Differentiation of Alpine marmot populations traced by DNA fingerprinting.

As revealed by allozyme studies, the genetic variation of the Alpine marmot (Marmota m. marmota) has been reduced by a species-wide bottleneck at the end of the last glaciation. Therefore the more variable microsatellite loci were used as a genetic marker system to investigate variablility and differentiation of four autochthonous and four allochthonous populations founded by the release of small numbers of individuals during the last 150 years. The microsatellite loci detected by the DNA-probe (ATCC)₄ were found to be polymorphic in all populations, but the amount of variation was lower than in comparable mammalian species. In spite of founder effects the variation in the allochthonous populations was not significanlty reduced compared to the autochthonous populations. The autochthonous populations from Austria and from the eastern part of Switzerland were genetically similar, only the population from western Switzerland was clearly differentiated from the others. In the allochthonous populations similarities in the microsatellite patterns reveal genetic affinities to putative autochthonous source populations of the founder individuals.     

Sex ratio variation in the cooperatively breeding alpine marmot Marmota marmota

In cooperatively breeding vertebrate species, a clear theoreticalprediction about the direction of sex ratio adjustment can bemade: mothers should bias the sex ratio of their offspring towardsthe helping sex when helpers are absent. A consistent trendin the direction predicted by theory exists in cooperative birds,but theory is still poorly tested in cooperative mammals. Here,multivariate analyses are applied to a long-term data set totest this prediction in two ways in the alpine marmot: (1) acrossfemales in a population and (2) in individual females acrossmultiple years. It was shown that in the alpine marmot offspringsex ratio was biased towards the helping sex (males) when helperswere absent, whereas helped mothers produced unbiased sex ratio.Unhelped mothers did not adjust the litter size but producedmore sons and fewer daughters than helped mothers. These resultssupport the theoretical prediction and explain well the malebias observed among juvenile alpine marmots at the populationlevel. The occurrence of possible sex ratio manipulations incooperatively breeding vertebrates is also discussed.     

Microsatellite variation in autochthonous and introduced populations of the Alpine marmot (Marmota marmota) along a European west–east transect

Microsatellite variation was studied in 11 populations of the Alpine marmot along a west–east transect through the present distribution range. The samples represent five autochthonous and six introduced populations. Eleven loci were analysed in nine populations and six loci in the two populations from France. In the populations from the Western Alps, there is no indication for reduced variability as has been assumed in previous studies. However, a decrease of variation in the autochthonous populations was observed from the west to the east. The introduced populations showed a heterogeneous pattern reflecting the geographic origin of the released individuals. The population from the Spanish Pyrenees harbours a high level of variation and is genetically closest to the French populations. In Austria, three of the introduced populations have low variation and are closely related to the autochthonous populations from the western part of Austria. In contrast, two introduced populations from the central part of Austria are highly variable and resemble the populations from France. At least for one of these populations an early introduction of founder individuals from the Western Alps has been documented.     

Is there an optimal number of helpers in Alpine marmot family groups?

The consequence of helping behavior on breeders fitness is stillcontroversial. We used multivariate analyses to investigatefor the effects of male and female subordinates on breeders'components of fitness in the Alpine marmot, Marmota marmota.We found that male and female subordinates, respectively, increasedand decreased juvenile survival during winter. Thus, we giveevidence that male subordinates should be considered as helpers,and that helpers provided breeders with immediate reproductivesuccess gains, whereas subordinates females were costly. Helpershad no positive effects on female body condition, on persistence(future survival) of dominants, and on future reproduction (occurrenceand size of a litter). Helpers thus did not provide breederswith delayed fitness benefits, and therfore, the load-lighteninghypothesis was not supported. On the contrary, helpers had delayedfitness cost for dominant males and, consequently, for dominantfemales. Immediate benefits counterbalanced by delayed costssuggested an optimal number of helpers in the family group bothfrom male and female perspectives. An optimality model wellpredicted the observed mean number of helpers in Alpine marmotfamily groups. Optimal numbers of helpers were slightly differentfor males and females, suggesting a potential conflict of interestbetween dominants. We finally discuss the possible mechanismsof helping that may explain the observed pattern in the Alpinemarmot.     

Male-biased sex ratio in litters of Alpine marmots supports the helper repayment hypothesis

In a French population of Alpine marmots (Marmota marmota),the sex ratio at weaning was biased in favor of males. Thisbias also seemed to exist at birth. Under Fisher's equal allocationprinciple, this means that daughters should be more costlyto produce than sons. Because the Alpine marmot can be considereda cooperative breeding species, we investigated whether thedifferential cost between sons and daughters may be explainedby the helper repayment hypothesis. The Alpine marmot usessocial thermoregulation during hibernation, allowing juvenilesto better survive over winter. In the study population, juvenilesurvival during winter increased with group size. More precisely,juvenile survival during winter increased with the number andwith the proportion of subordinate males in the hibernatinggroup, but juvenile survival did not depend on the number of subordinate females. As our results did not support alternativehypotheses to explain the observed bias in sex ratio amongoffspring at emergence, we conclude that the helper repaymenthypothesis is the best candidate to explain the observed offspringsex ratio bias in Alpine marmots. By participating in socialthermoregulation, subordinate males may repay part of the investment they received from their parents and thus become less costlyto produce. We suggest that only subordinate males helped becausethey may gain direct fitness benefits, whereas subordinatefemales may only expect indirect fitness benefits from helping.Finally, the offspring sex ratio per individual parent wasmale biased, but mothers adjusted the size and the sex compositionof their litters according to their phenotypic condition asexpected from the Trivers-Willard hypothesis.     

Male-caused failure of female reproduction and its adaptive value in alpine marmots (Marmota marmota)

We studied reproductive performance of free-living alpine marmots (Marmotamarmota) for 14 years in the National Park of Berchtesgaden, Germany.Female reproduction was influenced by body condition and social factors.Reproduction depleted fat reserves, and only females emergingfrom hibernation with sufficient body mass were able to reproducesuccessfully. Marmots lived in social groups in territoriesdefended by a dominant male and female. Subordinate femalesnever reproduced, regardless of body mass. Territory takeoversby males impaired reproduction of dominant females, but onlyif the takeover occurred after the mating period. Reproductivefailures occurred despite clear signs of pregnancy such enlargednipples or late molt. Decreasing progesterone levels after themating period and the lack of evidence for direct infanticideby new territorial males suggest a block of pregnancy as a likelyexplanation for reproductive failures in groups with male takeoversduring gestation. Rendering female reproduction impossible increasedfuture reproductive success of new territory owners. Nonparous femalessaved the energetic cost of maternal investment and thus emergedwith higher body mass in the following spring. In line withthis, females failing to wean young had higher reproductivesuccess in the subsequent year.     

Plucked hair samples as a source of DNA: reliability of dinucleotide microsatellite genotyping

To test whether plucked hairs are a reliable source of DNA for genotyping microsatellite loci, we carried out experiments using one, three, or 10 hairs per extract for 50 alpine marmots. For each extract, seven independent genotypings were performed for the same locus (multiple-tubes approach). Two types of genotyping errors were recorded: a false homozygote defined as the detection of only one allele of a true heterozygote, and a false allele defined as a PCR-generated allele that was not one of the alleles of the true genotype. Using DNA extracted from one, three, or 10 hairs, the overall error rate was 14.00%, 4.86%, and 0.29%, respectively. Based on our results, we conclude that 10 hairs should be used to obtain consistently reliable genotypings using the single-tube approach, and that a single plucked hair could represent a reliable source of DNA if the multiple-tubes approach is used. For future studies of dinucleotide repeat diversity using DNA extracted from one to three shed or plucked hairs, we strongly recommend initiating an appropriate pilot study to quantify the error rate and to determine the reliability of the single-tube approach.     

Global warming and positive fitness response in mountain populations of common lizards Lacerta vivipara

Recent global warming threatens many species and has already caused population‐ and species‐level extinctions. In particular, high risks of extinction are expected for isolated populations of species with low dispersal abilities. These predictions rely on widely used ‘climatic envelope’ models, while individual responses, the ultimate driver of a species response to climate change, have been most often neglected. Here, we report on some changes in life‐history traits of a dispersal‐limited reptile species (a poorly studied taxa) living in isolated populations. Using long‐term data on common lizards collected in southern France, we show that individual body size dramatically increased in all the four populations studied over the past 18 years. This increase in body size in all age classes appeared related to a concomitant increase in temperature experienced during the first month of life (August). Daily maximum temperature in August increased by 2.2°C and yearling snout‐vent‐length increased by about 28%. As a result, adult female body size increased markedly, and, as fecundity is strongly dependent on female body size, clutch size and total reproductive output also increased. For one population where capture–recapture data were available, adult survival was positively related to May temperature. All fitness components investigated therefore responded positively to the increase in temperature, such that it might be concluded that the common lizard has been advantaged by the shift in temperature. We contrast these short‐term results with the long‐term habitat‐based prediction that these populations located close to mountain tops on the southern margin of the species range should be unable to cope with the alteration of their habitat. To achieve a better prediction of a species persistence, one will probably need to combine both habitat and individual‐based approaches.     

Intriguing asexual life in marginal populations of the brown seaweed Fucus vesiculosus

Reproduction of attached large brown algae is known to occur only by sexual zygotes. Using microsatellites we show evolution of asexual reproduction in the bladder wrack promoting population persistence in the brackish water Baltic Sea (< 6 psu). Here a dwarf morph of Fucus vesiculosus is dominated by a single clone but clonal reproduction is also present in the common form of the species. We describe a possible mechanism for vegetative reproduction of attached algae, and conclude that clonality plays an important role in persistence and dispersal of these marginal populations, in which sexual reproduction is impaired by low salinity.