An understanding of food-web persistence from local to global scales

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1 An understanding of food-web persistence from local to global scales Daniel B. Stouffer Integrative Ecology Group Estación Biológica de Doñana - CSIC Sevilla, Spain stouffer@ebd.csic.es CABDyN Seminar Series

2 Invasive species in the Great Lakes

3 Invasive species in the Great Lakes Zebra mussel

4 Invasive species in the Great Lakes Zebra mussel

5 Invasive species in the Great Lakes Northern snakehead

6 Invasive species in the Great Lakes Asian carp

7 Invasive species in the Great Lakes Asian carp

8 Challenge of solving these problems

9 Challenge of solving these problems A true complex system

10 Challenge of solving these problems A true complex system Experimentation is impossible (or impractical)

11 Challenge of solving these problems A true complex system Experimentation is impossible (or impractical) A modeling approach is imperative

12 What makes a food web stable? Allesina and Pascual, Theor. Ecol. (2007) and Otto et al., Nature (2007)

13 What makes a food web stable? Complexity-stability debate Stability of small sub-webs Allesina and Pascual, Theor. Ecol. (2007) and Otto et al., Nature (2007)

14 What makes a food web stable? Complexity-stability debate More diversity and more connections = Greater stability More diversity and more connections = Lower stability Greater empirical realism increases stability Stability of small sub-webs Allesina and Pascual, Theor. Ecol. (2007) and Otto et al., Nature (2007)

15 What makes a food web stable? Complexity-stability debate More diversity and more connections = Greater stability More diversity and more connections = Lower stability Greater empirical realism increases stability Stability of small sub-webs Omnivory is a stabilizing force Weak interactions confer stability Predator-prey body size ratios Allesina and Pascual, Theor. Ecol. (2007) and Otto et al., Nature (2007)

16 What makes a food web stable? Complexity-stability debate More diversity and more connections = Greater stability More diversity and more connections = Lower stability Greater empirical realism increases stability Stability of small sub-webs Omnivory is a stabilizing force Weak interactions confer stability Predator-prey body size ratios What is meant by stability? Allesina and Pascual, Theor. Ecol. (2007) and Otto et al., Nature (2007)

17 What makes a food web stable? Complexity-stability debate More diversity and more connections = Greater stability More diversity and more connections = Lower stability Greater empirical realism increases stability Stability of small sub-webs Omnivory is a stabilizing force Weak interactions confer stability Predator-prey body size ratios What is meant by stability? Return to equilibrium after perturbation Stabilization of dynamics Greater species persistence Allesina and Pascual, Theor. Ecol. (2007) and Otto et al., Nature (2007)

18 Food-web structure and persistence What is the role of food-web structure on persistence?

19 Food-web structure and persistence What is the role of food-web modules?

20 Food-web structure and persistence What is the role of food-web modules? How does persistence of food-web modules in isolation relate to their influence within community food webs?

21 Universal function forms for distributions of numbers of prey, predators, and links A key to the success of leading static food-web models Cumulative distribution Number of prey, k/2z Number of predators, m/2z Number of links, r/2z Stouffer et al., Ecology (2005)

22 Ingredients for successful food-web model Stouffer et al., Ecology (2005)

23 Ingredients for successful food-web model Species can be ordered in one dimension Stouffer et al., Ecology (2005)

24 Ingredients for successful food-web model Species can be ordered in one dimension Prey selection Stouffer et al., Ecology (2005)

25 Ingredients for successful food-web model Species can be ordered in one dimension Prey selection: Random Stouffer et al., Ecology (2005)

26 Ingredients for successful food-web model Species can be ordered in one dimension Prey selection: Contiguous Stouffer et al., Ecology (2005)

27 Ingredients for successful food-web model Species can be ordered in one dimension Prey selection: Contiguous Stouffer et al., Ecology (2005)

28 Ingredients for successful food-web model Species can be ordered in one dimension Prey selection: Contiguous Stouffer et al., Ecology (2005)

29 Prey selection mechanism Random predation Contiguous predation Stouffer et al., PRSB (2007)

30 Prey selection mechanism Random predation Predators are indifferent to the identity of their prey Contiguous predation Stouffer et al., PRSB (2007)

31 Prey selection mechanism Random predation Predators are indifferent to the identity of their prey Contiguous predation Predators specialize on species which have some characteristic features Stouffer et al., PRSB (2007)

32 Prey selection mechanism Random predation Predators are indifferent to the identity of their prey Contiguous predation Predators specialize on species which have some characteristic features Is there a signature in the data indicating the empirically observed mechanism? Stouffer et al., PRSB (2007)

33 Network motifs Complete set of unique connected triplets of species Milo et al., Science (2002)

34 Significance of network motifs Just the appearance of motifs is not significant Compare to null hypothesis of a randomized network Stouffer et al., PRSB (2007)

35 Significance of network motifs Just the appearance of motifs is not significant Compare to null hypothesis of a randomized network Could the observed motif pattern occur at random? Stouffer et al., PRSB (2007)

36 Significance of network motifs Just the appearance of motifs is not significant Compare to null hypothesis of a randomized network Could the observed motif pattern occur at random? 1.0 Normalized profile S1 S2 S3 S4 S5 D1 D2 D3 D4 D5 D6 D7 D8 Stouffer et al., PRSB (2007)

37 Significance of network motifs Just the appearance of motifs is not significant Compare to null hypothesis of a randomized network Could the observed motif pattern occur at random? 1.0 Normalized profile S1 S2 S3 S4 S5 D1 D2 D3 D4 D5 D6 D7 D8 Stouffer et al., PRSB (2007)

38 Significance of network motifs Just the appearance of motifs is not significant Compare to null hypothesis of a randomized network Could the observed motif pattern occur at random? 1.0 Normalized profile S1 S2 S3 S4 S5 D1 D2 D3 D4 D5 D6 D7 D8 Stouffer et al., PRSB (2007)

39 Significance of network motifs Just the appearance of motifs is not significant Compare to null hypothesis of a randomized network Could the observed motif pattern occur at random? 1.0 Normalized profile S1 S2 S3 S4 S5 D1 D2 D3 D4 D5 D6 D7 D8 Stouffer et al., PRSB (2007)

40 Food-web modules and persistence

41 Food-web modules and persistence What is the relationship between persistence of modules in isolation and their influence on community food-web persistence?

42 Food-web modules and persistence How does a module s influence on community food-web persistence relate to its presence in community food-webs?

43 Food-web modules and persistence We will model module and food-web dynamics and examine the consequences of structure

44 Modeling food-web dynamics

45 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters db i dt db i dt = r i G i B i x k y k B k F ki e ki k=pred = x i B i + x i B i y i F ij j=prey k=pred x k y k B k F ki e ki Yodzis and Innes, Am. Nat. (1992) Brose et al., Ecol. Lett. (2006)

46 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs:

47 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Williams and Martinez, Nature (2000)

48 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions

49 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions

50 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Species body masses

51 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Species body masses

52 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Species body masses Interaction strengths

53 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Species body masses Interaction strengths

54 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Species body masses Interaction strengths What is the persistence after t timesteps?

55 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Species body masses Interaction strengths What is the persistence after t timesteps?

56 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Species body masses Interaction strengths What is the persistence after t timesteps?

57 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Species body masses Interaction strengths What is the persistence after t timesteps?

58 Modeling food-web dynamics Bioenergetic population dynamics model Allometric scaling of metabolic parameters Required inputs: Network of interactions Species body masses Interaction strengths What is the persistence after t timesteps?

59 Persistence of isolated modules Tri trophic chain Omnivory Exploitative competition Apparent competition

60 Persistence of isolated modules Tri trophic chain Omnivory Exploitative competition Apparent competition Constitute 95% of empirically observed modules

61 Persistence of isolated modules 1.00 Fraction of persistent modules Timesteps

62 Persistence of isolated modules Tri trophic chain Omnivory Exploitative competition Apparent competition

63 Persistence of isolated modules Tri trophic chain Omnivory Exploitative competition Apparent competition Expectation based upon isolated persistence

64 Persistence of isolated modules Tri trophic chain Omnivory Exploitative competition Apparent competition > > > Expectation based upon isolated persistence

65 Community food-web persistence How does presence of modules relate to persistence?

66 Community food-web persistence How does presence of modules relate to persistence? Generate a food web, assign masses, assign interaction strengths...

67 Community food-web persistence How does presence of modules relate to persistence? Generate a food web, assign masses, assign interaction strengths... The food web has some number of each module: Tri-trophic food chain Omnivory Exploitative competition Apparent competition

68 Community food-web persistence How does presence of modules relate to persistence? Generate a food web, assign masses, assign interaction strengths... The food web has some number of each module: Tri-trophic food chain Omnivory Exploitative competition Apparent competition How does the number of each module present influence the food web s persistence?

69 Community food-web persistence 0.4 Model residuals Conditional probability density N module

70 Community food-web persistence 0.4 Model residuals Conditional probability density N module

71 Community food-web persistence 0.5 Model residuals Conditional probability density N module

72 Community food-web persistence 0.2 Model residuals Conditional probability density N module

73 Community food-web persistence 0.2 Model residuals Conditional probability density N module

74 Persistence profile 0.2 Persistence factor

75 Conclusions

76 Conclusions Persistence of isolated modules is not the same as the effect within community food webs

77 Conclusions Presence of modules has clear influence on community food web persistence

78 Conclusions Presence of modules has clear influence on community food web persistence Strongly related to empirical observations

79 Implications

80 Implications There may be significant dynamic justifications for observed food-web structure

81 Implications There may be significant dynamic justifications for observed food-web structure Caution must be taken when attempting to scale up from modules to community food webs

82 Implications There may be significant dynamic justifications for observed food-web structure Caution must be taken when attempting to scale up from modules to community food webs Species appear to participate in interactions which maximize community persistence and not necessarily their own persistence

83 Implications 1.0 Normalized profile S1 S2 S3 S4 S5 D1 D2 D3 D4 D5 D6 D7 D8 Some empirical food webs exhibit fewer instances of omnivory and greater instances of exploitative and apparent competition

84 Implications 1.0 Normalized profile S1 S2 S3 S4 S5 D1 D2 D3 D4 D5 D6 D7 D8 We hypothesize that these food webs are less persistent and more vulnerable to perturbation

85 Implications Management decisions Invasive species

86 Implications Management decisions What motifs does a species participate in? Invasive species

87 Implications Management decisions What motifs does a species participate in? Invasive species What motifs is an invasive likely to participate in?

88 Acknowledgments Luís Amaral, Juan Camacho, Wenxin Jiang Jordi Bascompte Members of the Integrative Ecology Group and Amaral Lab NSF IGERT Graduate Research Fellowship NU ChBE Teaching Apprenticeship fellowship CSIC JAE Post-doctoral Fellowship Centro de Supercomputación de Galicia (CESGA)

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