Carl-Gustaf Thulin

The occurrence of mountain hare mitochondrial DNA in wild brown hares.

If interspecific hybrids are fertile and backcross to either parental species, transmission of mitochondrial DNA over the species barrier can occur. To investigate if such transmission has occurred between the brown hare Lepus europeus Pall and the mountain hare L. timidus L. in Scandinavia, an analysis of genetic variation in mitochondrial DNA from 36 hares, collected from 15 localities, was performed. Sequence divergence of mtDNA between species was estimated at 8 ± 1% (SD). Intraspecific mtDNA sequence divergence varied between 0.09 and 0.38% in brown hares and 0.10 and 1.44% in mountain hares. In six out of 18 brown hares examined, two different haplotypes of mountain hare origin were detected, demonstrating a transmission of mtDNA haplotypes from mountain hares to brown hares. The results indicate that interspecific hybridization between the two species occurs in wild populations.

Biased geographical distribution of mitochondrial DNA that passed the species barrier from mountain hares to brown hares (genus Lepus ): an effect of genetic incompatibility and mating behaviour?

Through interspecific hybridization and subsequent backcrossing, genes and genomes may be transferred over the species barrier. In Sweden, the introduced brown hare Lepus europaeus hybridizes with the native mountain hare L. timidus. To investigate the direction and the extent of transfer of mitochondrial DNA (mtDNA) between the species, the mtDNA haplotypes were screened in 522 brown hares and 149 mountain hares from areas of sympatry and allopatry. A total of 51 brown hares with mountain hare mtDNA, but no mountain hares with brown hare mtDNA were detected. Thus, mtDNA transfer over the species barrier is directed from mountain hares to brown hares. We argue that frequency-dependent hybridization and/or interspecific male competition mediates this directionality. Further, the percentage of brown hares with transmitted mountain hare mtDNA was lower in areas of former species sympatry (0.6%) compared to areas of current sympatry (15%). Thus, the transferred mtDNA may disappear from brown hare populations if there is no continuous input through hybridization. We suggest that specimens with an alien mtDNA experience a fitness reduction as a result of a functional incompatibility between the cytoplasmic mitochondrial genomes and the cell nucleus.

Genetic divergence in the small Indian mongoose (Herpestes auropunctatus), a widely distributed invasive species

The combination of founder events, random drift and new selective forces experienced by introduced species typically lowers genetic variation and induces differentiation from the ancestral population. Here, we investigate microsatellite differentiation between introduced and native populations of the small Indian mongoose (Herpestes auropunctatus). Many expectations based on introduction history, such as loss of alleles and relationships among populations, are confirmed. Nevertheless, when applying population assignment methods to our data, we observe a few specimens that are incorrectly assigned and/or appear to have a mixed ancestry, despite estimates of substantial population differentiation. Thus, we suggest that population assignments of individuals should be viewed as tentative and that there should be agreement among different algorithms before assignments are applied in conservation or management. Further, we find no congruence between previously reported morphological differentiation and the sorting of microsatellite variation. Some introduced populations have retained much genetic variation while others have not, irrespective of morphology. Finally, we find alleles from the sympatric grey mongoose (Herpestes edwardsii) in one small Indian mongoose within the native range, suggesting an alternative explanation for morphological differentiation involving a shift in female preferences in allopatry.

Species assignment and hybrid identification among Scandinavian hares Lepus europaeus and L. timidus

Abstract In Scandinavia, suspected hybrids between the native mountain hare Lepus timidus and the introduced brown hare L. europaeus have been observed by hunters since the first introductions of brown hares in the late 19th century. Several attempts to verify the status of these suspected hybrids have been unsuccessful. Recently, however, the transmission of mitochondrial DNA (mtDNA) from mountain hares to brown hares was documented and interpreted as a consequence of hybridisation and subsequent introgression. However, mtDNA markers alone will not allow identification of first-generation hybrids because of the strictly maternal inheritance of mitochondria. Here, we combine mtDNA data with analyses of variation in seven microsatellite loci among brown hares, mountain hares and putative hybrids. Our purpose was to determine species differentiation in nuclear DNA markers, elucidate the extent of interspecific gene flow, identify true hybrids within our sample and evaluate the ability of hunters to identify hybrids. The estimated genetic difference between species was low (FST = 0.18 – 0.24, RhoST = 0.09 – 0.16). We believe these low estimates result from a reticulated mode of evolution among hares, with repeated gene flow over the species barrier. Population assignment tests and randomly assembled, artificial, hybrid genotypes were used to classify individuals independently from the morphologically assessed species identity. More than half (57%) of the putative hybrid specimens were assigned unambiguously to either species and first-generation hybrids seem to be rare. Morphological plasticity and backcrossing, which confound species identification within the genus Lepus, might explain invalid classifications by hunters.

Temporal and spatial variation in Anaplasma phagocytophilum infection in Swedish moose (Alces alces)

SUMMARY The occurrence of Anaplasma phagocytophilum was investigated in spleen and serum samples from Swedish moose (Alces alces) in southern Sweden (island and mainland). Samples were analysed for presence of A. phagocytophilum DNA by real-time PCR (n = 263), and for Anaplasma antibodies with ELISA serology (n = 234). All serum samples had antibodies against A. phagocytophilum. The mean DNA-based prevalence was 26·3%, and significant (P < 0·01) temporal, and spatial variation was found. Island moose had significantly (P < 0·001) higher prevalence of A. phagocytophilum DNA than moose from the mainland areas. Two samples were sequenced to determine genetic variation in the 16S rRNA and groESL genes. Genetic sequence similarity with the human granulocytic anaplasmosis agent, equine granulocytic ehrlichiosis agent, and different wildlife-associated A. phagocytophilum variants were observed in the 16S rRNA and groESL genes. Our study shows that moose are exposed to A. phagocytophilum in Sweden, and represent a potential wildlife reservoir of the pathogen.

Detecting hybridization between willow grouse (Lagopus lagopus) and rock ptarmigan (L. muta) in Central Sweden through Bayesian admixture analyses and mtDNA screening

Willow grouse (L. lagopus) and rock ptarmigan (L. muta) are sibling species with similar phenotypic and life histories that coexist sympatrically in wide areas of their distribution range. These grouse are amongst the most popular game birds in Scandinavia but contrary to other quarry species, no restocking with captive-bred animals has ever been performed. The discovery of two individuals with intermediate plumage features evoked the question of possible hybridization events between both species, an idea that did not seem too unlikely on the basis of habitat overlap. Thus, to assess whether any genetic exchange is occurring, we used different Bayesian-based admixture analyses of multilocus genotypes determined at twelve microsatellite loci. We also obtained mitochondrial COI-sequences from a selected number of individuals to infer the maternal geneflow and potential introgression. The capacity of our panel of microsatellite markers to detect hybridization was verified using assignments of simulated genotypes. We then evaluated the extent of hybridization in an actual sample of 111 individuals collected in a 100-km2 area in the Scandinavian mountain range. An admixed condition was verified in one of the suspected hybrids, that seemed to carry a L. muta genotype with partial L. lagopus introgression. In addition, more than 4% of L. lagopus showed signs of hybridization under the most conservative scenario with respect to discrepancies between population assignment methods. This was unexpected, given that no L. lagopus displayed any apparent intermediate plumage features. Furthermore, interspecific geneflow of mtDNA haplotypes was lower than expected; which suggests that Haldane’s rule might apply for these two grouse species. Hence, plumage identification of hybrid ancestry is not always reliable and might lead to biases in the estimation of hybridization rates. Hybridisation may be expected to increase if the climate gets warmer as the habitat overlap between the species will become more extensive. We discuss whether hybridisation is a threat to the long-term survival of any of the two species.

High mtDNA haplotype diversity among introduced Swedish brown haresLepus europaeus

The brown hareLepus europaeus Pallas, 1778 occurs naturally in central Eurasia, but has been introduced to parts of northern Europe, South- and North America, Australia and New Zealand. Brown hares were introduced to Sweden from central Europe for hunting purposes during the 19th century. We investigated how the human--mediated brown hare colonisation of Sweden is reflected in the amount of genetic variation present by assessing variation and composition of mitochondrial DNA (mtDNA) lineages among Swedish brown hares. MtDNA from a total of 40 brown hare specimens from 15 localities were analysed for Restriction Fragment Length Polymorphisms. The haplotype diversity is surprisingly high (0.893 ± 0.002) when compared to the mtDNA diversity among brown hares on the European continent as well as to other mammalian species. Admixture of haplotypes from different source populations combined with a reduced effect of random genetic drift and a relaxed selection pressure due to rapid population growth after introduction are mechanisms that are likely to account for the observed high mtDNA haplotype diversity.

Variable flight initiation distance in incubating Eurasian curlew

Flight initiation distances (FIDs) of nesting birds approached by a predator likely reflect evolutionary stable strategies in which birds make trade-offs between adult survival and reproductive success. Here, we test if FID (a) had an impact on hatching success, (b) was adjusted to current conditions, and (c) was consistent for individual nests. All experiments were performed with a human approaching incubating Eurasian curlews Numenius arquata, a ground-nesting wader species under high egg predation pressure. Our results show that hatching success was higher in nests where the incubating parent left at intermediate FIDs compared to short and long ones, and that FID decreased with date and time of the evening. Further, FIDs from repeated approaches were not consistent within nests. We suggest that incubating Eurasian curlews follow a “surprise” strategy, where an element of randomness is superimposed on a context-adjusted norm to prevent predators from predicting their FID behaviour.

Microsatellite investigation of roe deer (Capreolus capreolus) in Scandinavia reveals genetic differentiation of a Baltic Sea Island population

After a decline in the early 19th century, a remnant population of the Scandinavian roe deer (Capreolus capreolus) was protected in southern Sweden around 1840. The roe deer quickly recovered and recolonized most of the Scandinavian Peninsula. In this study, I analyze microsatellite variation in 14 roe deer populations in Scandinavia to (1) investigate if all Swedish roe deer trace their ancestry to southern Sweden, (2) define other areas where roe deer may have remained during the population decline, and (3) examine if the large geographic distances during the expansion phase have influenced the degree and sorting of genetic variation. The results show that the mainland roe deer most likely trace their ancestry to southern Sweden, but interestingly, the two neighboring populations Övedskloster and Wittskövle in the south are significantly different from each other (p<0.01). Furthermore, the population in Wittskövle and roe deer from Köpingsvik at the Baltic Sea Island Öland are significantly different from all other populations (p<0.1). Thus, potentially, there are more than one founder population that contributed to the current Swedish population. The particular differentiation of roe deer from Öland might be an effect of random population genetic processes during the population expansion phase, potentially in combination with introductions of roe deer from other areas than the Swedish mainland. The results are discussed in relation to management of roe deer in Sweden.

Opportunities and challenges with growing wildlife populations and zoonotic diseases in Sweden

In many parts of Europe and North America, populations of large mammals and birds have recovered during recent decades. In Sweden, this has resulted in more wildlife than was historically recorded. This positive development provides a number of opportunities for ecosystem services and for biodiversity. More wildlife also means more interactions with humans, as many birds and mammals may interfere with human interests in the landscape, such as natural resource use. Thus, more wildlife may shift the baseline for societal perception of wildlife. Wildlife species may host pathogens with potential for infecting humans and domestic animals. With increasing knowledge about zoonotic diseases and their dynamics, more scientific, media, and public attention is given to zoonotic processes. We are concerned with how the public image of the wild animals is affected, because many of the recent zoonotic outbreaks connect animal groups to diseases such as avian and swine influenza, lyme disease, and tick-borne encephalitis. The societal focus on zoonotic diseases may increase the fear of the wild and will separate the public further from the outdoors in general and wildlife in particular. Ultimately, we risk a juxtaposition of the overall acceptance of biological diversity and a shifting societal perception of wildlife that could be harmful for life on earth. We therefore suggest multidisciplinary research on societal awareness of zoonotic diseases and its implications for public acceptance for wildlife and biological diversity.