Population Structure as Revealed by mtDNA and Microsatellites in Northern Fur Seals, Callorhinus ursinus

W
Document Sample
scope of work template
							                                                    Population Structure as Revealed by mtDNA and
                                                Microsatellites in Northern Fur Seals, Callorhinus ursinus
                                                                                                          B. R. Dickerson1, R. R. Ream1, S. N. Vignieri2,
                                                                                                                 P. Bentzen3 and G. A. Antonelis4
                                                               1Molecular   Ecology Research Laboratory, National Marine Mammal Laboratory, Alaska Fisheries Science Center, National Marine Fisheries Service, Seattle, WA 98115, USA
                                                                            2Departmentof Organismic and Evolutionary Biology, Harvard University, Museum of Comparative Zoology, 26 Oxford St., Cambridge, MA 02138, USA
                                                                                                                       3Dalhousie University, Halifax, Nova Scotia, B3H 4J1, Canada
                                                                                        4Pacific Islands Fisheries Science Center, National Marine Fisheries Service, NOAA, 2570 Dole St., Honolulu, Hawaii, USA




                               Introduction
The northern fur seal, Callorhinus ursinus, is a widely                                                                                                                                                                                                                                                                                                   132


distributed philopatric species that breeds on a limited
                                                                                                                                                                                                                                                                                                                                                    165


                                                                                                                                                                              Results
                                                                                                                                                                                                                                                                                                                                                                     70




number of islands across the North Pacific Ocean (Fig. 1).                                                                                                                                                                                                                                                                                                  1



The species has undergone large fluctuations in abundance                                                    Microsatellites
during the past two centuries. To date there have been no                                                    - Maximum likelihood tests looking at number of potential populations using
molecular studies of genetic variation or population structure.                                              an admixture model showed the islands best grouped into one population.                                                                                         71
                                                                                                                                                                                                                                                                    11


The goal of this project was to examine population structure                                                 - FST and RST were extremely low and insignificant following Bonferroni
in northern fur seals, throughout their entire range, using                                                  corrections:                                                                                                                                           41
                                                                                                                                                                                                                                                                             15
                                                                                                                                                                                                                                                                                   83

                                                                                                                                                                                                                                                                                                                                               13



both mtDNA and microsatellites.
                                                                                                                                                                                                                                                                                                                                                                33


                                                                                                                 - All populations FST = 0.0004, RST = -0.0041                                                                                                                                    87               74                  66




                                                                                                                 - Population pairs FST ≤ 0.0043 , RST ≤ 0.0373                                                                                                                              134

                                                                                                                                                                                                                                                                                                                                                                      31



                                                                                                             -Tests for isolation by distance were also insignificant.
                                                                                                                                                                                                                                                                                             10


                                                                                                                                                                                                                                                                                  172
                                                                                                                                                                                                                                                                                        75

                                                                                                                                                                                                                                                                                                                             1 Substitution




                                                                                                             mtDNA
                                                                                                             -Tests for differentiation between the 6 regional population groups (Robben I.,                                                                                                           136



                                                                                                             Kuril Islands (Lovushki and Srednev), Commander Islands (Bering and Medny), Bogoslof I., Pribilof Islands (St
                                                                                                             Paul and St George) and San Miguel I.) were not significant (AMOVA, p = 0.87).

                                                                                                             - Pairwise comparisons of conventional FST values revealed little
                                                                                                             differentiation but ΦST values were higher and showed some difference
                                                                                                             between the Russian animals and those in the eastern range (Table 1).                                                                                Figure 3. Minimum spanning network of 112 core mitochondrial DNA haplotypes of northern fur seals
                                                                                                                                                                                                                                                                  samples for this study. Branch lengths are the minimum number of steps between haplotypes. The size
                                                                                                                                                                                                                                                                  of the circle representing the individual haplotypes corresponds to the abundance of that haplotype.
                                                                                                             - Although not significant there does appear to be a trend toward isolation                                                                          Numbers identify the most abundant haplotypes. Dashed lines represent alternative groupings.

                                                                                                             by distance (p = 0.27; Figure 2).
                                                                                                             - Phylogeny analysis results (neighbor joining tree) were inconclusive with                                                                          0.25
                                                                                                             few nodes (7%) supported by >60% in the bootstrap analysis and no
                                                                                                             geographic concordance.                                                                                                                               0.2
                                                                                                             - The minimum spanning network showed 4 distinct maternal lineages,
                                                                                                             although there was no geographic concordance (Figure 3).




                                                                                                                                                                                                                                                      Frequency
                                                                                                                                                                                                                                                                  0.15
                                                                                                                                                                                                                                                                                                                                               observed
 Figure 1. Distribution of northern fur seal breeding sites.                                                 - The nucleotide frequency mismatch distribution matched that of a model                                                                                                                                                          expected
                                                                                                             of sudden expansion (Figure 4).                                                                                                                       0.1


                                                                                                                                                                                                                                                                  0.05
                                                                                                             Table 1. mtDNA based population differentiation and structure for population pairs (estimates of FST above diagonal
                                 Methods                                                                     and ΦST below diagonal). Bold indicates significant P-values (P < 0.05).

                                                                                                                                                                                                                                                                         0
                                                                                                                             Robben    Lovushki    Srednev     Bering       Medny       Bogoslof    St Paul   St George    San Miguel                                         0          5             10               15                    20           25
Flipper punches were taken from 728 individuals during the 1993-1998                                                                                                                                                                                                                                         Pairwise Differences
                                                                                                                   Robben                 -0.001      -0.001   -0.002         -0.001       -0.003     0.000       -0.003         0.000
and 2005 summer breeding seasons from nine islands: Bering Island (N
                                                                                                                Lovushki       0.013                   0.000        0.001     0.001         0.001     0.002       0.002          0.001
= 55), Bogoslof Island (N = 99), Lovushki Island (N = 61), Srednev Rock
(N = 50), Medny Island (N = 56), Robben Island (N = 50), St. George                                                Srednev     0.018      -0.003                    0.002     0.002         0.002     0.004       0.000          0.002                            Figure 4. Observed pairwise mismatch distribution of mtDNA in northern
                                                                                                                                                                                                                                                                  fur seals compared with the distribution of a model of sudden population
Island (N = 100), St. Paul Island (N = 113) and San Miguel Island (N =                                              Bering     0.017       0.010        0.02                  0.000        -0.002     0.000       -0.001         0.002
                                                                                                                                                                                                                                                                  expansion.
144; Fig. 1).                                                                                                       Medny      0.010      -0.007       0.006        0.012                  -0.001     0.002       0.000          0.002

                                                                                                                Bogoslof       0.010      -0.001       0.015         0.02     -0.007                  0.002       -0.001         0.002
                                                                                                                                                                                                                                                                                                  Discussion
To test for genetic variability and population differentiation between                                             St Paul     0.019      -0.002      -0.002        0.017     -0.001        0.007                 0.003          0.005
islands, seven microsatellite loci and 381 bp from the mtDNA control                                         St George         0.012      -0.003       0.006        0.025     -0.008       -0.002     0.000                      0.000
region were used. Allelic variability for the microsatellite loci were high                                   San Miguel       0.024      -0.000       0.011        0.024     -0.004        0.009     0.003       -0.001                 Our microsatellite analysis did not show any structure across the range but based on
with heterozygosities ranging from 0.25 to 0.93. mtDNA haplotype                                                                                                                                                                         pairwise comparisons of ΦST values there does appear to be some separation
variability was also high with overall haplotypic diversity at 0.994 and                                                                                                                                                                 between the Russian and eastern populations based on the mtDNA analysis. Since
nucleotide diversity moderate at 2.4%. From the 619 individuals we were                                                                                                                                                                  mtDNA is maternally inherited movement between populations by males does not
able to sequence we found 87 variable sites resulting in 332 haplotypes                                                                                                                                                                  influence an examination of population structure. So, under conditions of high male
of which 227 were represented by a single individual.                                                                                                                                                                                    migration and low female migration you would expect to see higher levels of structure
                                                                                                              0.03                                                                                                                       when looking at mtDNA compared to microsatellites which are inherited in the
Estimates of FST , ΦST (mtDNA), and RST (microsatellites), were                                                                                                                                                                          Mendialian fashion. These results suggest that as in other pinniped species males
calculated and pairwise comparisons and AMOVA were used to asses                                            0.025                                                                                                                        are probably more apt to immigrate than females. Although, given the low levels of
population structure. The maximum likelihood program Structure was                                            0.02                                                                                                                       structure that were found and the observed levels of philopatry in northern fur seals,
used to estimate the most likely number of populations the islands                                                                                                                                                                       these results cannot be explained by male migration alone. The shapes of the
                                                                                                            0.015                                                                                                                        minimum spanning network (Figure 3) and the pairwise mismatch distribution (Figure
represented based on the microsatellite results. Isolation by distance
                                                                                            ΦST \1- ΦST




was examined for both markers and a neighbor joining tree, nucleotide                                         0.01                                                                                                                       4) suggest that suggest that the population has recently (evolutionarily) undergone
mismatch distribution and minimum spanning network were created for                                                                                                                                                                      rapid population expansion, possibly as a result of populations expansion and the
                                                                                                            0.005
the mtDNA data.                                                                                                                                                                                                                          subsequent recolonization of modern day breeding sites following the retreat of the
                                                                                                                    0                                                                                                                    Pleistocene ice sheets and in a more recent time frame, expansion since the
                                                                                                                                           2                   4                    6                    8                 10            cessation of hunting pressures.
                                                                                                          -0.005

                                                                                                            -0.01                                                                                                                        The results do not prove that population structure is absent in the northern fur seal.
                                                                                                                                                                                                                                         The extensive geographic separation of breeding islands and the foraging habitat
                                                                                                          -0.015
                                                                                                                                                                                                                                         used around these islands, differences in population dynamics, high levels of
                                                                                                                                                                   Ln Distance (km)                                                      philopatry, the slight heterogeneity observed in microsatellite allele frequencies over
                                                                                                                                                                                                                                         all populations, and low levels of differentiation in mtDNA contradict this notion. When
                                                                                                                        Figure 2. Isolation by distance based on mitochondrial DNA analysis in northern
                                                                                                                        fur seals. Relationship between genetic distance, pairwise comparisons of
                                                                                                                                                                                                                                         examining population structure it is important to keep in mind that changes in F-
                                                                                                                        rookeries (ΦST) and the natural log of the geographic distance between the rookery                               statistics increase very slowly after reductions in migration occur and even the use of
                                                                                                                        pairs.                                                                                                           markers with high mutation rates still result in a long lag time between demographic
                                                                                                                                                                                                                                         and genetic differentiation. It is important, particularly when using neutral markers, to
                                                                                                                                                                                                                                         look at more than just genetic differentiation to determine structure. Overall, further
                                                                                                                                                                                                                                         investigation into the population structure present in this species may allow for the
                                                                              The recommendations and general content presented in this poster do not necessarily represent the views or                                                 identification of the population processes which are actually occurring, be they due to
                                                                              official position of the Department of Commerce, the National Oceanic and Atmospheric Administration, or the                                               migration (both resent and historic), mating strategy, high population numbers, or a
                                                                              National Marine Fisheries Service.                                                                                                                         complicated small-scale polymorphic pattern.