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Quantifying social attributes and their
consequences in yellow-bellied marmots
Tina W. Wey1
& Daniel T. Blumstein2
1
University of California, Davis; twwey@ucdavis.edu
2
University of California, Los Angeles; marmots@ucla.edu
Tom Uhlman
Causes and consequences of
sociality
 Sociality is widespread and varied
 Questions
 Why is there social variation?
 What are the consequences of social variation?
 Multiple levels
 Proximate and ultimate causes of sociality
 Individual and group variation
 Many ways to quantify sociality
Interactions are important
INTERACTIONS
RELATIONSHIPS
SOCIAL STRUCTURE
Hinde 1976
Social network variation
Outline
 Social network structure and consequences in
yellow-bellied marmots
 How are marmot social networks structured at
multiple levels?
 Do individual social attributes have fitness
correlates?
Yellow-bellied marmots
Rocky Mountain Biological Laboratory
Marmota flaviventris
Fieldwork
 Data
 Trapping
Adriana Maldonado-Chaparo
Karisa Tang
Fieldwork
 Data
 Observations
 2003-2008
Brian Smith
Carol Glazer
Marmot social networks
Quantifying network variation
 Affiliative (cohesive, “friendly”) vs. agonistic (competitive,
“unfriendly”) interactions
 Multiple levels of analysis
 Networks
 Dyads
 Individuals
vs
Network structure – “Friendly” and
“unfriendly” networks differ
Measure Affiliative Agonistic
Network structure – “Friendly” and
“unfriendly” networks differ
Measure Affiliative Agonistic
Density
N = 24
Z = -3.741
P < 0.001
Network structure – “Friendly” and
“unfriendly” networks differ
Measure Affiliative Agonistic
Density
Reciprocity
N = 24
Z = -3.741
P < 0.001
N = 24
Z = -3.657
P < 0.001
Network structure – “Friendly” and
“unfriendly” networks differ
Measure Affiliative Agonistic
Density
Reciprocity
Clustering
coefficient
N = 24
Z = -3.741
P < 0.001
N = 24
Z = -3.657
P < 0.001
N = 24
Z = -0.343
P < 0.001
Dyadic structure
Wey & Blumstein 2010 Animal Behaviour
 Age and kinship influence affiliation preference
 Marmots are more friendly to others in the same age class and
closer relatives
 Less consistent “rules” for who fights
Friendship Fighting
Age class
Sex
Kinship
Individual variation
 Age and sex influence social tendencies
 Older and male marmots are crankier
 Older marmots receive fewer affiliations, and initiate more and
receive less aggression
 Females receive affiliation from more partners, and males initiate
aggression to more
Affiliative Agonistic
Initiation Reception Initiation Reception
Age
Sex
Wey & Blumstein 2010 Animal Behaviour
Social variation can have
associated consequences
 Individual social variation influences reproductive
success
 Social bonds can improve fitness, especially for females
 Aggression and dominance can result in differential fitness
 Increased parasitism is often a cost of sociality
 Increase with group size (Côté & Poulin 1995, Bordes et al. 2007)
 Social behavior is expected to influence parasites (Møller et al.
1993, Altizer et al. 2003)
Consequences of individual social
variation in marmots
 Background:
 Dominance rank is associated with reproductive success in
males but not females (Huang et al. 2011 Ethology)
 Larger males tend to be dominant
 Social cohesion predicts female dispersal (Blumstein et al. 2009 Proc B)
 Social variation is heritable (Lea et al. 2010 PNAS)
 Predictions:
 Social cohesion would improve female reproductive success
 Aggression would predict males reproductive success
 More social individuals would have more parasites
Consequences of individual social
variation in marmots
 Defined social attributes
 Measured fitness correlates
 Annual reproductive success
 Parasites – fleas and intestinal parasites
 Fit separate mixed models with social variables as
predictors
 Female and male reproductive success analyzed separately
 Included a spatial component
 Burrow overlap
Defining social attributes
 Many possibly measures of connectivity, which reflect different but
related aspects of social connectivity
Degree Number of social partners
Out-degree Number social partners initiating to
In-degree Number social partners receiving from
Strength Total interactions (weighted degree)
Out-strength Total interactions initiated
In-strength Total interactions received
Closeness Overall connectivity in the network
Betweenness Importance as a connection point in the network
Embeddedness Social integration
Defining social attributes
Network Social Variable Description
Affiliative PC1: Connectivity Overall affiliative
connectedness
PC2: Affiliation
strength
Weighted affiliation
Agonistic PC1: Victimization Received aggression,
embeddedness
PC2: Bullying Initiated aggression,
Betweenness
Spatial Overlap Crowding experienced
Social cohesion did not improve
female reproductive success
 Friendlier females had fewer offspring
 Crowded females had fewer offspring
Affiliation
strength
Annual
reproductive
success
Spatial
overlap
Annual
reproductive
success
N = 228
P < 0.001
N = 228
P = 0.014
Size and aggression were important
for male reproductive success
 Bigger males had more offspring
 More aggressive males had more offspring
Relative
mass
Annual
reproductive
success
Bullying
Annual
reproductive
success
N = 44
P < 0.001
N = 44
P = 0.044
Parasites were not a cost of
sociality
 Social factors did not influence ectoparasite or
endoparasite infection
 Ectoparasites: fleas (N = 185)
 Males had more fleas (P = 0.030)
 Endoparasites: gut protozoa (N = 428)
 No significant trends
Sociality Parasites
Summary
 Social attributes explained reproductive success, but not
parasites
 Friendlier females had lower reproductive success
 Contrasts with more highly social species
 Constraints and competition?
 Male bullies had higher reproductive success
 Consistent with results from dominance
 Parasites do not appear to be a cost of sociality in marmots
 Results differ from findings in other systems (Corner et al. 2003)
 Consistent with previous research on group measures in marmots
(Arnold & Lichtenstein 1993, Van Vuren 1996)
Conclusions
 Quantified and distinguished biologically
important attributes of sociality examine
structure and possible consequences
 Examining social network variation offered
complimentary insights to other approaches
Acknowledgements
Collaborators
Brian Huang
Ferenc Jordán
Amanda Lea
Julien Martin
Weiwei Shen
Karisa Tang
Adriana Maldonado
Help & Feedback
Rick Grannis
Peter Nonacs
Peter Narins
Blumstein lab
UCLA ATS stats consulting

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Social attributes in yellow-bellied marmots

  • 1. Quantifying social attributes and their consequences in yellow-bellied marmots Tina W. Wey1 & Daniel T. Blumstein2 1 University of California, Davis; twwey@ucdavis.edu 2 University of California, Los Angeles; marmots@ucla.edu Tom Uhlman
  • 2. Causes and consequences of sociality  Sociality is widespread and varied  Questions  Why is there social variation?  What are the consequences of social variation?  Multiple levels  Proximate and ultimate causes of sociality  Individual and group variation  Many ways to quantify sociality
  • 5. Outline  Social network structure and consequences in yellow-bellied marmots  How are marmot social networks structured at multiple levels?  Do individual social attributes have fitness correlates?
  • 6. Yellow-bellied marmots Rocky Mountain Biological Laboratory Marmota flaviventris
  • 7. Fieldwork  Data  Trapping Adriana Maldonado-Chaparo Karisa Tang
  • 8. Fieldwork  Data  Observations  2003-2008 Brian Smith Carol Glazer
  • 10. Quantifying network variation  Affiliative (cohesive, “friendly”) vs. agonistic (competitive, “unfriendly”) interactions  Multiple levels of analysis  Networks  Dyads  Individuals vs
  • 11. Network structure – “Friendly” and “unfriendly” networks differ Measure Affiliative Agonistic
  • 12. Network structure – “Friendly” and “unfriendly” networks differ Measure Affiliative Agonistic Density N = 24 Z = -3.741 P < 0.001
  • 13. Network structure – “Friendly” and “unfriendly” networks differ Measure Affiliative Agonistic Density Reciprocity N = 24 Z = -3.741 P < 0.001 N = 24 Z = -3.657 P < 0.001
  • 14. Network structure – “Friendly” and “unfriendly” networks differ Measure Affiliative Agonistic Density Reciprocity Clustering coefficient N = 24 Z = -3.741 P < 0.001 N = 24 Z = -3.657 P < 0.001 N = 24 Z = -0.343 P < 0.001
  • 15. Dyadic structure Wey & Blumstein 2010 Animal Behaviour  Age and kinship influence affiliation preference  Marmots are more friendly to others in the same age class and closer relatives  Less consistent “rules” for who fights Friendship Fighting Age class Sex Kinship
  • 16. Individual variation  Age and sex influence social tendencies  Older and male marmots are crankier  Older marmots receive fewer affiliations, and initiate more and receive less aggression  Females receive affiliation from more partners, and males initiate aggression to more Affiliative Agonistic Initiation Reception Initiation Reception Age Sex Wey & Blumstein 2010 Animal Behaviour
  • 17. Social variation can have associated consequences  Individual social variation influences reproductive success  Social bonds can improve fitness, especially for females  Aggression and dominance can result in differential fitness  Increased parasitism is often a cost of sociality  Increase with group size (Côté & Poulin 1995, Bordes et al. 2007)  Social behavior is expected to influence parasites (Møller et al. 1993, Altizer et al. 2003)
  • 18. Consequences of individual social variation in marmots  Background:  Dominance rank is associated with reproductive success in males but not females (Huang et al. 2011 Ethology)  Larger males tend to be dominant  Social cohesion predicts female dispersal (Blumstein et al. 2009 Proc B)  Social variation is heritable (Lea et al. 2010 PNAS)  Predictions:  Social cohesion would improve female reproductive success  Aggression would predict males reproductive success  More social individuals would have more parasites
  • 19. Consequences of individual social variation in marmots  Defined social attributes  Measured fitness correlates  Annual reproductive success  Parasites – fleas and intestinal parasites  Fit separate mixed models with social variables as predictors  Female and male reproductive success analyzed separately  Included a spatial component  Burrow overlap
  • 20. Defining social attributes  Many possibly measures of connectivity, which reflect different but related aspects of social connectivity Degree Number of social partners Out-degree Number social partners initiating to In-degree Number social partners receiving from Strength Total interactions (weighted degree) Out-strength Total interactions initiated In-strength Total interactions received Closeness Overall connectivity in the network Betweenness Importance as a connection point in the network Embeddedness Social integration
  • 21. Defining social attributes Network Social Variable Description Affiliative PC1: Connectivity Overall affiliative connectedness PC2: Affiliation strength Weighted affiliation Agonistic PC1: Victimization Received aggression, embeddedness PC2: Bullying Initiated aggression, Betweenness Spatial Overlap Crowding experienced
  • 22. Social cohesion did not improve female reproductive success  Friendlier females had fewer offspring  Crowded females had fewer offspring Affiliation strength Annual reproductive success Spatial overlap Annual reproductive success N = 228 P < 0.001 N = 228 P = 0.014
  • 23. Size and aggression were important for male reproductive success  Bigger males had more offspring  More aggressive males had more offspring Relative mass Annual reproductive success Bullying Annual reproductive success N = 44 P < 0.001 N = 44 P = 0.044
  • 24. Parasites were not a cost of sociality  Social factors did not influence ectoparasite or endoparasite infection  Ectoparasites: fleas (N = 185)  Males had more fleas (P = 0.030)  Endoparasites: gut protozoa (N = 428)  No significant trends Sociality Parasites
  • 25. Summary  Social attributes explained reproductive success, but not parasites  Friendlier females had lower reproductive success  Contrasts with more highly social species  Constraints and competition?  Male bullies had higher reproductive success  Consistent with results from dominance  Parasites do not appear to be a cost of sociality in marmots  Results differ from findings in other systems (Corner et al. 2003)  Consistent with previous research on group measures in marmots (Arnold & Lichtenstein 1993, Van Vuren 1996)
  • 26. Conclusions  Quantified and distinguished biologically important attributes of sociality examine structure and possible consequences  Examining social network variation offered complimentary insights to other approaches
  • 27. Acknowledgements Collaborators Brian Huang Ferenc Jordán Amanda Lea Julien Martin Weiwei Shen Karisa Tang Adriana Maldonado Help & Feedback Rick Grannis Peter Nonacs Peter Narins Blumstein lab UCLA ATS stats consulting

Editor's Notes

  1. Multiple levels of interest
  2. Often assumed, but rarely measured explicitly.
  3. Multiple levels of interest
  4. And with this data we then start to measure and test questions about social variation.
  5. Predictions based on previous work in marmots and results form other animal and human social networks
  6. Patterns consistent with other systems. Implications. Dyadic relationships.
  7. Interpret in conjunction with earlier results.
  8. Mention dispersal results
  9. Mention dispersal results
  10. Mention dispersal results
  11. Interestingly, higher agonistic reception was not associated with lower reproductive success
  12. Despite predictions from theory, an empirical test in this system found no relationship, although it remains to be seen in other systems. May be incorrect time scale or resolution to capture dynamics