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松田裕之（横浜国大・環境情報） Hiroyuki Matsuda (Yokohama Nat’l Univ)

スイッチング捕食と天敵特異的防御がもたらす食物網構造と群集動態 More stories of community ecology with adaptive fish behaviors and adaptive fisheries management. 松田裕之（横浜国大・環境情報） Hiroyuki Matsuda (Yokohama Nat’l Univ). predator. carnivore. 3. 3. 2. 3. herbivore. 2. 1. 2. 1. 1. prey. plant.

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松田裕之（横浜国大・環境情報） Hiroyuki Matsuda (Yokohama Nat’l Univ)

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  1. スイッチング捕食と天敵特異的防御がもたらす食物網構造と群集動態More stories of community ecology with adaptive fish behaviors and adaptive fisheries management 松田裕之（横浜国大・環境情報） Hiroyuki Matsuda (Yokohama Nat’l Univ)

  2. predator carnivore 3 3 2 3 herbivore 2 1 2 1 1 prey plant 3 types of 3 species system

  3. 3 2 1 The paradox of pesticides Pesticide also attacks carnivore Pesticide attacks herbivore Herbivore will increase and the plant will decrease.

  4. 間接効果Indirect Effects • 個体数変化を通じた間接効果 Density-Mediated Indirect Effects • 行動や形質変化を通じた間接効果 Trait-Mediated Indirect Effects

  5. Increase predator 2 2 Decrease prey Decrease predator 3 1 3 Exploitative Competition 2 3 1

  6. Exploitative Competition • dN1/dt = (-d1 - b1N1 + a1R)N1 • dN2/dt = (-d2 - b2N2 + a2R)N2 • dR/dt = (d0 - a1N1 + a2N2)R • N1 = (a1b2d0+a1a2d2 -a22d1)/(a22b1+a12b2), • N2 = (a2b1d0+a1a2d1 -a12d2)/(a22b1+a12b2) • R = (b1b2d0+a1b2d1 -a2b1d2)/(a22b1+a12b2) • dN1*/dd2 > 0

  7. Increase prey 2 1 Increase predator Decrease prey 3 2 3 Apparent Competition 1 2 3

  8. Increase prey 2 1 Predator focuses on prey 2 Increase prey 3 2 3 Apparent mutualismAbrams & Matsuda 1996 Ecology 77:610-616 1 2 3

  9. Apparent Mutualism • Suppose Prey A & B and 1 Predator. • Prey A increases. • Predator focuses on A, consequently ignores B (Predator switching). • Fitness of Prey B may increase with A. • Few empirical data,

  10. Increase predator 2 2 3 Watch more against 2 Increase predator 3 Exploitative Mutualism(Matsuda et al. Oikos 1993, 68:549-559) 2 3 1

  11. Antipredator effort against predator 1 is … • [Nonspecific defense] effective against both predator species (types) 1 & 2; • [Partly-specific] partly effective against 2; • [Perfect-specific] not effective against 2 at all; • [overly-specific] riskier against 2 than when it pais no attention to any predator.

  12. How many points can you watch for simultaneously? Quiz by Japan Automobile Fedaration JAF News, the recent issue

  13. Yodzis(1988)の間接効果理論 • dN/dt = f(N, p)　　　群集動態 • dN/dt = (f/N) (N-N*) 　線形近似 • = C (N-N*) 群集行列 • f/p+(f/N)(N*/p)=0陰関数微分 • N*/p = –(f/N)-1(f/p) • = – C-1(f/p)

  14. Example: indirect effects in a 10 species system 10 8 9 5 6 7 1 2 3 4

  15. 群集行列Community Matrix 10 8 9 5 6 7 1 2 3 4

  16. 種 1 2 3 4 5 6 7 8 9 10 1 1000 101 953 511 0 934 511 653 658 157 2 101 1000 267 959 101 81 959 722 738 53 3 953 267 1000 112 953 81 112 747 728 51 10 4 511 959 112 1000 511 941 0 669 643 140 5 1000 899 47 489 1000 66 489 347 342 843 8 9 6 66 919 919 59 66 1000 59 402 420 733 5 6 7 7 489 41 888 1000 489 59 1000 331 357 860 8 653 722 747 669 653 598 669 1000 12 202 1 2 3 4 9 658 738 728 643 658 580 643 12 1000 188 10 843 947 949 860 843 733 860 798 812 1000 Sensitivity frequencyMatrix “–C-1 ”

  17. Kyoto Declaration and Plan of Action on the Sustainable Contribution of Fisheries to Food Security in 1992 (FAO) • Article 14 “When and where appropriate, consider harvesting multiple trophic levels in a manner consistent with sustainable development of these resources”. http://www.fao.org/fi/agreem/kyoto/kyoe.asp

  18. イワシxとマグロyの数理模型 • dx/dt = (r - a x - b y - f) x • dy/dt = (-d + e b x - g) y • Maximize total yield fx+pgy at the equilibrium

  19. Paradox of Kyoto Declaration • Optimal solution is either • to catch sardine after tuna goes extinct; or • to catch tuna only.

  20. 6 6 4 5 4 4 5 5 4 4 3 3 3 3 2 Examples of biological community at MSY (Matsuda & Abrams in review) Solution maximizing total yield from community MSY solution often reduces species and links; (d) (e) (c) (b) (a) 6 6 4 5 6 6 5 5 5 4 4 3 3 3 1 2 1 2 1 2 1 2 1 2

  21. 4 5 6 4 5 4 5 5 4 3 3 3 2 2 100% 92% 61% 12% 6% Examples of biological community at MSY (Matsuda & Abrams in review) Constrained MSY that guarantee coexistence exploit more species, more trophic levels. (d) (e) (c) (b) (a) 6 6 6 6 6 4 5 5 5 4 4 4 3 3 3 3 1 1 2 1 2 1 2 1 2

  22. Conclusion of story 2 • MSY theory does not guarantee species coexistence • Fisheries must take care of biodiversity conservation explicitly = Foodweb constraint to reconciling fisheries with conservation

  23. Requiem to Maximum Sustainable Yield Theory • Ecosystems are uncertain, non-equilibrium and complex. • MSY theory ignores all the three. • Does MSY theory guarantee species persistence? - No!! surplusproduction Stockabundance

  24. N* N* N* Feedback control in fishing effort is powerful... A straw man says; • Even though the MSY level is unknown, the feedback control stabilizes a broad range of target stock level. f(N) Stock size N

  25. 9 10 8 7 5 6 1 4 2 3 Feedback control with community interactions also result in undesired outcomes.(M & A in preparation) r = (0.454,1.059,1.186,0.247,-0.006,-0.028,-0.059,-0.704,-0.308,-0.238) A = (aji) = e9 = 0.1, ei = 0

  26. Feedback control may result in extinction of other species (sp. 6). ratio de9/dt = u(N9-N9*)

  27. Conclusion of story 3 • Single stock monitoring is dangerous • Target stock level is much more sensitive than we have considered in single stock models. • We must monitor not only stock level of target species, but also the “entire” ecosystem.

  28. Only 5 to 10 percent of us succeed of the weight-loss industry lantern fish deep sea Wasp-waist is a classic dream... , is this illusion? birds seals tunas • Anyway, we need to investigate how to fluctuate the total biomass of small pelagics. sardine/anchovy pelagic copepods krill ....

  29. 非定常群集nonequilibrial community • 環境が変化する Changing Environment • 個体数が変化する Unstable Population • 行動や形質が変化する • Change in Behavior & Traits • 餌選好や住み場所が変われば、群集構造も変化する • Change in Community Structure

  30. 共進化的に安定な群集Coevolutionarily stable community • dNj/dt = [-dj + ΣifjiajiRi]Nj • dRi/dt = [ri - biRi -ΣjfjiajiNj]Ri • tradeoff Σifji=1 • optimal prey preference ΣifjiajiRi maximize • at CSC, ajiRi = ajkRkｒif fji>0, fjk>0 • # equations = #links - #predator species

  31. Link-species scaling law (1) • # equations = # links - # predator species • # unknowns = # prey species R1 R2 R3

  32. Link-species scaling law (2) • # equations < # unknowns • # links (L) < # prey + # predator species • L < 2S (Cohen et al. 1993).

  33. Predator-specific defense enhances • Coexistence of predators. • A more complex community strucutre Food web in Lake Tanganyika Matsuda with Abrams & Hori (1994, 1996, Evol. Ecol)

  34. Polis’ opinion • Food web is • L is proportional to S2 • link-species scaling law is an artifact from short-term, narrow range observation.

  35. Foodweb changes temporallyMatsuda, H. & Namba, T. (1991) Ecology 72(1):267-276. predator prey

  36. 長期と短期を分けて考えようImportance of short-term structure • Temporal niche overlap is reasonable for abundant resource • Predator may avoid short-term competition. (behavioral response) • It is different from long-term coexistence and population dynamics

  37. 非定常群集nonequilibrial community • 環境が変化する Changing Environment • 個体数が変化する Unstable Population • 行動や形質が変化する Change in Behavior & Traits • 餌選好や住み場所が変われば、群集構造も変化するChange in Community Structure

  38. back back Lateral dimorphism of scale eating cichlids in Lake Tanganyika Hori 1991 Science 267 Righty Lefty

  39. “Antisymmetry” Fluctuating asymmetry (FA) Directive asymmetry (DA) Three types of Asymmetries(van Valen 1962)  

  40. Antisymmetry in fishes • Scale-eating cichlid in Lake Tanganyika • Lefties feed on scales of the right side, righties feed on scales of the left side • Frequency dependent natural selection • Hori 1991 Science 267: • Maintained by predator-specific defense

  41. More Story in Fish Laterality…. • Another Tanganyikan fish has lateral asymmetry (Mboko et al. 1998: Zool. Sci. 15) • A fresh water goby has lateral asymmetry in a Japanese river(Seki et al. 2000: Zool. Sci. 17) • Many fishes and other aquatic invertebrates have lateral antisymmetry! (Hori unpublished) • In these fishes, lefty is dominant heritage. • Far too counterintuitive! • We need more evidence and theoretical reason...

  42. Coexistence of laterality dimorphism (antisymmetry) Scale eaters in Lake Tanganyika (Hori unpublished) Frequencies of lefties Year of birth

  43. Righty predators eat lefty prey, and vice versa. • Lefties of scale-eating fish feed only on left side scales of lefties, righties feed only on right side scales of righties (Hori 1993 stomach contents, unpublished lab experiment). • Circa 75% of the stomach contents of righty and lefty piscivorous predators (Lamprologus spp.) were the lefty and righty, respectively (Hori unpublished field data).

  44. Why does a lefty catch a righty?(Michio Hori’s idea)

  45. Definition of Antisymmetric Predation • Both prey and predator have anti-symmetric traits (laterality); • “Lefty” predators feed on “righty” prey; “Righty” predators feed on “lefty” prey.

  46. Two-platoon lineups in MLB No fluctuation is reported in the frequency of lefty pitchers and batters in MLB or College baseball

  47. Question… %lefties • Does it really fluctuate? • Statistically significant (Hori unpubl) • Does it really synchronize? • If so, what mechanism promote fluctuation?

  48. Omnivory is common in Lake Tanganyika Fish Community Piscivores Scale eaters Hori 1997 Algal eaters

  49. We must apply our model to the entire community (Hori unpublished)

  50. x y z Extension to Holt and Polis (1997) • Where k = K/2

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