Multiple geographic origins and high genetic differentiation of the Alpine marmots reintroduced in the Pyrenees

Reintroductions inherently involve a small number of founders leading reintroduced populations to be prone to genetic drift and, consequently, to inbreeding depression. Assessing the origins as the genetic diversity and structure of reintroduced populations compared to native populations are thus crucial to foresee their future. Here, we aim to clarify the origins of the Alpine marmots reintroduced in the Pyrenees and to evaluate the genetic consequences of this reintroduction after almost 30 years without monitoring. We search for the origins and compare the genetic structure and the genetic variability of three reintroduced Pyrenean and eight native Alpine populations using pairwise genetic distances, Bayesian clustering method and multivariate analyses. Our results reveal that the Alpine marmots reintroduced in the Pyrenees originated both from the Northern and the Southern Alps, and that, despite these multiple origins, none of the current Pyrenean marmots are admixed. The reintroduction led to a strong genetic differentiation and to a decrease in genetic diversity. This pattern likely results from the small number of founders and the low dispersal capacities of Alpine marmots and thus, highlight the necessity to consider both genetic characteristics and natural history when reintroducing a species.     

Social structure and facultative mating systems of hoary marmots (Marmota caligata)

Mate-choice theory predicts different optimal mating systems depending on resource availability and habitat stability. Regions with limited resources are thought to promote monogamy. We tested predictions of monogamy in a social rodent, the hoary marmot (Marmota caligata), at the northern climatic extreme of its distribution. Mating systems, social structure and genetic relationships were investigated within and among neighbouring colonies of marmots within a 4 km(2) valley near Kluane National Park, Yukon, Canada, using 21 microsatellite loci. While both monogamous and polygynous populations of hoary marmots have been observed in the southern reaches of this species' range; northern populations of this species are thought to be predominantly monogamous. Contrary to previous studies, we did not find northern hoary marmot social groups to be predominantly monogamous; rather, the mating system seemed to be facultative, varying between monogamy and polygyny within, as well as among, social groups. These findings reveal that the mating systems within colonies of this species are more flexible than previously thought, potentially reflecting local variation in resource availability.     

Matrilineal genetic structure within and among populations of the cooperatively breeding common marmoset,Callithrix jacchus

Common marmosets are members of the family Callitrichidae, South American primates characterized by highly social group living and cooperative breeding. In this study we analysed 1112 base pairs (bp) of the mitochondrial control region in 59 Callithrix jacchus individuals, sampled mainly from two geographically distinct field sites in N.E. Brazil. Analysis of molecular variation revealed a highly significant genetic structuring of haplotypes between social groups and between populations. Examination of matrilineal genetic structure within social groups revealed that seven of nine recorded breeding pairs were from different maternal lineages, indicating assortative mating and outbreeding. In addition to the breeders, at least six of 10 groups contained adult individuals from different matrilines, with five haplotypes present in one social group of nine animals. Groups of mixed lineages raise questions about potential reproductive conflicts of interest, and the extent of kin-selected altruism in the evolution and maintenance of cooperative breeding in this species.     

Genetic discontinuity, breeding-system change and population history of Arabis alpina in the Italian Peninsula and adjacent Alps

Arabis alpina is a widespread plant of European arctic and alpine environments and belongs to the same family as Arabidopsis thaliana. It grows in all major mountain ranges within the Italian glacial refugia and populations were sampled over a 1300 km transect from Sicily to the Alps. Diversity was studied in nuclear and chloroplast genome markers, combining phylogeographical and population genetic approaches. Alpine populations had significantly lower levels of nuclear genetic variation compared to those in the Italian Peninsula, and this is associated with a pronounced change in within-population inbreeding. Alpine populations were significantly inbred (F(IS) = 0.553), possibly reflecting a change to the self-incompatibility system during leading edge colonization. The Italian Peninsula populations were approaching Hardy-Weinberg equilibrium (outbreeding, F(IS) = 0.076) and genetic variation was highly structured, consistent with independent local 'refugia within refugia' and the fragmentation of an established population by Quaternary climate oscillations. There is very little evidence of genetic exchange between the Alps and the Italian Peninsula main distribution ranges. The Alps functioned as a glacial sink for A. alpina, while the Italian Peninsula remains a distinct and separate long-term refugium. Comparative analysis indicated that inbreeding populations probably recolonized the Alps twice: (i) during a recent postglacial colonization of the western Alps from a Maritime Alps refugium; and (ii) separately into the central Alps from a source outside the sampling range. The pronounced geographical structure and inbreeding discontinuities are significant for the future development of A. alpina as a model species.     

Social behaviour and individual differences in free-living Alpine marmots (Marmota marmota)

I studied the behaviour of free-living Alpine marmots, Marmota marmota, from 1 June to 19 August 1973 in Vanoise National Park, France. Comparing animals of the same sex and age class, individual differences in behaviour generally were greater between colonies than within colonies. Individual differences among adults exceeded those of yearlings, and greater individual differences occurred with regard to social behaviour than with relatively asocial behaviour. General patterns of social biology resembled those of the North American marmots, exceptions being a tendency for adult males to remain somewhat away from and at higher elevation than their colonies, and a distinctly lower frequency of greeting behaviour. Implications of these findings for the evolution of marmot social behaviour are discussed.     

Ectoparasites,Fitness, and Social Behaviour of Yellow-Bellied Marmots

Parasites can cause a loss of fitness for their hosts, potentially influencing social behaviour patterns of the host that promote or hinder parasite transmission. I studied yellow-bellied marmots (Marmota flaviventris) and their ectoparasites to determine if ectoparasites reduce the fitness of marmots and to test whether ectoparasite loads differ according to social behaviour. Three taxa of ectoparasites were identified, fleas (Oropsylla Stanfordi), lice (Linognathoides marmotae), and mites (family Dermanyssidae). High ectoparasite loads were related to slower growth, lower overwinter survival, and reduced reproduction, suggesting that ectoparasites are a fitness cost for marmots. Ectoparasite loads were not higher in colonial than in noncolonial marmots, nor in polygynous than in monogamous adult males. There was a trend, however, toward higher ectoparasite loads in marmots that were dispersing rather than philopatric. Further, ectoparasite loads differed among groups of marmots that nested or hibernated communally, indicating that spatial scale is important in understanding the relationships between parasites and social behaviour.     

Early and adult social environments have independent effects on individual fitness in a social vertebrate

Evidence that the social environment at critical stages of life-history shapes individual trajectories is accumulating. Previous studies have identified either current or delayed effects of social environments on fitness components, but no study has yet analysed fitness consequences of social environments at different life stages simultaneously. To fill the gap, we use an extensive dataset collected during a 24-year intensive monitoring of a population of Alpine marmots (Marmota marmota), a long-lived social rodent. We test whether the number of helpers in early life and over the dominance tenure length has an impact on litter size at weaning, juvenile survival, longevity and lifetime reproductive success (LRS) of dominant females. Dominant females, who were born into a group containing many helpers and experiencing a high number of accumulated helpers over dominance tenure length showed an increased LRS through an increased longevity. We provide evidence that in a wild vertebrate, both early and adult social environments influence individual fitness, acting additionally and independently. These findings demonstrate that helpers have both short- and long-term effects on dominant female Alpine marmots and that the social environment at the time of birth can play a key role in shaping individual fitness in social vertebrates.     

Paternity and dominance loss in male breeders: the cost of helpers in a cooperatively breeding mammal

Paternity insurance and dominance tenure length are two important components of male reproductive success, particularly in species where reproduction is highly skewed towards a few individuals. Identifying the factors affecting these two components is crucial to better understand the pattern of variation in reproductive success among males. In social species, the social context (i.e. group size and composition) is likely to influence the ability of males to secure dominance and to monopolize reproduction. Most studies have analyzed the factors affecting paternity insurance and dominance tenure separately. We use a long term data set on Alpine marmots to investigate the effect of the number of subordinate males on both paternity insurance and tenure of dominant males. We show that individuals which are unable to monopolize reproduction in their family groups in the presence of many subordinate males are likely to lose dominance the following year. We also report that dominant males lose body mass in the year they lose both paternity and dominance. Our results suggest that controlling many subordinate males is energetically costly for dominant males, and those unable to support this cost lose the control over both reproduction and dominance. A large number of subordinate males in social groups is therefore costly for dominant males in terms of fitness.     

Socio-spatial Organization and Activity Distribution of the Alpine Marmot Marmota marmota: Preliminary Results

The behavioural ecology of the Alpine marmot Marmota marmota (Linné, 1758) was studied Jun.—Sep. 1990 in the Vanoise National Park (French Alps). We describe the socio-spatial structure of a high-altitude population, to give additional information on the formerly unclear marmot social organization. The social unit was the family group, with a common home range between 0.9 and 2.8 ha; a slight overlapping occurred (9 to 12.5%). Each home range had a central area of main burrows, where the hibernaculum was located, and peripheral areas. Space utilization and distribution of activities during the season were analysed. Some activities took place exclusively in the centre area while peripheral parts were used for foraging. This latter increased in Aug. and decreased in Sep., marmots tending to remain more and more at the centre area as the hibernation period approached.     

Nouvelles données sur le déroulement de la mycose àNeozygites fumosa sur la cochenille du maniocPhenacoccus manihoti

Résumé Cet article présente les résultats obtenus en 1987 sur des points non encore abordés précédemment dans le cadre de l'étude des circonstances épidémiologiques favorisant le déroulement de l'Entomophthorose àNeozygites fumosa (Speare) Remaudière et Keller, pathogène de la cochenille du maniocPhenacoccus manihoti Matile-Ferrero (Hom.: Pseudococcidae). Il montre notamment que l'évolution de la maladie, très rapide, correspond à la succession de 2 phases: une première phase, d'implantation, fortement liée à la taille et à la structure des colonies et une deuxième phase (épizootique au sens strict) indépendante de leur taille et de leur structure. Son évolution appara?t plus liée à la régularité des pluies qu'à la quantité d'eau. Les conditions sont très favorables quand l'humidité relative est supérieure à 90% pendant au moins 5 heures par jour de fa?on régulière. Le r?le important joué par la durée d'humectation du feuillage est montré ici pour la première fois. La densité des conidies du pathogène dans l'air est proportionnelle aux taux de mycose observés dans les populations deP. manihoti.      

Lateglacial and Holocene vegetation history in the mountain range of central Pyrenees

We present the pollen analysis of a new sedimentary sequence taken at La Pouretère ( 1720 m), in the mountain vegetation zone of the Marcadau valley (central Pyrenees). The Lateglacial and Holocene chronology is supported by six 14C-dating results. The complementary analysis of some vegetal macroremains, stomata, pollen-clusters and the use of pollen influx allows us to elucidate the dynamic of mountain species such as Pinus and specially Abies but also to infer the unusual part played by Betula at the beginning of the Postglacial period.     

Préférence de ponte et niveaux d’adaptation de différentes souches de Caryedon serratus Ol. (Coleoptera: Bruchidae) à l’arachide (Arachis hypogaea L., Fabaceae)

Résumé

La sélection du substrat de ponte par des souches de la bruche de l’arachide, Caryedon serratus, issues d’hôtes différents est étudiée au laboratoire. Des tests de choix sur des gousses de différentes plantes ont été d’abord effectués. Les distributions des oeufs sur les différents substrats de ponte sont comparées. Les niveaux d’adaptation des souches dites sauvages (Bauhinia rufescens, Cassia sieberiana, Piliostigma reticulatum et Tamarindus indica) sur l’arachide et les capacités de développement de la souche ? arachide ? sur les graines des plantes hôtes ? sauvages ? sont aussi testés. Les résultats révèlent que, d’une manière générale, les souches évoluent selon le principe de sélection d’hôtes de Hopkins puisque la ponte se fait de manière privilégiée sur la gousse de la plante d’où est issue la femelle. La souche ? Piliostigma ? dépose ses oeufs de manière indifférenciée sur gousses de P. reticulatum et sur gousses d’arachide. Les résultats des tests sur les niveaux d’adaptation révèlent du point de vue de la durée de développement, qu’on peut considérer les différentes souches comme identiques quant à leurs possibilités intrinsèques de développement sur l’arachide. Le paramètre poids des adultes, montre que la souche ? Bauhinia ? est intrinsèquement moins bien adaptée que les autres souches sur l’arachide. Les taux fécondité révèlent que les différentes souches sont identiques quant à leurs possibilités intrinsèques de développement sur l’arachide mais, ici également, la souche ? Bauhinia ? paraît moins bien adaptée à l’arachide. Le passage de la souche ? arachide ? aux plantes hôtes sauvages est possible mais, élevée sur graines plantes hôtes sauvages, celle-ci se comporte généralement moins bien que sur arachide.     

Mate choice for neutral and MHC genetic characteristics in Alpine marmots: different targets in different contexts?

Sexual selection through female mate choice for genetic characteristics has been suggested to be an important evolutionary force maintaining genetic variation in animal populations. However, the genetic targets of female mate choice are not clearly identified and whether female mate choice is based on neutral genetic characteristics or on particular functional loci remains an open question. Here, we investigated the genetic targets of female mate choice in Alpine marmots (Marmota marmota), a socially monogamous mammal where extra‐pair paternity (EPP) occurs. We used 16 microsatellites to describe neutral genetic characteristics and two MHC loci belonging to MHC class I and II as functional genetic characteristics. Our results reveal that (1) neutral and MHC genetic characteristics convey different information in this species, (2) social pairs show a higher MHC class II dissimilarity than expected under random mate choice, and (3) the occurrence of EPP increases when social pairs present a high neutral genetic similarity or dissimilarity but also when they present low MHC class II dissimilarity. Thus, female mate choice is based on both neutral and MHC genetic characteristics, and the genetic characteristics targeted seem to be context dependent (i.e., the genes involved in social mate choice and genetic mate choice differ). We emphasize the need for empirical studies of mate choice in the wild using both neutral and MHC genetic characteristics because whether neutral and functional genetic characteristics convey similar information is not universal.     

Endémisme insulaire et cleptoparasitisme chez les Lasioglossum Curtis 1833 (Hymenoptera : Apoidea : Halictidae) de Nouvelle-Calédonie

Huit espèces de Lasioglossum sont inventoriées de Nouvelle-Calédonie. Un nouveau sous-genre cleptoparasite endémique, avec une nouvelle espèce, Lasioglossum (Parachilalictus n. subg. ) neocaledonicum n. sp., est décrit. Les espèces endémiques suivantes du sous-genre Chilalictus sont nouvelles : Lasioglossum (Chilalictus) alticola n. sp., L. (C.) tchambae n. sp. et L. (C.) webbi n. sp. Une nouvelle sous-espèce, Lasioglossum (Chilalictus) polygoni austrocaledonicum n. ssp., est décrite de Nouvelle-Calédonie et d'Australie. Les espèces australiennes L. (Chilalictus) lanarium (Smith 1853) et L. (C.) instabilis (Cockerell 1914) sont découvertes pour la première fois en Nouvelle-Calédonie. Lasioglossum (Parasphecodes) sulthicum (Smith 1853) est la première espèce de ce sous-genre australien signalé en Nouvelle-Calédonie. L’émergence du parasitisme et la spéciation en condition insulaire font l'objet d'une discussion. Le cleptoparasitisme du nouveau sous-genre Parachilalictus n. subg. peut être établi en examinant un ensemble de caractères morphologiques convergents chez les groupes parasites de Halictidae dont les principaux sont la réduction de la brosse collectrice de pollen sur les pattes postérieures et l'aplatissement de la quille du labre. La forte ressemblance de Parachilalictus n. subg. avec les espèces de Chilalictus endémiques de Nouvelle-Calédonie et son absence en Australie suggère une spéciation locale et récente. Les deux nouvelles espèces endémiques décrites dans le sous-genre Chilalictus et dont Parachilalictus n. subg. pourrait avoir émergé ont une distribution montagnarde (L. alticola n. sp. , L. tchambae n. sp.). Dans le complexe L. polygoni (Cockerell 1929), la spéciation s'exprime par la variation de la densité et la force de la ponctuation. A basse altitude et le long de la côte ouest plus sèche, on trouve la sous-espèce typique polygoni. A des altitudes variant de 5 à 1150m, il existe clairement deux autres sous-espèces. La nouvelle sous-espèce austrocaledonica ne se trouve que dans la partie sud de l’île, en milieu ouvert de « maquis » tandis que la sous-espèce delobeli Pauly & Munzinger 2003 a une distribution plus étendue sur les massifs montagneux de la partie est, et a été rencontrée essentiellement le long de pistes forestières en forêt dense humide. Cette dernière sous-espèce habite aussi l'Australie.     

Révision des Xylocopinae (Hymenoptera : Apidae) de France et de Belgique

Cette publication présente les résultats d’une étude de 6.263 spécimens de Xylocopinae récoltés en France métropolitaine (France continentale et Corse) et en Belgique et qui proviennent de récoltes personnelles et de 63 institutions et collections privées. Les Xylocopinae de France sont représentées par 11 espèces du genre Ceratina et 4 espèces du genre Xylocopa. Seules deux de ces espèces sont présentes en Belgique : Xylocopa violacea (L.) et Ceratina cyanea (Kirby). Un nouveau sous-genre pour le genre Ceratina est décrit : Dalyatina n. subg. Il comporte l’espèce méditerranéenne Ceratina parvula Smith présente en France, ainsi que six autres espèces d’Afrique subsaharienne. Pour chaque genre, sous-genre et espèce, les auteurs fournissent une diagnose, une diagnose différentielle, la liste des fleurs visitées, la liste des sites de nidification, la carte de distribution en France métropolitaine, un diagramme phénologique et une clé d’identifi cation des genres, sous-genres et espèces. La systématique, la biogéographie, l’écologie et le sexe ratio des espèces sont présentés et discutés. Les Xylocopinae apparaissent comme très largement polylectiques mais montrent une très nette affinité envers les Lamiaceae, les Asteraceae Cardueae et, pour le genre Xylocopa, envers les Fabaceae. Toutes les espèces présentent une phénologie estivale qui s’étend d’avril à septembre. Le sexe-ratio de la plupart des espèces est biaisé vers les femelles. Aucun mâle de C. parvula, pour 120 femelles, n’a été observé ce qui suggère que, en France du moins, l’espèce pourrait se reproduire par parthénogenèse thélytoque comme c’est le cas de C. dallatorreana Friese. La publication comprend 62 dessins au trait, 18 photos au microscope électronique à balayage, 17 cartes, 14 graphiques de phénologie, une liste de 232 espèces de fleurs visitées par les Xylocopinae, dont 176 observations originales, et 171 références.     

Outbreeding effects in an inbreeding insect,Cimex lectularius

In some species, populations with few founding individuals can be resilient to extreme inbreeding. Inbreeding seems to be the norm in the common bed bug, Cimex lectularius, a flightless insect that, nevertheless, can reach large deme sizes and persist successfully. However, bed bugs can also be dispersed passively by humans, exposing inbred populations to gene flow from genetically distant populations. The introduction of genetic variation through this outbreeding could lead to increased fitness (heterosis) or be costly by causing a loss of local adaptation or exposing genetic incompatibility between populations (outbreeding depression). Here, we addressed how inbreeding within demes and outbreeding between distant populations impact fitness over two generations in this re‐emerging public health pest. We compared fitness traits of families that were inbred (mimicking reproduction following a founder event) or outbred (mimicking reproduction following a gene flow event). We found that outbreeding led to increased starvation resistance compared to inbred families, but this benefit was lost after two generations of outbreeding. No other fitness benefits of outbreeding were observed in either generation, including no differences in fecundity between the two treatments. Resilience to inbreeding is likely to result from the history of small founder events in the bed bug. Outbreeding benefits may only be detectable under stress and when heterozygosity is maximized without disruption of coadaptation. We discuss the consequences of these results both in terms of inbreeding and outbreeding in populations with genetic and spatial structuring, as well as for the recent resurgence of bed bug populations.     

Reproductive suppression in female Alpine marmots, Marmota marmota

We studied mechanisms responsible for reproductive skew in female Alpine marmots. In each social group, only dominant females produced weaned young although subordinate females had similar body condition, were fertile as indicated by high levels of oestradiol during the mating season, and occasionally started pregnancies. During the period of gestation, dominant females initiated significantly more agonistic interactions against subordinate females, resulting in significantly increased levels of glucocorticoids and decreased levels of progesterone in subordinates. Results suggested that reproductive suppression in female Alpine marmots is mediated by the negative effects of stress (glucocorticoids) on the activity of the hypothalamic-pituitary-gonadal axis. The strength of competition between subordinate and dominant females was affected by their relatedness. Dominant females attacked unrelated subordinate females more, whereas amicable behaviour was observed mainly between dominant females and their daughters. These differences could be explained by differences in indirect fitness: related subordinate females benefited from warming the offspring of the dominant female during hibernation. Reproductive suppression was apparently costly for dominant females, because their reproductive success decreased as the number of adult subordinate females in a group increased. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

On the increase of chromosome mutations under random mating.

After a short introduction on karyotypes and chromosome mutations, we review the ways by which a chromosome mutation can increase in a random mating population, despite the mutation's deleterious effect on the fertility of heterozygotes. Random drift, segregation distortion, viability advantage, and recombination modification are the mechanisms considered. When possible, the models are illustrated with examples of chromosome mutations involving autosomes in mammals, but the arguments apply, of course, to any genetic factor in any outbreeding species that causes a fertility decrease in heterozygotes.     

Large-scale genotyping of highly polymorphic loci by next-generation sequencing: how to overcome the challenges to reliably genotype individuals?

Studying the different roles of adaptive genes is still a challenge in evolutionary ecology and requires reliable genotyping of large numbers of individuals. Next-generation sequencing (NGS) techniques enable such large-scale sequencing, but stringent data processing is required. Here, we develop an easy to use methodology to process amplicon-based NGS data and we apply this methodology to reliably genotype four major histocompatibility complex (MHC) loci belonging to MHC class I and II of Alpine marmots (Marmota marmota). Our post-processing methodology allowed us to increase the number of retained reads. The quality of genotype assignment was further assessed using three independent validation procedures. A total of 3069 high-quality MHC genotypes were obtained at four MHC loci for 863 Alpine marmots with a genotype assignment error rate estimated as 0.21%. The proposed methodology could be applied to any genetic system and any organism, except when extensive copy-number variation occurs (that is, genes with a variable number of copies in the genotype of an individual). Our results highlight the potential of amplicon-based NGS techniques combined with adequate post-processing to obtain the large-scale highly reliable genotypes needed to understand the evolution of highly polymorphic functional genes.