attain sexual maturity from 4 and 6 years of age, from fork lengths of 15 and 23 cm, respectively. This phenotypic polymorphism is associated with habitat utilization and diet of the fish, and has probably arisen within the lake system through diversification and niche specialization. The pelagic morphs apparently stem from a single population, and are possibly diversified through conditional niche shifts which affect ontogeny. Juveniles reaching a body length of 23 cm may change from zooplankton to fish feeding. Asymptotic length increases thereby from 20.5 cm in planktivorous charr to 30.2 cm in piscivorous charr. The benthic morphs appear to represent separate populations, although both feed chiefly on the gastropod Lymnaea peregra. Their co-existence seems to be facilitated by size dependent constraints on habitat use. The small morph (asymptotic length 13.3 cm) exploit the interstitial crevices in the lava block substratum, whereas the large morph (asymptotic length 55.4 cm) live epibenthically.     

Indirect genetic effects in a sex‐limited trait: the case of breeding time in red‐billed gulls

Female reproductive performance can be strongly affected by male care, so that breeding time, a trait expressed only by females, can be seen as one trait determined by both male and female genotypes. Animal model analyses of a 46‐year study of red‐billed gulls (Larus novaehollandiae scopulinus) revealed that laying date was not heritable in females (h² = 0.001 ± 0.030), but significantly so in males (h² = 0.134 ± 0.029). Heritability of breeding time in males probably reflects genetic variability in some other trait such as courtship feeding ability. In line with predictions of evolutionary models incorporating indirect genetic effects, the strong and consistent directional selection for advanced breeding time has not resulted in detectable selection response in males. Our results demonstrate that a female trait is largely determined by genetic characteristics of its mate, and hence, any evolutionary change in red‐billed gull breeding time depends critically on genetic variation in males.     

The Impact of male reproductive skew on kin structure and sociality in multi‐male groups

Patterns of within‐group relatedness are expected to affect the prospects for cooperation among group members through kin selection. It has long been established that dispersal patterns determine the availability of kin and there is ample evidence of matrilineal kin biases in social behavior across primate species. However, in 1979, Jeanne Altmann¹ suggested that mating patterns also influence the structure of within‐group relatedness; high male reproductive skew and the frequent replacement of breeding males leads to relatively high levels of paternal relatedness and age‐structured paternal sibships within groups. As a consequence of frequent replacement of breeding males, relatedness among offspring of a given female will be reduced to the half‐ rather than full‐sibling level. Depending on the number of sires and degree of relatedness among mothers, members of the same birth cohort may be as closely related as maternal siblings. If animals are able to recognize their paternal kin and exhibit biases in favor of them, this may influence the distribution of cooperation and the intensity of competition within groups of primates. Here, I summarize the evidence that serves as the basis for Altmann's predictions and review evidence regarding whether or not the availability of paternal kin also leads to paternal kin bias among primates.     

Population status of Common Guillemots Uria aalge at a colony in western Alaska: results and simulations

Colony-wide boat-based censuses suggest that numbers of Common Guillemots at Bluff, Alaska declined markedly in the late 1970s and remained low until 1983. Annual land-based counts of study plots since 1979 generally showed no trend. However, numbers declined substantially in one study area between 1978 and 1983. Reproductive success varied considerably among years and was highest in moderately warm summers. Simulation analyses suggest that the changes in numbers cannot be attributed to changes in reproductive success several years previously. Thus, factors affecting survival, presumably in winter, are apparently implicated in the declines.     

Division of labour and colony efficiency in social insects: effects of interactions between genetic architecture, colony kin structure and rate of perturbations

The efficiency of social insect colonies critically depends on their ability to efficiently allocate workers to the various tasks which need to be performed. While numerous models have investigated the mechanisms allowing an efficient colony response to external changes in the environment and internal perturbations, little attention has been devoted to the genetic architecture underlying task specialization. We used artificial evolution to compare the performances of three simple genetic architectures underlying within-colony variation in response thresholds of workers to five tasks. In the 'deterministic mapping' system, the thresholds of individuals for each of the five tasks is strictly genetically determined. In the second genetic architecture ('probabilistic mapping'), the genes only influence the probability of engaging in one of the tasks. Finally, in the 'dynamic mapping' system, the propensity of workers to engage in one of the five tasks depends not only on their own genotype, but also on the behavioural phenotypes of other colony members. We found that the deterministic mapping system performed well only when colonies consisted of unrelated individuals and were not subjected to perturbations in task allocation. The probabilistic mapping system performed well for colonies of related and unrelated individuals when there were no perturbations. Finally, the dynamic mapping system performed well under all conditions and was much more efficient than the two other mapping systems when there were perturbations. Overall, our simulations reveal that the type of mapping between genotype and individual behaviour greatly influences the dynamics of task specialization and colony productivity. Our simulations also reveal complex interactions between the mode of mapping, level of within-colony relatedness and risk of colony perturbations.     

Finding your mate in a seabird colony: contrasting strategies of the Guillemot Uria aalge and King Penguin Aptenodytes patagonicus

Capsule King Penguins recognize their mates by voice, but Guillemots do not need acoustic cues even though their calls show individual variation.

Aims To determine whether the structure of Guillemot calls could allow individual recognition, as with King Penguin, and whether acoustic cues are used to locate mates among a dense mass of conspecifics at a colony.

Methods Observations were made on breeding Guillemots and King Penguins. Calls made by birds returning to their mates were recorded, the signals digitized and the calls analysed. Calls were later played back to the mates of the birds concerned and the effects noted on both them and their neighbours.

Results Both Guillemots and King Penguins emitted calls on return to the breeding site which contained individual signatures and were therefore potentially usable for mate recognition. In King Penguins, auditory recognition was essential for finding a mate, whereas in Guillemots most of the arriving birds located their mate in a dense crowd of conspecifics without the help of acoustic signals. Guillemots could differentiate neighbours from strangers without auditory cues.

Conclusion Calls are essential for the successful identification of mates by King Penguins but not by Guillemots.     

Arctic fisheries catches in Russia, USA, and Canada: baselines for neglected ecosystems

The Amerasian Arctic, covering northern Siberia (Russia), Arctic Alaska (USA), and the Canadian Arctic, extends over seven coastal Large Marine Ecosystems and makes up the seasonally ice-free part of FAO Statistical Area 18 (Arctic Sea). Historically, the harsh climate has limited marine fisheries (here excluding marine mammal hunting) to small-scale operations conducted mainly in estuaries and river deltas. Most of the catches have traditionally not been reported to FAO, with the result that total catch estimated here from 1950 to 2006 is 75 times higher than the sum of the catches reported for FAO 18 (Arctic). Catches were reconstructed from data on fishing communities in Canada and Alaska, and from various government and non-government sources for Siberia. Based on national data supplied to FAO since 1950, catches have been reported as 12,700 t in toto (reported on behalf of the former Soviet Union). This compares with our reconstructed catches of over 950,000 t, i.e., 770,000, 89,000, and 94,000 t by Russia, USA, and Canada, respectively for the same time period. The reconstructed catch (mainly whitefishes in Siberia, various salmonids in Alaska, and Arctic char in Canada) was 24,100 t year⁻¹ in 1950, but declined to 10,200 t year⁻¹ by the mid-2000s. Reasons for these trends are discussed by country, along with sources of uncertainty (particularly large for Siberia). Catches were allocated to Large Marine Ecosystems to present ecosystem-relevant baselines for the impact of fisheries on the Arctic, which can be expected to increase, as industrial fisheries move into a warming Arctic following the invasion of boreal species, unless countries apply precautionary ecosystem-based management approaches.     

Ecological niche specialization inferred from morphological variation and otolith strontium of Arctic charr Salvelinus alpinus L. found within open lake systems of southern Baffin Island, Nunavut, Canada

The presence of two morphotypes of Arctic charr Salvelinus alpinus was confirmed via morphological variation and otolith strontium (Sr) within three open-lake systems of southern Baffin Island, Nunavut, Canada: Qinngu (LH001), Iqalugaarjuit Lake (PG082) and Qasigiat (PG015). Analysis of otolith Sr indicates that a component of each S. alpinus population within lakes LH001 and PG082 is migratory (large–maturing S. alpinus ), whereas another component is lake-resident (small–maturing S. alpinus ). Alternatively, small and large maturing S. alpinus may both inhabit tidal habitats during their lifetime in lake PG015. Three morphological characters were identified by principal factor analysis (PFA) as characters that were different between maturity groups for all lakes studied: eye diameter, pectoral fin length and pelvic fin length. As well, upper jaw length (LH001 and PG082) and fork depth (PG015) were identified in PFA as traits that differed between morphs. Univariate tests of morphological characters identified by PFA demonstrated maturity group differences with the exception of eye diameter in Lake PG015 and upper jaw length and pelvic fin length in lake LH001. No difference was found in the MANOVA test of upper and lower gill raker number between small–maturing and undeveloped fish within all lakes studied. Clear morphological variation observed between small–maturing and undeveloped fish in all three lakes of the study suggests ecological niche separation between morphotypes. This is the first documented case of lake-resident S. alpinus use of the tidal habitat in the presence of a migratory large–maturing morphotype.     

Population genetic structuring of the king weakfish, Macrodon ancylodon (Sciaenidae), in Atlantic coastal waters of South America: deep genetic divergence without morphological change

Phylogeographic patterns in Macrodon ancylodon sampled from 12 locations across all its range were investigated using mitochondrial DNA cytochrome b sequences, and analysed together with patterns of morphometric differentiation. Populations of the North Brazil and the Brazil currents, with warmer waters, form a clade (tropical clade) separated by 23 fixed mutations from the populations that inhabit regions of colder waters influenced by the Brazil and Malvinas currents (subtropical clade). No gene flow exists between the tropical and subtropical clades, and most likely also between the two groups of the tropical clade. Distribution of these clades and groups is correlated with flow of currents and their temperatures, and is facilitated by larval retention and low adult migration. Despite differentiation at the molecular level, fishes analysed from all these current-influenced regions are morphometrically homogeneous. Throughout its range M. ancylodon inhabits the same, or very similar niche; thus, stabilizing selection probably promotes the retention of highly conserved morphology despite deep genetic divergence at the mitochondrial DNA cytochrome b.     

Analogous aquatic and terrestrial food webs in the high Arctic: The structuring force of a harsh climate

Understanding food web structure and dynamics remains a central theme in ecology. Whilst differences between aquatic and terrestrial food webs have been the focus of several studies, we aim to reveal similarities where abiotic conditions are particularly extreme such as in the high Arctic. We propose that here, the combination of a short growing season, low temperature and low light, leads to the absence of predator control and the development of typically two-trophic, grazer-dominated food webs with high plant quality in terms of elemental ratios. Moreover, we advocate that this mechanism is evident in both aquatic and terrestrial high-Arctic environments, allowing the build-up of herbivore densities that consume a large fraction of plant primary production and tightly recycle nutrients. Thus, the particular abiotic conditions that characterise the high Arctic give rise to a unique environment that allows biotic factors to orchestrate food web structure and influence ecosystem function. Specially, the short growing season, low temperatures and low light levels collectively constrain the accumulation of structural autotroph tissue that, in temperate regions, effectively keeps herbivores at bay. While fundamental differences between terrestrial and aquatic ecosystems have been frequently advocated, we show here that harsh live-constraining conditions in the high Arctic have led to analogous, grazer-dominated, food web dynamics in both terrestrial and freshwater ecosystems.     

Predator‐induced morphological defences as by‐products of prey behaviour: a review and prospectus

Predator‐induced morphological defences (PIMDs) are ubiquitous. Many PIMDs may be mediated by prey behaviour rather than directly cued by predators. A survey of 92 studies indicated 40 that quantified prey behaviour, all of which document positive associations between defence production and activity reduction. Thus, PIMDs are associated with changes in prey activity, which could have caused the morphological change. We propose two possible mechanisms: 1) decreased activity reduces feeding rate, resulting in lower growth and morphological change; and 2) activity reduction conserves energy, which is reallocated for growth, subsequently changing morphology. Resource availability also causes similar morphological change to predator presence, suggesting confounding effects of resources and predators with current methodology. Future studies should estimate food ingestion, assimilation efficiency, and growth rate in the presence and absence of predators, crossing predator presence with resource levels. Not all PIMDs will be behaviourally‐mediated, but consideration of causal linkages between prey behaviour and PIMDs is warranted.     

Population history of Berthelot's pipit: colonization, gene flow and morphological divergence in Macaronesia

The fauna of oceanic islands provide exceptional models with which to examine patterns of dispersal, isolation and diversification, from incipient speciation to species level radiations. Here, we investigate recent differentiation and microevolutionary change in Berthelot's pipit (Anthus berthelotii), an endemic bird species inhabiting three Atlantic archipelagos. Mitochondrial DNA sequence data and microsatellite markers were used to deduce probable colonization pathway, genetic differentiation, and gene flow among the 12 island populations. Phenotypic differentiation was investigated based on eight biologically important morphological traits. We found little mitochondrial DNA variability, with only one and four haplotypes for the control region and cytochrome b, respectively. However, microsatellite data indicated moderate population differentiation (FST=0.069) between the three archipelagos that were identified as genetically distinct units with limited gene flow. Both results, combined with the estimated time of divergence (2.5 millions years ago) from the Anthus campestris (the sister species), suggest that this species has only recently dispersed throughout these islands. The genetic relationships, patterns of allelic richness and exclusive alleles among populations suggest the species originally colonized the Canary Islands and only later spread from there to the Madeiran archipelago and Selvagen Islands. Differentiation has also occurred within archipelagos, although to a lesser degree. Gene flow was observed more among the eastern and central islands of the Canaries than between these and the western islands or the Madeiran Islands. Morphological differences were also more important between than within archipelagos. Concordance between morphological and genetic differentiation provided ambiguous results suggesting that genetic drift alone was not sufficient to explain phenotypic differentiation. The observed genetic and morphological differences may therefore be the result of differing patterns of selection pressures between populations, with Berthelot's pipit undergoing a process of incipient differentiation.     

Genetic, morphological, and feather isotope variation of migratory willow warblers show gradual divergence in a ring

The circular distribution of the willow warbler Phylloscopus trochilus around the Baltic Sea shares many features with the classic examples of ring species; however, the system is much younger. It has previously been shown that a secondary contact zone is located in central Scandinavia, where there are narrow clines for several morphological traits coincident with a migratory divide. Here we analyse multiple traits and genes from > 1700 males captured on breeding territories at 77 sites spread around the Baltic Sea to test the following hypothesis. If the secondary contact zone in Scandinavia is a result of divergence in two allopatric refuge populations during the last glaciation, we expect to find a similar secondary contact zone somewhere else around the circular distribution. Our results show that the trait clines were wider and displaced from each other along the eastern side of the Baltic Sea. Analyses of 12 microsatellite loci confirmed that the genome is very similar between the terminal forms ( F _ST = 0). Two AFLP-derived markers filtered out from a genomic scan instead appear to be maintained by selection. These markers exhibited steep clines at the secondary contact zone in Scandinavia, but as for the phenotypic traits, had vastly different cline centres east of the Baltic Sea. The trait clines along the ring distribution outside the Scandinavian secondary contact zone thus seem to have been shaped by independent action of selection or drift during the process of postglacial colonization.     

Philopatry in the alpine grasshopper, Podisma pedestris: a novel experimental and analytical method

Abstract.

• 1 Local dispersal and philopatric behaviour of the alpine grasshopper, Podisma pedestris, were studied at two sites in the Alpes Maritimes using a new mark—release—resight technique featuring marking in situ, and multiple resighting without handling. The time-consuming nature of the design was justified by the quantity and quality of the data yielded.
• 2 Philopatry is defined independently of any concept of home range as a phenomenon by which movements over longer periods are less than would be expected by extrapolation of measures over shorter periods.
• 3 By this definition, the grasshoppers are shown to exhibit philopatry. Nymphs comprise two heterogenous classes of those which move little and those which move considerable distances.
• 4 Daily dispersal estimates were obtained from movements of 1055 nymphs and adults at one site, and of 593 adults at another.
• 5 Differences are demonstrated in daily dispersal distances for age, sex and site combinations.
• 6 Some differences in microhabitat preference were observed.
• 7 The findings are compared with previous measurements of dispersal in this species, and the merits of the new technique are discussed.