High Toxocara cati prevalence in wild, free-ranging Eurasian lynx (Lynx lynx) in Finland, 1999-2015.

In Finland, free-ranging Eurasian lynx (Lynx lynx) population has grown from 30 to 40 individuals to 2800 individuals since the species became partly protected in 1962. Changes in host population size are known to have an impact on host-parasite dynamics, and the Eurasian lynx population in Finland provides a unique opportunity for studying the potential effects of dramatic population increase and expansion of a solitary apex predator on their parasite prevalence and abundance. Toxocara cati is a zoonotic gastrointestinal parasite infecting domestic cats and wild felids worldwide. We studied T. cati infection prevalence and worm burden in 2756 Eurasian lynx individuals from Finland, covering the years 1999-2015. Toxocara cati worms that had been collected from intestinal contents were identified based on morphology. We performed regression analyses to investigate possible associations of age, sex, and host population density with T. cati infection. We found T. cati from 2324 (84.3%, 95% confidence interval 82.9-86.0) of the examined lynx. Each year, the infection prevalence was higher than 75% and not density dependent. The parasites were strongly aggregated, with older individuals harboring fewer T. cati than younger ones did. Old females aged 9-15 years had higher T. cati abundance than males of the same age group. Our results indicate that T. cati was a common and abundant parasite of Eurasian lynx throughout the study period, regardless of the changing population size and density.

Diversity of the MHC class II DRB gene in the wolverine (Carnivora: Mustelidae: Gulo gulo) in Finland.

The wolverine (Gulo gulo) in Finland has undergone significant population declines in the past. Since major histocompatibility complex (MHC) genes encode proteins involved in pathogen recognition, the diversity of these genes provides insights into the immunological fitness of regional populations. We sequenced 862 amplicons (242 bp) of MHC class II DRB exon 2 from 32 Finnish wolverines and identified 11 functional alleles and three pseudogenes. A molecular phylogenetic analysis indicated trans-species polymorphism, and PAML and MEME analyses indicated positive selection, suggesting that the Finnish wolverine DRB genes have evolved under balancing and positive selection. In contrast to DRB gene analyses in other species, allele frequencies in the Finnish wolverines clearly indicated the existence of two regional subpopulations, congruent with previous studies based on neutral genetic markers. In the Finnish wolverine, rapid population declines in the past have promoted genetic drift, resulting in a lower genetic diversity of DRB loci, including fewer alleles and positively selected sites, than other mustelid species analyzed previously. Our data suggest that the MHC region in the Finnish wolverine population was likely affected by a recent bottleneck.

Unique hip and stifle extensor muscle patterns in the Eurasian lynx, Lynx lynx (Carnivora: Felidae).

The Eurasian lynx (Lynx lynx) is a medium-sized felid, with a tendency to hunt for prey larger than itself. We studied the lynx hindlimb musculoskeletal anatomy in order to determine possible anatomical adaptations to hunting large prey. In our previous work, we had found characters of both large and small felids in the lynx forelimb. The crouched limbs, typical of all felids, increase the energy demands for the antigravity muscles during locomotion. As a powerful pounce is required for the smaller felid to bring down large prey, strong hindquarters may be needed. We hypothesized that the muscle attachments are more mechanically advantageous and muscles heavier in the lynx as compared to other felids to compensate for the energy requirements. In support of this, we found unique patterns in the hindlimb musculature of the lynx. Insertion of the m. gluteus medius was large with a short moment arm around the hip joint, providing mechanical disadvantage, but rapid movement. The musculus vastus medialis was relatively heavier than in other felids emphasizing the role of the m. quadriceps femoris as a powerful stifle extensor. The extensor muscles support the crouched hind limbs, which is crucial when tackling large prey, and they are also responsible for the swift powerful pounce brought by extending the hindlimbs. However, we cannot rule out the possibility the characters are shared with other Lynx spp. or they are adaptations to other aspects of the locomotor strategy in the Eurasian lynx.

Genetic analysis indicates spatial-dependent patterns of sex-biased dispersal in Eurasian lynx in Finland.

Conservation and management of large carnivores requires knowledge of female and male dispersal. Such information is crucial to evaluate the population's status and thus management actions. This knowledge is challenging to obtain, often incomplete and contradictory at times. The size of the target population and the methods applied can bias the results. Also, population history and biological or environmental influences can affect dispersal on different scales within a study area. We have genotyped Eurasian lynx (180 males and 102 females, collected 2003-2017) continuously distributed in southern Finland (~23,000 km2) using 21 short tandem repeats (STR) loci and compared statistical genetic tests to infer local and sex-specific dispersal patterns within and across genetic clusters as well as geographic regions. We tested for sex-specific substructure with individual-based Bayesian assignment tests and spatial autocorrelation analyses. Differences between the sexes in genetic differentiation, relatedness, inbreeding, and diversity were analysed using population-based AMOVA, F-statistics, and assignment indices. Our results showed two different genetic clusters that were spatially structured for females but admixed for males. Similarly, spatial autocorrelation and relatedness was significantly higher in females than males. However, we found weaker sex-specific patterns for the Eurasian lynx when the data were separated in three geographical regions than when divided in the two genetic clusters. Overall, our results suggest male-biased dispersal and female philopatry for the Eurasian lynx in Southern Finland. The female genetic structuring increased from west to east within our study area. In addition, detection of male-biased dispersal was dependent on analytical methods utilized, on whether subtle underlying genetic structuring was considered or not, and the choice of population delineation. Conclusively, we suggest using multiple genetic approaches to study sex-biased dispersal in a continuously distributed species in which population delineation is difficult.

Genetic evidence of female kin clusters in a continuous population of a solitary carnivore, the Eurasian lynx.

Large terrestrial carnivores can sometimes display strong family bonds affecting the spatial distribution of related individuals. We studied the spatial genetic relatedness and family structure of female Eurasian lynx, continuously distributed in southern Finland. We hypothesized that closely related females form matrilineal assemblages, clustering together with relatives living in the neighboring areas. We evaluated this hypothesis using tissue samples of 133 legally harvested female lynx (from year 2007 to 2015), genotyped with 23 microsatellite markers, and tested for possible spatial genetic family structure using a combination of Bayesian clustering, spatial autocorrelation, and forensic genetic parentage analysis. The study population had three potential family genetic clusters, with a high degree of admixture and geographic overlap, and showed a weak but significant negative relationship between pairwise genetic and geographic distance. Moreover, parentage analysis indicated that 64% of the females had one or more close relatives (sister, mother, or daughter) within the study population. Individuals identified as close kin consistently assigned to the same putative family genetic cluster. They also were sampled closer geographically than females on average, although variation was large. Our results support the possibility that Eurasian lynx forms matrilineal assemblages, and comparisons with males are now required to further assess this hypothesis.

Musculoskeletal anatomy of the Eurasian lynx, Lynx lynx (Carnivora: Felidae) forelimb: Adaptations to capture large prey?

Mammalian carnivores adhere to two different feeding strategies relative to their body masses. Large carnivores prey on animals that are the same size or larger than themselves, whereas small carnivores prey on smaller vertebrates and invertebrates. The Eurasian lynx (Lynx lynx) falls in between these two categories. Lynx descend from larger forms that were probably large prey specialists, but during the Pleistocene became predators of small prey. The modern Eurasian lynx may be an evolutionary reversal toward specializing in large prey again. We hypothesized that the musculoskeletal anatomy of lynx should show traits for catching large prey. To test our hypothesis, we dissected the forelimb muscles of six Eurasian lynx individuals and compared our findings to results published for other felids. We measured the bones and compared their dimensions to the published material. Our material displayed a well-developed pectoral girdle musculature with some uniquely extensive muscle attachments. The upper arm musculature resembled that of the pantherine felids and probably the extinct sabertooths, and also the muscles responsible for supination and pronation were similar to those in large cats. The muscles controlling the pollex were well-developed. However, skeletal indices were similar to those of small prey predators. Our findings show that lynx possess the topographic pattern of muscle origin and insertion like in large felids. J. Morphol. 277:753-765, 2016. © 2016 Wiley Periodicals, Inc.

Recovery of large carnivores in Europe's modern human-dominated landscapes.

The conservation of large carnivores is a formidable challenge for biodiversity conservation. Using a data set on the past and current status of brown bears (Ursus arctos), Eurasian lynx (Lynx lynx), gray wolves (Canis lupus), and wolverines (Gulo gulo) in European countries, we show that roughly one-third of mainland Europe hosts at least one large carnivore species, with stable or increasing abundance in most cases in 21st-century records. The reasons for this overall conservation success include protective legislation, supportive public opinion, and a variety of practices making coexistence between large carnivores and people possible. The European situation reveals that large carnivores and people can share the same landscape.

Endoparasites of the Eurasian lynx (Lynx lynx) in Finland.

We sampled 339 fecal samples, 296 intestines, and 82 lungs from 371 lynx hunted during the 2010-2011 season in Finland. The fecal samples were analyzed for endoparasites by a quantitative flotation method, and helminths from intestines were studied morphologically, while lungs were investigated for pulmonary parasites. From fecal samples, eggs and oocysts of at least 6 different endoparasite species were identified, with a mean of 1.5 (range 0-4) parasite species per host. In the intestines, at least 4 different helminth species were found, with the mean of 2.0 (range 1-4) species per infected host. The prevalence of eggs in feces and the prevalence of worms in intestines were 71% and 93% for Toxocara cati , 29% and 68% for Taenia spp., and 5% and 2% for Diphyllobothrium sp., respectively. Only eggs were detected for Capillaria sp. (46%) and Uncinaria sp. (0.6%) nematodes, and only adults were detected for Mesocestoides sp. cestodes (0.3%). Significant positive correlations were evident between the number of T. cati (r = 0.664; P = 0.01) and Diphyllobothrium sp. (r = 0.645; P = 0.01) eggs per gram of feces and adult worms detected in intestine. In addition to the metazoan parasites, protozoan Isospora sp. oocysts were also found (0.6%). Pulmonary samples were all negative for parasites. These data demonstrate that lynx commonly harbor various endoparasites, some of which are zoonotic.

Free-ranging Eurasian lynx (Lynx lynx) as host of Toxoplasma gondii in Finland.

We investigated the presence of Toxoplasma gondii infections in Eurasian lynx (Lynx lynx) in Finland by analyzing samples from 337 lynx that were legally hunted during the 2010-2011 season and by performing a retrospective nationwide database search of postmortem toxoplasmosis diagnoses in this species. We detected specific anti-T. gondii IgG antibodies in 290 (86.1%) of the 337 lynx. The method used was a direct agglutination test, and samples positive at the used dilution 1:40 were defined as antibody positive. Older lynx had 14.3 times higher odds of being antibody-positive than did lynx of the presumed age of 7-10 mo, and lynx weighing >15 kg had 16.7 times higher odds of being antibody positive than did those ≤ 15 kg. Lynx from the southwest were more often antibody positive, with an odds ratio 6.3, than lynx from the northeast. None of the 332 fecal samples available was positive for the presence of T. gondii-like oocysts with a quantitative MgSO4 flotation technique, and none of the 167 free-ranging Eurasian lynx examined postmortem by veterinary pathologists from January 2000 to May 2010 had died from toxoplasmosis. Although Finnish lynx were confirmed to commonly encounter T. gondii, we found no evidence of an ongoing contribution to the environmental oocyst burden nor of the lynx dying from the infection.

Rabies in northeastern Europe--the threat from invasive raccoon dogs.

The red fox (Vulpes vulpes) is the main terrestrial wildlife rabies vector in Europe. However, recently the raccoon dog (Nyctereutes procyonoides)--an invasive species originating from East Asia--has become increasingly important as secondary host, especially in the Baltic States. This imposes problems on neighboring rabies-free countries (such as Finland), where the density of each of the two vector species on its own might be too low to sustain a long-term rabies epizootic, but the community of vectors could be large enough to support a rabies epizootic. In this modeling study, we analyzed rabies epizootics in a community of foxes and raccoon dogs. We focused on the impact of density and behavioral differences (hibernation) between the two vector species. We found that rabies could persist in the community, even if the disease would not spread in the single vector species because its density was too low. Epizootics in the community were stronger than expected for single species, and raccoon dogs were usually the major rabies host. If raccoon dog territory density was high, invasive raccoon dogs could even outcompete native foxes because of apparent competition via the rabies virus. The enhancement in disease risk and disease intensity caused by raccoon dogs suggests that current strategies to control wildlife rabies in Europe should be reviewed, and that oral rabies vaccination also should target raccoon dogs after they emerge from hibernation.