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Abstract

Grey wolves (Canis lupus) are one of the few large terrestrial carnivores that have maintained a wide geographic distribution across the Northern Hemisphere throughout the Pleistocene and Holocene. Recent genetic studies have suggested that, despite this continuous presence, major demographic changes occurred in wolf populations between the late Pleistocene and early Holocene, and that extant wolves trace their ancestry to a single late Pleistocene population. Both the geographic origin of this ancestral population and how it became widespread remain unknown. Here, we used a spatially and temporally explicit modelling framework to analyse a dataset of 90 modern and 45 ancient mitochondrial wolf genomes from across the Northern Hemisphere, spanning the last 50,000 years. Our results suggest that contemporary wolf populations trace their ancestry to an expansion from Beringia at the end of the Last Glacial Maximum, and that this process was most likely driven by Late Pleistocene ecological fluctuations that occurred across the Northern Hemisphere. This study provides direct ancient genetic evidence that long‐range migration has played an important role in the population history of a large carnivore, and provides an insight into how wolves survived the wave of megafaunal extinctions at the end of the last glaciation. Moreover, because late Pleistocene grey wolves were the likely source from which all modern dogs trace their origins, the demographic history described in this study has fundamental implications for understanding the geographical origin of the dog.
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www.zora.uzh.ch
Year: 2019
Ancient DNA suggests modern wolves trace their origin to a Late
Pleistocene expansion from Beringia
Loog, Liisa ; Thalmann, Olaf ; Sinding, Mikkel-Holger S ; Schuenemann, Verena J ; et al
Abstract: Grey wolves (Canis lupus) are one of the few large terrestrial carnivores that have maintained a
wide geographical distribution across the Northern Hemisphere throughout the Pleistocene and Holocene.
Recent genetic studies have suggested that, despite this continuous presence, major demographic changes
occurred in wolf populations between the Late Pleistocene and early Holocene, and that extant wolves
trace their ancestry to a single Late Pleistocene population. Both the geographical origin of this ancestral
population and how it became widespread remain unknown. Here, we used a spatially and temporally
explicit modelling framework to analyse a data set of 90 modern and 45 ancient mitochondrial wolf
genomes from across the Northern Hemisphere, spanning the last 50,000 years. Our results suggest that
contemporary wolf populations trace their ancestry to an expansion from Beringia at the end of the Last
Glacial Maximum, and that this process was most likely driven by Late Pleistocene ecological uctuations
that occurred across the Northern Hemisphere. This study provides direct ancient genetic evidence that
long-range migration has played an important role in the population history of a large carnivore, and
provides insight into how wolves survived the wave of megafaunal extinctions at the end of the last
glaciation. Moreover, because Late Pleistocene grey wolves were the likely source from which all modern
dogs trace their origins, the demographic history described in this study has fundamental implications
for understanding the geographical origin of the dog.
DOI: https://doi.org/10.1111/mec.15329
Posted at the Zurich Open Repository and Archive, University of Zurich
ZORA URL: https://doi.org/10.5167/uzh-180219
Journal Article
Published Version
The following work is licensed under a Creative Commons: Attribution 4.0 International (CC BY 4.0)
License.
Originally published at:
Loog, Liisa; Thalmann, Olaf; Sinding, Mikkel-Holger S; Schuenemann, Verena J; et al (2019). Ancient
DNA suggests modern wolves trace their origin to a Late Pleistocene expansion from Beringia. Molecular
Ecology:Epub ahead of print.
DOI: https://doi.org/10.1111/mec.15329
Molecular Ecology. 2019;00:1–15.
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
FROM THE COVER
Ancient DNA suggests modern wolves trace their origin to a
Late Pleistocene expansion from Beringia
Liisa Loog1,2,3,4 | Olaf Thalmann5| Mikkel-Holger S. Sinding6, 7, 8 |
Verena J. Schuenemann9,10,11| Angela Perri12| Mietje Germonpré13|
Herve Bocherens10,14| Kelsey E. Witt15| Jose A. Samaniego Castruita6|
Marcela S. Velasco6| Inge K. C. Lundstrøm6| Nathan Wales6,16| Gontran Sonet17 |
Laurent Frantz2| Hannes Schroeder6| Jane Budd18| Elodie-Laure Jimenez13|
Sergey Fedorov19| Boris Gasparyan20| Andrew W. Kandel21|
Martina Lázničková-Galetová22,23,24| Hannes Napierala25| Hans-Peter Uerpmann9|
Pavel A. Nikolskiy26,27| Elena Y. Pavlova27, 2 8| Vladimir V. Pitulko27|
Karl-Heinz Herzig5,29| Ripan S. Malhi30| Eske Willerslev2,31,32| Anders J. Hansen8,31|
Keith Dobney33,34,35| M. Thomas P. Gilbert6,36| Johannes Krause9,37| Greger Larson1|
Anders Eriksson2,38,39 | Andrea Manica2
1 
2
 
4
5
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7  
8
9 
10
11  
12
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14
15
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17
18
19  
20
21
22



Molecular Ecology
      
2 
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   LOOG et aL.
24
25

27
28 
29

 


 
 




Correspondence














Funding information


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  


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
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 
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


   

      
    





          
          



          



 

  
 
  


 
  

Abstract
  Canis lupus        
        
             


      

 




   
              

   
    
 


KEY WORDS


1 | INTRODUCTION


       
Canis lupus
      
           
  
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LOOG et aL .
 
        
           

            


 

       
  


         


            
         
        
 
      
  
   

            
      



2 | MATERIALS AND METHODS
2.1 | Data preparation
         
     
  
        
     


       
            



            
     







         


 






        
     
        
            


 

  
    
             
              


        
            
            


       
   
    
           
clustalw




      
   

2.2 | Phylogenetic analysis
    
         
       
     

4 
|
   LOOG et aL.
     
            
      



       


         
             

          


Partition Finder
and bea st       
 
      



FIGURE 1 


Sample age [1,000 years BP]
020406080
*
*
*
*
*
*
*
(a)
(b)
CGG16
TU14
TH14
TU4
TU8
TU5
CGG20
CGG21
TU13
TU11
TH11
TH15
TH7
TH5
CGG33
CGG19
TH10
TU15
TH1
TH8
CGG17
TH3
TH6
CGG34
TU10
TU9
CGG26
CGG22
CGG25
CGG28
TU3
CGG14
CGG27
CGG15
TU2
TH12
SK1
TU1
TU6
CGG18
TU7
TH4
CGG12
CGG32
CGG29
  
|
 5
LOOG et aL .
bea st
             
           

  bea st run
             
be ast



R2
        
              
 

      
              
bea st
          N     
    
        
 
    
bea st
2.3 | Isolation by distance analysis
             
       
    

      
         
             
         

Gi and j; φi
and φji and jλi
and λji and j
and r

        dist.dna
aPe
2.4 | Geographical deme definitions
 

  European
        
             
         
 
  Middle-Eastern      


        


  Central Nor th Eurasian
              




  East Eurasian     
         

 

  Beringia
              

        
   

  Arctic North America
            


  North America



2.5 |amovas

  
(amova          
       n  
      

           
    
         
    amova     R   
            
(p   


G
ij =2rarcsin (
sin ((𝜑j𝜑i)2)2+cos (𝜑i) cos (𝜑j) sin ((𝜆i𝜆j)2)2
)
6 
|
   LOOG et aL.
2.6 | Demographic scenarios


 
   
      
             
        

            
       
          



    
      

 
2.7 | Population genetic coalescent framework


      
           K   
K


m 



K
K (K1K2 and
K       

    

         
    

           

              

time T
     

ΔT




       


       



 x 
x = 1

and x


K

2.8 | Approximate Bayesian Computation analysis
         
abctoolbox 
  
         

  
beast

        
         
         
          
     

       

         
F

                
      






        


  
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 7
LOOG et aL .
              


  


              
 





 mK
    m    K1 K2 K in

     m    K    x in
T    ΔT         
    

T

       m in
K1K2KxTΔT in

m
TΔTKK1K2 and K
xT and ΔT       

           
     
   

abc tool box

    
abc-
tool box manual.
            
         
     
    ΔTT10K1 10K2
10K10mx


2.9| Map plots

         
     
     
       

3 | RESULTS
3.1 | Population structure of grey wolf across the
Northern Hemisphere

           
              
           
ρ p  
        
              



             

   

    

     




a mova
   
 amova p     
   


     
      

3.2 | Bayesian phylogenetic analysis



       
 be ast
       


8 
|
   LOOG et aL.


R2
     
              

      

    

       
        

       
     



             



  
    
  
              


  
          

             
    



          

3.3 | Spatiotemporal reconstruction of past grey
wolf demography


 
       


    

     
            

 

         
        
         
       
      


   


            







       


      
              


         
           
 

       

             
            

              
     
 
         

            

4 | DISCUSSION
4.1 | Geographical origin of the ancestral wolf
population
     
Canis
  
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 9
LOOG et aL .
lupus  

         
    
     

      






    

        
              

     
 

       
FIGURE 2 bea st tree





bea st





Age [1,000 years BP]
200406080 100 120
Europe
Middle-East
Central-North Eurasia
East Eurasia
Beringia
North-America
Arctic North-America
Outgroup
Age [1,000 years BP]
Effective population size
(a)
(b)
020406080100 120140
160
10
3
10
4
10
5
10
6
FIGURE 3 







(b)
Europe
Central
North
Eurasia
Beringia
Middle-
East
East Asia
North
America
Arctic
North
America
(a)
10 
|
   LOOG et aL.
 



  

       

            
       
      


        
 

  


         

       
 
       



 

4.2 | Implications for the evolution of grey
wolf morphology




     
             
 
        
             
     
FIGURE 4 

staticexpansion
population size change





Static Population size
changes
Expansions
Expansions and
population
size changes
Time
Past
Present
Static
Population
size
changes
Expansions Expansions and population
size changes
Expansion origins
Europe
Middle-East
Central-North Eurasia
East Eurasia
Beringia
North-America
Arctic North-America
(a)
(b)
Bayes factor
0.0 0.20.4 0.6 0.81.0
  
|
 11
LOOG et aL .



        
         
             




      
        
      
     
     
   
      
   


         

       
         
           
   
              
      
     


            


         
  
     
      
  

FIGURE 5 




(a)
(b)
Ef
fective population size
Time [1,000 years BP]
Migration rate
[Nr. per 1,000 years]
Starting time of expansion
[1,000 years BP]
Prob. density
(c)
(d)
Prob. density
12 
|
   LOOG et aL.
       
       
  


        



     
             
    
        
      

              
    

     


          


4.3 | Implications for the study of wolf
domestication
             
  
  
   

            

    
          

    

       


      
        
   
       



         





into account.
ACKNOWLEDGEMENTS
        

             
            
        

              
       
        


        
   

            
        
  
        
        
             
              
    
       
   
       
            
 
       
     
       
     
 

            
 


AUTHOR CONTRIBUTIONS
           

        
            
    
           
      
  
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LOOG et aL .
      
        
  

        
        
   





  

DATA AVA ILAB ILITY STATE MEN T
       
    
         
           





ORCID
Liisa Loog 
Mikkel-Holger S. Sinding 

Gontran Sonet 
Anders Eriksson 
REFERENCES
          
       
     Journal of Zoological
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   
     ETOPO1 1 Arc-Minute Global Relief
Model: Procedures, Data Sources and Analysis  
     

        
        
  Science 306  



Russian Journal of Theriology8 

          
     Genetics 162

             
Quaternary Science Reviews117

    
 
 Proceedings of the National Academy of Sciences 109
 




           
     
Journal of Archaeological Science39
                
          
        Panthera spelaea    
       PLoS ONE 11    


   
          Scientific Reports 5 


 
Genetics161


Molecular Biology and
Evolution29
 
    
PLoS ONE8

                  
  
     
Proceedi ngs of the National Academy of Sciences 109 
 

             
 Proceedings of the National Academy
of Science s109

        
  Physical Biology 1  
 
  

  Genetic s
131
      
        
 Molecular Biology and Evolution 16  

 
        
Genome Research26  

            
     Quaternary
Science Reviews 96  
irev.2014.04.015
           

 
  Palaeogeography, Palaeoclimatology, Palaeoecology
261
14 
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   LOOG et aL.

        
       Science 352 

         
        
     PLoS Genetics 10
 
                 
Nature534

          
        
  Molecular Ecology 13  


          
       
Journal of Archaeological Science78

          
             
 Arctic Anthropolog y 44  
arc.2011.0 015
     
    
      Quaternary International 359–
360
        
     
Journal of Archaeological Science39


   
             
   Journal of Archaeological Science
36
                  
         
        
  Molecular Ecology 20     
   
       
       
         Canis.
Current Biology 28  
cub.2018.08.041
        
     
Homo sapiensEvolutionary Anthropology24

Current Biology19
R584–R594.
Paleoecology of Beringia



Genetics49
       Stochastic Processes
and their Applications 13  

         

  
(Canis lupusJournal of Biogeography43

       Materiali po
mezozoickoi i kainozoickoi istorii nazemnykh pozvonochnykh
         


Molecular Biolog y and Evolution29

 
    
Bioinformatics23   

 
              
     Proceedings of
the National Academy of Sciences 109  

Evolutionary
Ecology Research16 
       
     
Current Biology17



  Comptes Rendus Geoscience 340 


        
    Proceedings of the National Academy
of Science s 114  

 
 
   Nature 479
 
          
       Canis lupus 
   Molecular Ecology 13    

             
        
     Theoretical Population
Biology104
               
       
   
  Heredity 116  

            
 Journal of Archaeological Science 52  



 International Journal of Osteoarchaeology14
 
            
 Molecular Ecology 18   
   
       

      Journal of Mammalogy 94 

     
              
  
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 15
LOOG et aL .
       Proceedings of the Royal Society
B: Biological Sciences 280   
 
          
   Bioinformatics20 

      
Journal of Archaeological Science68


                                               
Molecular Ecology
15 
 
   
       
     Current Biology 26   

          
        
        
Quaternary International 420  

               
             
       
Science 349  

        
    
  Bioinformatics 16  


   
        Molecular Biology and Evolution
31
                
Current Anthropolog y43

      
     Proceedings of the Royal Society of
London. Series B: Biological Sciences271

       
 
             
 PLoS Genetics 14   

Sk y and Telescope68
                

   Current Biology25 

              
  
 Quaternary International 212  

                  
        
 Nature 431  
natur e02890
                
 

Science342
                                                      
   Canis
lupusMolecular Ecology
15


Cell Research26
 


     Proceedings of the National
Academy of Sciences 109  

       
       
  BMC Bioinformatics 11  
 
SUPPORTING INFORMATION
        
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How to cite this article:

Mol Ecol.
2019;00:1–15. 
... The species has a Holarctic distribution covering a wide range of environments, from tundra to deserts. Based on mitogenomes, Loog et al. (2020) suggested that extant wolf populations trace their origin to an expansion of the Beringian population starting ca. 25 thousand years ago (ka) towards Eurasia and later, ca. 15 ka, towards North America. This expansion led to the replacement of indigenous Pleistocene wolf populations (Loog et al., 2020). ...
... Based on mitogenomes, Loog et al. (2020) suggested that extant wolf populations trace their origin to an expansion of the Beringian population starting ca. 25 thousand years ago (ka) towards Eurasia and later, ca. 15 ka, towards North America. This expansion led to the replacement of indigenous Pleistocene wolf populations (Loog et al., 2020). An alternative scenario for North America is that extant wolves on the continent descend from a pre-LGM expansion out of Beringia followed by a post- ...
... LGM northward expansion from a glacial refugium south of the Laurentide/Cordilleran ice sheet complex (Koblmüller et al., 2016;Meachen et al., 2020). Overall, mitochondrial signatures illustrate a complex phylogeographic history for extant North American and Eurasian populations and high genetic similarity between the wolves of Alaska and Eastern Russia (Ersmark et al., 2016;Koblmüller et al., 2016;Loog et al., 2020). Nuclear data, however, show two reciprocally monophyletic wolf clades (Fan et al., 2016;Pilot et al., 2019;Sinding et al., 2018), with signs of inter-continental admixture between East Siberian and Alaskan populations (Sinding et al., 2018), consistent with their higher geographic proximity to the area of common origin. ...
Article
Glacial and interglacial periods throughout the Pleistocene have been substantial drivers of change in species distributions. Earlier analyses suggested that modern grey wolves (Canis lupus) trace their origin to a single Late Pleistocene Beringian population that expanded east and westwards, starting ca. 25,000 years ago (ya). Here, we examined the demographic and phylogeographic histories of extant populations around the Bering Strait with wolves from two inland regions of the Russian Far East (RFE) and one coastal and two inland regions of North‐western North America (NNA), genotyped for 91,327 single nucleotide polymorphisms. Our results indicated that RFE and NNA wolves had a common ancestry until ca. 34,400 ya, suggesting that these populations diverged before the previously proposed expansion out of Beringia. Coastal and inland NNA populations diverged ca. 16,000 ya, concordant with the minimum proposed date for the ecological viability of the migration route along the Pacific Northwest coast. Demographic reconstructions for inland RFE and NNA populations reveal spatial and temporal synchrony, with large historical effective population sizes that declined throughout the Pleistocene, possibly reflecting the influence of broad‐scale climatic changes across continents. In contrast, coastal NNA wolves displayed a consistently lower effective population size than the inland populations. Differences between the demographic history of inland and coastal wolves may have been driven by multiple ecological factors, including historical gene flow patterns, natural landscape fragmentation, and more recent anthropogenic disturbance.
... To date, however, studies of ancient DNA from Pleistocene wolf fossils in East Asia, including Japan, have been limited because of extensive DNA degradation in the hot and humid environment. 14,15 Although the Pleistocene wolves, for which ancient DNA analyses have been carried out, were quite biased toward high latitudes (e.g., remains found in Russia and Alaska), 16,17 little is known about their phylogenetic positions and population dynamics in East Asia. Therefore, the Pleistocene wolves in Asia offer an opportunity to use genomics to further our understanding of the population dynamics of ancient wolves throughout the northern hemisphere. ...
... ka). In agreement with previous studies, 16,17 the maternal lineage of the Japanese wolf was relatively one of the earlydiverging groups among the major extant populations of modern wolves in North America (Alaska, Canada, Mexico, etc.) and Eurasia (Siberia, East Asia, North East, etc.) including dogs ( Figure S2A). Because coalescent times can, in general, change depending on the dataset, the evolutionary timescale is also estimated based on the two datasets for the 13 protein-coding genes and the third codon positions of those genes, the latter of which is suggested to be little affected by the time dependency of the evolutionary rates. ...
... Previous analyses of ancient mtDNA have suggested that the Pleistocene gray wolves of the stem groups, which diverged 90-45 ka from the common ancestral lineage of the modern wolf clade, have been replaced by modern wolves. 16,17,27 However, we here demonstrated that both the Pleistocene (35 ka) and Holocene (5 ka) Japanese wolf specimens showed a genetic association with the Siberian Pleistocene wolves. Furthermore, we unexpectedly found that the Japanese wolf reflects an ancient hybridization event between the Japanese Pleistocene wolves and ancestral wolf populations from the continent rather than a simple replacement. ...
Article
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Little is known about the spatiotemporal dynamics of gray wolves in the Pleistocene across low-latitude regions of Eurasia. In Japan, a small-bodied endemic subspecies of Japanese wolves existed and went extinct in the early 1900s. The fossil record indicates that a giant wolf, which reached 70 cm in body height, inhabited Japan during the Pleistocene, but its evolutionary relationship, if any, with the Japanese wolf remains uncertain. Here, to reveal the genetic origin of the Japanese wolf, we analyzed ancient DNA from remains (recovered in Japan) of one Pleistocene wolf that lived 35,000 years ago and one Holocene wolf from 5,000 years ago. The analysis of the mitochondrial DNA revealed that the Pleistocene wolf was not part of the Japanese wolf clade but rather an earlier-diverging lineage. The analysis of the nuclear DNA of the Holocene Japanese wolf revealed that it was an admixture of the Japanese Pleistocene wolf and continental wolf lineages. These findings suggest that the Japanese wolf originated via waves of colonization of multiple Pleistocene wolf populations at 57–35 and 37–14 ka, respectively, followed by interpopulation hybridization.
... C. lupus occupies large parts of Europe, with C. l. signatus (Loog et al., 2020) found in the Iberian peninsula and C. l. italicus (Altobello, 1921) in Italy, France and Switzerland (Boitani et al., 2018). C. lupus dominates Asia according to Boitani et al. (2018) from Mongolia across China and the Himalayan Mountains, and C. l. chanco, the Himalayan wolf, is mentioned as proposed for the Himalayan range. ...
... Frontiers in Ecology and Evolution | www.frontiersin.org ecological changes of the time (Koblmüller et al., 2009;Ersmark et al., 2016;Loog et al., 2020). Within the Holarctic gray wolf complex, the highest diversity is found in wolves from Europe, China, and Russia (Ersmark et al., 2016). ...
... The Indian wolf shows divergent mtDNA haplotypes, mitogenomes, and evolutionary distinct genomes forming a monophyletic lineage basal to the Holarctic gray wolf complex (Figure 2; Sharma et al., 2004;Aggarwal et al., 2007;Pilot et al., 2010;Fan et al., 2016;Werhahn et al., 2017b;Loog et al., 2020;Hennelly et al., 2021). Aggarwal et al. (2007) analyzed five samples for D-loop mtDNA (1,140 bp) and 16S rRNA gene (560 bp), and two samples for cytochrome b (1,300 bp). ...
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Taxa belonging to the Genus Canis can challenge taxonomists because species boundaries and distribution ranges are often gradual. Species delineation within Canis is currently not based on consistent criteria, and is hampered by geographical bias and lack of taxonomic research. But a consistent taxonomy is critical, given its importance for assigning legal protection, conservation priorities, and financial resources. We carried out a qualitative review of the major wolf lineages so far identified from Asia from historical to contemporary time and considered relevant morphological, ecological, and genetic evidence. We present full mitochondrial phylogenies and genetic distances between these lineages. This review aims to summarize the available data on contemporary Asian wolf lineages within the context of the larger phylogenetic Canis group and to work toward a taxonomy that is consistent within the Canidae. We found support for the presence and taxon eligibility of Holarctic gray, Himalayan/Tibetan, Indian, and Arabian wolves in Asia and recommend their recognition at the taxonomic levels consistent within the group.
... Present-day sled dogs are closely related to Siberian 9,500-year-old dogs and 33,000year-old Pleistocene Siberian wolves, but not modern American wolves. The importance of the Beringia wolf expansion during the last glacial maximum is demonstrated by Loog et al. (2019): ...
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This paper provides an overview of wolf research in Russia at the beginning of the 21st century. Wolf research covered various directions, including population density estimation, management methods and minimization of human-wildlife conflicts, general and behavioral ecology, behavior, wolf population genetics and morphology, paleontology, dog domestication, helminthology and the wolves’ role in the rabies transmission. Some studies are performed with state-of-art methodology using molecular genetics, mathematical modeling, camera traps, and GPS telemetry.
... Future applications could include examining ancient Ychromosome and mitochondrial lineages in ancient cattle (Verdugo et al. 2019), wolves, and dogs (Bergström, Frantz, et al. 2020;Loog et al. 2020), for which large turnovers have occurred. ...
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Joint phylogenetic analysis of ancient DNA (aDNA) with modern phylogenies is hampered by low sequence coverage and post-mortem deamination, often resulting in over-conservative or incorrect assignment. We provide a new efficient likelihood-based workflow, pathPhynder, that takes advantage of all the polymorphic sites in the target sequence. This effectively evaluates the number of ancestral and derived alleles present on each branch and reports the most likely placement of an ancient sample in the phylogeny and a haplogroup assignment, together with alternatives and supporting evidence. To illustrate the application of pathPhynder, we show improved Y chromosome assignments for published ancient DNA sequences, using a newly compiled Y variation dataset (120,908 markers from 2,014 samples) that significantly enhances Y haplogroup assignment for low coverage samples. We apply the method to all published male ancient DNA samples from Africa, giving new insights into ancient migrations and the relationships between ancient and modern populations. The same software can be used to place samples with large amounts of missing data into other large non-recombining phylogenies such as the mitochondrial tree.
... Nevertheless, to increase statistical power for the monitoring of genetic erosion in subsequent studies and reduce potential effects of ascertainment bias, we recommend to aim for a higher number of SNP markers (min. several hundred; e.g.,Cammen et al., 2018;Ewart et al., 2019;Stronen et al., 2019a), consider secondgeneration sequencing of microsatellites(Curto et al., 2019), or data from WGS(Larsson et al., 2019;Loog et al., 2020;White et al., 2018). ...
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Like many carnivore species, European wildcats (Felis silvestris) have suffered severe anthropogenic population declines in the past, resulting in a strong population bottleneck at the beginning of the 20th century. In Germany, the species has managed to survive its near extinction in small isolated areas and is currently recolonizing former habitats owing to legal protection and concerted conservation efforts. Here, we SNP-genotyped and mtDNA-sequenced 56 historical and 650 contemporary samples to assess the impact of massive persecution on genetic diversity, population structure, and hybridization dynamics of wildcats. Spatiotemporal analyses suggest that the presumed postglacial differentiation between two genetically distinct meta-populations in Germany is in fact the result of the anthropogenic bottleneck followed by re-expansion from few secluded refugia. We found that, despite the bottleneck, populations experienced no severe genetic erosion, nor suffered from elevated inbreeding or showed signs of increased hybridization with domestic cats. Our findings have significant implications for current wildcat conservation strategies, as the data analyses show that the two presently recognized wildcat population clusters should be treated as a single conservation unit. Although current populations appear under no imminent threat from genetic factors, fostering connectivity through the implementation of forest corridors will facilitate the preservation of genetic diversity and promote long-term viability. The present study documents how museum collections can be used as essential resource for assessing long-term anthropogenic effects on natural populations, for example, regarding population structure and the delineation of appropriate conservation units, potentially informing todays' species conservation.
... In conjunction with ABC-based analyses, we identified patterns of post-glacial colonization in formerly glaciated regions that contradict previously proposed routes for the red fox and other similar species. Given the genomic resources now available (Kukekova et al., 2018), the application of ancient genomics on the extensive fossil material available for this species (Sommer & Benecke, 2005) should fall into line with other charismatic carnivores (Loog et al., 2020) to fully understand re-colonization and temporal patterns that have not been captured in previous studies of ancient red fox specimens Teacher et al., 2011). ...