High levels of inbreeding with spatial and host-associated structure in lice of an endangered freshwater seal.

Host-specialist parasites of endangered large vertebrates are in many cases more endangered than their hosts. In particular, low host population densities and reduced among-host transmission rates are expected to lead to inbreeding within parasite infrapopulations living on single host individuals. Furthermore, spatial population structures of directly-transmitted parasites should be concordant with those of their hosts. Using population genomic approaches, we investigated inbreeding and population structure in a host-specialist seal louse (Echinophthirius horridus) infesting the Saimaa ringed seal (Phoca hispida saimensis), which is endemic to Lake Saimaa in Finland, and is one of the most endangered pinnipeds in the world. We conducted genome resequencing of pairs of lice collected from 18 individual Saimaa ringed seals throughout the Lake Saimaa complex. Our analyses showed high genetic similarity and inbreeding between lice inhabiting the same individual seal host, indicating low among-host transmission rates. Across the lake, genetic differentiation among individual lice was correlated with their geographic distance, and assignment analyses revealed a marked break in the genetic variation of the lice in the middle of the lake, indicating substantial population structure. These findings indicate that movements of Saimaa ringed seals across the main breeding areas of the fragmented Lake Saimaa complex may in fact be more restricted than suggested by previous population-genetic analyses of the seals themselves.

SealID: Saimaa Ringed Seal Re-Identification Dataset.

Wildlife camera traps and crowd-sourced image material provide novel possibilities to monitor endangered animal species. The massive data volumes call for automatic methods to solve various tasks related to population monitoring, such as the re-identification of individual animals. The Saimaa ringed seal (Pusa hispida saimensis) is an endangered subspecies only found in Lake Saimaa, Finland, and is one of the few existing freshwater seal species. Ringed seals have permanent pelage patterns that are unique to each individual and that can be used for the identification of individuals. A large variation in poses, further exacerbated by the deformable nature of seals, together with varying appearance and low contrast between the ring pattern and the rest of the pelage makes the Saimaa ringed seal re-identification task very challenging, providing a good benchmark by which to evaluate state-of-the-art re-identification methods. Therefore, we make our Saimaa ringed seal image (SealID) dataset (N = 57) publicly available for research purposes. In this paper, the dataset is described, the evaluation protocol for re-identification methods is proposed, and the results for two baseline methods-HotSpotter and NORPPA-are provided. The SealID dataset has been made publicly available.

Mercury and Selenium Balance in Endangered Saimaa Ringed Seal Depend on Age and Sex.

The endangered Saimaa ringed seal (Pusa hispida saimensis) is exposed to relatively high concentrations of mercury (Hg) in freshwaters poor in selenium (Se), a known antagonist of Hg. The impact of age and sex on the bioaccumulation of Hg and Se was studied by analyzing liver, muscle, and hair samples from seals of different age groups. Adult females were found to accumulate significantly more Hg in the liver (with ca. 60% as HgSe), and less Hg in the muscles compared to adult males, which may be explained by accelerated metabolism during gestation and lactation. In adult seals, molar Se:Hg ratios in the muscles fall below one, which is considered a threshold for the emergence of adverse effects. As a result, Saimaa ringed seals may be at risk of developing health and reproductive problems. According to mass balance calculations, the pups are exposed to considerable amounts (µg/d) of mercury during gestation, although lactation is their main exposure route. In lanugo pups, Hg concentrates in the hair, and molting serves as a main detoxification route. For other age groups, demethylation followed by the formation of HgSe is the main detoxification route, and the demethylation capability develops in pups by the time of weaning.

Causes and consequences of fine-scale population structure in a critically endangered freshwater seal.

BACKGROUND: Small, genetically uniform populations may face an elevated risk of extinction due to reduced environmental adaptability and individual fitness. Fragmentation can intensify these genetic adversities and, therefore, dispersal and gene flow among subpopulations within an isolated population is often essential for maintaining its viability. Using microsatellite and mtDNA data, we examined genetic diversity, spatial differentiation, interregional gene flow, and effective population sizes in the critically endangered Saimaa ringed seal (Phoca hispida saimensis), which is endemic to the large but highly fragmented Lake Saimaa in southeastern Finland. RESULTS: Microsatellite diversity within the subspecies (HE = 0.36) ranks among the lowest thus far recorded within the order Pinnipedia, with signs of ongoing loss of individual heterozygosity, reflecting very low effective subpopulation sizes. Bayesian assignment analyses of the microsatellite data revealed clear genetic differentiation among the main breeding areas, but interregional structuring was substantially weaker in biparentally inherited microsatellites (FST = 0.107) than in maternally inherited mtDNA (FST = 0.444), indicating a sevenfold difference in the gene flow mediated by males versus females. CONCLUSIONS: Genetic structuring in the population appears to arise from the joint effects of multiple factors, including small effective subpopulation sizes, a fragmented lacustrine habitat, and behavioural dispersal limitation. The fine-scale differentiation found in the landlocked Saimaa ringed seal is especially surprising when contrasted with marine ringed seals, which often exhibit near-panmixia among subpopulations separated by hundreds or even thousands of kilometres. Our results demonstrate that population structures of endangered animals cannot be predicted based on data on even closely related species or subspecies.

Osteological profiling of femoral diaphysis and neck in aquatic, semiaquatic, and terrestrial carnivores and rodents: effects of body size and locomotor habits.

The increased limb bone density documented previously for aquatic tetrapods has been proposed to be an adaptation to overcome buoyancy during swimming and diving. It can be achieved by increasing the amount of bone deposition or by reducing the amount of bone resorption, leading to cortical thickening, loss of medullary cavity, and compaction of trabecular bone. The present study examined the effects of locomotor habit, body size, and phylogeny on the densitometric, cross-sectional, and biomechanical traits of femoral diaphysis and neck in terrestrial, semiaquatic, and aquatic carnivores, and in terrestrial and semiaquatic rodents (12 species) by using peripheral quantitative computed tomography, three-point bending, and femoral neck loading tests. Groupwise differences were analyzed with the univariate generalized linear model and the multivariate linear discriminant analysis supplemented with hierarchical clustering. While none of the individual features could separate the lifestyles or species adequately, the combinations of multiple features produced very good or excellent classifications and clusterings. In the phocid seals, the aquatic niche allowed for lower femoral bone mineral densities than expected based on the body mass alone. The semiaquatic mammals mostly had high bone mineral densities compared to the terrestrial species, which could be considered an adaptation to overcome buoyancy during swimming and shallow diving. Generally, it seems that different osteological properties at the levels of mineral density and biomechanics could be compatible with the adaptation to aquatic, semiaquatic, or terrestrial niches.

Seasonal changes in diel haul-out patterns of a lacustrine ringed seal (Pusa hispida saimensis).

Seasonal changes in diel haul-out patterns of the lacustrine Saimaa ringed seal (Pusa hispida saimensis) were studied using a combination of satellite telemetry and camera traps during 2007-2015. We found the haul-out activity patterns to vary seasonally. Our results show that during the ice-covered winter period before the seals start their annual molt, the peak in haul-out generally occurs at midnight. Similarly, during the postmolt season of summer and autumn when the lake is free of ice, the haul-out is concentrated in the early hours of the morning. In contrast, over the spring molting period, Saimaa ringed seals tend to haul out around the clock. The spring molt is also the only period when a slight difference in haul-out behavior between the sexes is observed, with females having a haul-out peak at nighttime while the males have a less visible diel pattern. According to our results, the diel haul-out patterns of Saimaa ringed seals are similar to the ones of marine ringed seals. Gathering information on haul-out activity is important in order to safeguard the natural patterns of Saimaa ringed seals in areas that are prone to disturbance from human activities.

Museum specimens of a landlocked pinniped reveal recent loss of genetic diversity and unexpected population connections.

The Saimaa ringed seal (Pusa hispida saimensis) is endemic to Lake Saimaa in Finland. The subspecies is thought to have originated when parts of the ringed seal population of the Baltic region were trapped in lakes emerging due to postglacial bedrock rebound around 9000 years ago. During the 20th century, the population experienced a drastic human-induced bottleneck. Today encompassing a little over 400 seals with extremely low genetic diversity, it is classified as endangered. We sequenced sections of the mitochondrial control region from 60 up to 125-years-old museum specimens of the Saimaa ringed seal. The generated dataset was combined with publicly available sequences. We studied how genetic variation has changed through time in this subspecies and how it is phylogenetically related to other ringed seal populations from the Baltic Sea, Lake Ladoga, North America, Svalbard, and the White Sea. We observed temporal fluctuations in haplotype frequencies and loss of haplotypes accompanied by a recent reduction in female effective population size. In apparent contrast with the traditionally held view of the Baltic origin of the population, the Saimaa ringed seal mtDNA variation also shows affinities to North American ringed seals. Our results suggest that the Saimaa ringed seal has experienced recent genetic drift associated with small population size. The results further suggest that extant Baltic ringed seal is not representative of the ancestral population of the Saimaa ringed seal, which calls for re-evaluation of the deep history of this subspecies.

Loss of species and genetic diversity during colonization: Insights from acanthocephalan parasites in northern European seals.

Studies on host-parasite systems that have experienced distributional shifts, range fragmentation, and population declines in the past can provide information regarding how parasite community richness and genetic diversity will change as a result of anthropogenic environmental changes in the future. Here, we studied how sequential postglacial colonization, shifts in habitat, and reduced host population sizes have influenced species richness and genetic diversity of Corynosoma (Acanthocephala: Polymorphidae) parasites in northern European marine, brackish, and freshwater seal populations. We collected Corynosoma population samples from Arctic, Baltic, Ladoga, and Saimaa ringed seal subspecies and Baltic gray seals, and then applied COI barcoding and triple-enzyme restriction-site associated DNA (3RAD) sequencing to delimit species, clarify their distributions and community structures, and elucidate patterns of intraspecific gene flow and genetic diversity. Our results showed that Corynosoma species diversity reflected host colonization histories and population sizes, with four species being present in the Arctic, three in the Baltic Sea, two in Lake Ladoga, and only one in Lake Saimaa. We found statistically significant population-genetic differentiation within all three Corynosoma species that occur in more than one seal (sub)species. Genetic diversity tended to be high in Corynosoma populations originating from Arctic ringed seals and low in the landlocked populations. Our results indicate that acanthocephalan communities in landlocked seal populations are impoverished with respect to both species and intraspecific genetic diversity. Interestingly, the loss of genetic diversity within Corynosoma species seems to have been less drastic than in their seal hosts, possibly due to their large local effective population sizes resulting from high infection intensities and effective intra-host population mixing. Our study highlights the utility of genomic methods in investigations of community composition and genetic diversity of understudied parasites.

Application of change-point analysis to determine winter sleep patterns of the raccoon dog (Nyctereutes procyonoides) from body temperature recordings and a multi-faceted dietary and behavioral study of wintering.

BACKGROUND: A multi-faceted approach was used to investigate the wintertime ecophysiology and behavioral patterns of the raccoon dog, Nyctereutes procyonoides, a suitable model for winter sleep studies. By utilizing GPS tracking, activity sensors, body temperature (Tb) recordings, change-point analysis (CPA), home range, habitat and dietary analyses, as well as fatty acid signatures (FAS), the impact of the species on wintertime food webs was assessed. The timing of passive bouts was determined with multiple methods and compared to Tb data analyzed by CPA. RESULTS: Raccoon dogs displayed wintertime mobility, and the home range sizes determined by GPS were similar or larger than previous estimates by radio tracking. The preferred habitats were gardens, shores, deciduous forests, and sparsely forested areas. Fields had close to neutral preference; roads and railroads were utilized as travel routes. Raccoon dogs participated actively in the food web and gained benefit from human activity. Mammals, plants, birds, and discarded fish comprised the most important dietary classes, and the consumption of fish could be detected in FAS. Ambient temperature was an important external factor influencing Tb and activity. The timing of passive periods approximated by behavioral data and by CPA shared 91% similarity. CONCLUSIONS: Passive periods can be determined with CPA from Tb recordings without the previously used time-consuming and expensive methods. It would be possible to recruit more animals by using the simple methods of data loggers and ear tags. Hunting could be used as a tool to return the ear-tagged individuals allowing the economical extension of follow-up studies. The Tb and CPA methods could be applied to other northern carnivores.

Seasonal and diel activity patterns of the endangered taiga bean goose (Anser fabalis fabalis) during the breeding season, monitored with camera traps.

Taiga bean goose (Anser fabalis fabalis) is an endangered subspecies that breeds sporadically in remote habitats in the arctic and boreal zones. Due to its elusive behaviour, there is a paucity of knowledge on the behaviour of taiga bean goose during the breeding season, and survey methods for monitoring numbers in the breeding areas are lacking. Camera traps are a useful tool for wildlife monitoring, particularly when there is a need for non-invasive methods due to the shy nature of the species. In this study, we tested the use of camera traps to investigate seasonal and diel activity patterns of taiga bean goose in Finland over two successive breeding seasons, 2018 and 2019. We did this by modelling counts of geese from images with generalized linear and additive mixed models. The camera type (cameras placed by experts specialized in bean goose ecology vs randomly placed cameras) did not influence the count of taiga bean goose (p = 0.386). However, the activity varied significantly by region, Julian day, time of day and temperature, with the study site (individual peatland) and year adding substantial random variation and uncertainty in the counts. Altogether, the best fitting model explained nearly 70% of the variation in taiga bean goose activity. The peak in activity occurred about a month later in the southernmost region compared to the more northern regions, which may indicate behaviours related to migration rather than breeding and moulting. Our results show that long-term monitoring with game camera traps provide a potential unobtrusive approach for studying the behavioural patterns of taiga bean goose and can increase our ecological knowledge of this little-known subspecies. The results can be applied to planning of the annual censuses and finding the optimal time frame for their execution.

Patterns of Microbiome Variation Among Infrapopulations of Permanent Bloodsucking Parasites.

While interspecific variation in microbiome composition can often be readily explained by factors such as host species identity, there is still limited knowledge of how microbiomes vary at scales lower than the species level (e.g., between individuals or populations). Here, we evaluated variation in microbiome composition of individual parasites among infrapopulations (i.e., populations of parasites of the same species living on a single host individual). To address this question, we used genome-resolved and shotgun metagenomic data of 17 infrapopulations (balanced design) of the permanent, bloodsucking seal louse Echinophthirius horridus sampled from individual Saimaa ringed seals Pusa hispida saimensis. Both genome-resolved and read-based metagenomic classification approaches consistently show that parasite infrapopulation identity is a significant factor that explains both qualitative and quantitative patterns of microbiome variation at the intraspecific level. This study contributes to the general understanding of the factors driving patterns of intraspecific variation in microbiome composition, especially of bloodsucking parasites, and has implications for understanding how well-known processes occurring at higher taxonomic levels, such as phylosymbiosis, might arise in these systems.

DNA barcoding reveals different cestode helminth species in northern European marine and freshwater ringed seals.

Three subspecies of the ringed seal (Pusa hispida) are found in northeastern Europe: P. h. botnica in the Baltic Sea, P. h saimensis in Lake Saimaa in Finland, and P. h. ladogensis in Lake Ladoga in Russia. We investigated the poorly-known cestode helminth communities of these closely related but ecologically divergent subspecies using COI barcode data. Our results show that, while cestodes from the Baltic Sea represent Schistocephalus solidus, all worms from the two lakes are identified as Ligula intestinalis, a species that has previously not been reported from seals. The observed shift in cestode communities appears to be driven by differential availability of intermediate fish host species in marine vs. freshwater environments. Both observed cestode species normally infect fish-eating birds, so further work is required to elucidate the health and conservation implications of cestode infections in European ringed seals, whether L. intestinalis occurs also in marine ringed seals, and whether the species is able to reproduce in seal hosts. In addition, a deep barcode divergence found within S. solidus suggests the presence of cryptic diversity under this species name.

Sex, age, and tissue specific accumulation of eight metals, arsenic, and selenium in the European hedgehog (Erinaceus europaeus).

Many insectivores have been shown to be sensitive to heavy metals and therefore suitable for biomonitoring purposes. In Finland, the hibernation period of the European hedgehog (Erinaceus europaeus) is long, and during hibernation the stress caused by environmental toxins may be crucial. Concentrations of cadmium (Cd), copper (Cu), iron (Fe), magnesium (Mg), manganese (Mn), molybdenum (Mo), nickel (Ni), lead (Pb), arsenic (As), and selenium (Se) were measured in a population of hedgehogs in the town of Joensuu in eastern Finland during the summers of 2004 and 2005. The analyzed tissues were kidney, liver, hair, and spine. The sampled hedgehogs (n = 65) were mainly road-killed animals. As expected, the concentrations of heavy metals were low because the hedgehogs were living in a comparatively unpolluted area. Significant increases with age were found in Cd concentrations (kidney, liver, and spine) and some essential elements (Se in spine, kidney, and liver; Mo in kidney and liver; Cu in spine; Fe in liver; and Mn in spine). Age accumulation and correlations between Se and Cd and between Mo and Cd may indicate the protective roles of Se and Mo against Cd toxicity in hedgehogs, in which Cd is already at comparatively low concentrations. Sex had no significant effect on concentrations of the elements studied. In conclusion, age is an important parameter to be taken into account when studying heavy-metal concentrations in hedgehogs and other insectivores.

Determinants of genetic variation across eco-evolutionary scales in pinnipeds.

The effective size of a population (Ne), which determines its level of neutral variability, is a key evolutionary parameter. Ne can substantially depart from census sizes of present-day breeding populations (NC) as a result of past demographic changes, variation in life-history traits and selection at linked sites. Using genome-wide data we estimated the long-term coalescent Ne for 17 pinniped species represented by 36 population samples (total n = 458 individuals). Ne estimates ranged from 8,936 to 91,178, were highly consistent within (sub)species and showed a strong positive correlation with NC ([Formula: see text] = 0.59; P = 0.0002). Ne/NC ratios were low (mean, 0.31; median, 0.13) and co-varied strongly with demographic history and, to a lesser degree, with species' ecological and life-history variables such as breeding habitat. Residual variation in Ne/NC, after controlling for past demographic fluctuations, contained information about recent population size changes during the Anthropocene. Specifically, species of conservation concern typically had positive residuals indicative of a smaller contemporary NC than would be expected from their long-term Ne. This study highlights the value of comparative population genomic analyses for gauging the evolutionary processes governing genetic variation in natural populations, and provides a framework for identifying populations deserving closer conservation attention.

A mark-recapture approach for estimating population size of the endangered ringed seal (Phoca hispida saimensis).

Reliable population estimates are fundamental to the conservation of endangered species. We evaluate here the use of photo-identification (photo-ID) and mark-recapture techniques for estimating the population size of the endangered Saimaa ringed seal (Phoca hispida saimensis). Photo-ID data based on the unique pelage patterns of individuals were collected by means of camera traps and boat-based surveys during the molting season in two of the species' main breeding areas, over a period of five years in the Pihlajavesi basin and eight years in the Haukivesi basin. An open model approach provided minimum population estimates for these two basins. The results indicated high survival rates and site fidelity among the adult seals. More accurate estimates can be obtained in the future by increasing the surveying effort both spatially and temporally. The method presented here proved effective for evaluating population size objectively, whereas the results of the current snow lair censuses are dependent on varying winter conditions, for instance. We therefore suggest that a photo-ID-based non-invasive mark-recapture method should be used for estimating Saimaa ringed seal abundances in order to ensure reliable, transparent population monitoring under changing climatic conditions.

Winter behavior of Saimaa ringed seals: Non-overlapping core areas as indicators of avoidance in breeding females.

Climate change, together with increasing human activity, poses a threat to the breeding success of endangered landlocked ringed seals (Phoca hispida saimensis). In this study, we estimated the spatial ecology of Saimaa ringed seals during the breeding season in the ice-covered period of December-April. The telemetry data on tagged seals (n = 20), with a total of 25 separate tracking periods and birth lair locations (n = 59) of non-tagged seals, were studied to estimate the movement ecology and breeding density. The movements of the ringed seals were more restricted during the ice-covered season; the total home range size (average 7.4 km2) in winter was 13 times smaller than that in summer. Individual tagged seals occupied an average of 5 ± 3 SD subnivean haul outs (snow lairs or ice cavities), and the mean distance between the haul outs was 1.6 ± 1.1 SD km (range 0.2-5.9 km). Moreover, our data indicated that ringed seal females likely exhibited breeding time avoidance of each other's core areas, which may indicate some degree of territoriality. This was supported by the findings that the core areas (mean 1.2 km2) of tagged adult females (n = 9), did not overlap with each other. Also data on non-tagged seals showed that females did not give birth to pups within the core area radius of other parturient females. This study, together with earlier findings on the home ranges of nursed pups and perinatal mortality rates, has implications into land usage planning in Lake Saimaa by highlighting the need of undisturbed area between seal lairs and anthropogenic disturbances.

Identifying foraging habitats of Baltic ringed seals using movement data.

BACKGROUND: Identification of key foraging habitats of aquatic top predators is essential for designing effective management and conservation strategies. The Baltic ringed seal (Phoca hispida botnica) interacts with anthropogenic activities and knowledge of its spatial ecology is needed for planning population management and mitigating interactions with coastal fisheries. We investigated habitat use and foraging habitats of ringed seals (n = 26) with satellite telemetry in the northern Baltic Sea during autumn, which is important time for foraging for ringed seals. We used first passage time (FPT) approach to identify the areas of high residency corresponding to foraging areas. RESULTS: Tracked seals showed considerable movement; mean (±SD) home ranges (95 % adaptive local nearest-neighbour convex hull, a-LoCoH) were 8030 ± 4796 km(2). Two seals moved randomly and foraging areas could not be identified for them. The majority (24/26) of the studied seals occupied 1-6 main foraging areas, where they spent 47 ± 22 % of their total time. Typically the foraging areas of individuals had a mean distance of 254 ± 194 km. Most of the seals (n = 17) were "long-range foragers" which occupied several spatially remote foraging areas (mean distance 328 ± 180 km) or, in the case of two individuals, did not concentrate foraging to any particular area. The other seals (n = 9) were "local foragers" having only one foraging area or the mean distance between several areas was shorter (67 ± 26 km). Foraging areas of all seals were characterised by shallow bathymetry (median ± SD: 13 ± 49 m) and proximity to the mainland (10 ± 14 km), partly overlapping with protected areas and coastal fisheries. CONCLUSIONS: Our results indicate that in general the ringed seals range over large areas and concentrate feeding to different-often remote-areas during the open water season. Therefore, removal of individuals near the fishing gear may not be a locally effective method to mitigate seal depredation. Overlap of foraging areas with protected areas indicate that management of key foraging and resting habitats could to some extent be implemented within the existing network of marine protected areas.

A novel tool to mitigate by-catch mortality of baltic seals in coastal fyke net fishery.

Developing methods to reduce the incidental catch of non-target species is important, as by-catch mortality poses threats especially to large aquatic predators. We examined the effectiveness of a novel device, a "seal sock", in mitigating the by-catch mortality of seals in coastal fyke net fisheries in the Baltic Sea. The seal sock developed and tested in this study was a cylindrical net attached to the fyke net, allowing the seals access to the surface to breathe while trapped inside fishing gear. The number of dead and live seals caught in fyke nets without a seal sock (years 2008-2010) and with a sock (years 2011-2013) was recorded. The seals caught in fyke nets were mainly juveniles. Of ringed seals (Phoca hispida botnica) both sexes were equally represented, while of grey seals (Halichoerus grypus) the ratio was biased (71%) towards males. All the by-caught seals were dead in the fyke nets without a seal sock, whereas 70% of ringed seals and 11% of grey seals survived when the seal sock was used. The seal sock proved to be effective in reducing the by-catch mortality of ringed seals, but did not perform as well with grey seals.

Demographic histories and genetic diversities of Fennoscandian marine and landlocked ringed seal subspecies.

Island populations are on average smaller, genetically less diverse, and at a higher risk to go extinct than mainland populations. Low genetic diversity may elevate extinction probability, but the genetic component of the risk can be affected by the mode of diversity loss, which, in turn, is connected to the demographic history of the population. Here, we examined the history of genetic erosion in three Fennoscandian ringed seal subspecies, of which one inhabits the Baltic Sea 'mainland' and two the 'aquatic islands' composed of Lake Saimaa in Finland and Lake Ladoga in Russia. Both lakes were colonized by marine seals after their formation c. 9500 years ago, but Lake Ladoga is larger and more contiguous than Lake Saimaa. All three populations suffered dramatic declines during the 20th century, but the bottleneck was particularly severe in Lake Saimaa. Data from 17 microsatellite loci and mitochondrial control-region sequences show that Saimaa ringed seals have lost most of the genetic diversity present in their Baltic ancestors, while the Ladoga population has experienced only minor reductions. Using Approximate Bayesian computing analyses, we show that the genetic uniformity of the Saimaa subspecies derives from an extended founder event and subsequent slow erosion, rather than from the recent bottleneck. This suggests that the population has persisted for nearly 10,000 years despite having low genetic variation. The relatively high diversity of the Ladoga population appears to result from a high number of initial colonizers and a high post-colonization population size, but possibly also by a shorter isolation period and/or occasional gene flow from the Baltic Sea.