In this paper, we develop and analyze a model that studies the interaction between a specialist predator (one that relies exclusively on a single prey species), a generalist predator (one that takes advantage of alternative food sources in addition to consuming the focal prey species), and their common prey in a two-trophic ecosystem featuring three timescales. We assume that the prey operates on a faster timescale, while the specialist and generalist predators operate on slow and superslow timescales respectively. Treating the predation efficiency of the generalist predator as the primary varying parameter and the proportion of its diet formed by the prey species under study as the secondary parameter, we obtain a host of rich and interesting dynamics, including relaxation oscillations, mixed-mode oscillations (MMOs), subcritical elliptic bursting patterns, torus canards, and mixed-type torus canards. By grouping the timescales into two classes and using the timescale separation between classes, we apply one-fast/two-slow and two-fast/one-slow analysis techniques to gain insights about the dynamics. Using the geometric properties and flows of the singular subsystems, in combination with bifurcation analysis and numerical continuation of the full system, we classify the oscillatory dynamics and discuss the transitions from one type of dynamics to the other. The types of oscillatory patterns observed in this model are novel in population models featuring three-timescales; some of which qualitatively resemble natural cycles in small mammals and insects. Furthermore, oscillatory dynamics displaying torus canards, mixed-type torus canards, and MMOs experiencing a delayed loss of stability near one of the invariant sheets of the self-intersecting critical manifold before getting attracted to the adjacent attracting sheet of the critical manifold have not been previously reported in three-timescale models.
| Citation: |
Figure 4. Geometric structure and stable periodic orbits $ \Gamma $ of system (6) for different values of $ \beta_2 $ and $ \alpha $ and other parameters as in (8). Shown are the critical manifold $ \mathcal{M}_1 = \Pi \cup S $, the superslow manifold $ \mathcal{Z} $ (red) and the coexistence equilibrium state (blue dot). The plane $ \Pi $ is divided into an attracting component $ \Pi^a $ and a repelling component $ \Pi^r $ joined at the transcritical curve $ TC $. The surface $ S $ can consist of one or more attracting and repelling sheets joined at the folds $ \mathcal{F}^{0} $ and $ \mathcal{F}^{\pm} $ (also see table 5). Note that $ \Gamma $ displays three distinct timescales during the course of its cycle. (A) $ \beta_2 = 0.005 $ and $ \alpha = 0.6 $. (B) $ \beta_2 = 0.0245 $ and $ \alpha = 0.8 $
Figure 6. Geometric structure of the surface $ S $ of system (6) for different values of $ \beta_2 $ with $ \alpha = 0.6 $ and other parameters as in (8). The black dotted lines divide $ S $ into distinct regions, characterized by the number of attracting and repelling branches that the surface possesses. (A) The surface $ S $ is divided into four regions for $ \beta_2 = 0.048 $. Also shown are different cross-sections of $ S $ (in red) for constant $ y $ values chosen from each region. (B) The surface $ S $ is divided into three regions for $ \beta_2 = 0.025 $
Figure 9. Two parameter bifurcation diagram of (15) in $ (z, \beta_2) $ parameter plane with $ \alpha = 1 $. The HC curves intersect with each other at the "gluing bifurcation" denoted by a star and intersect with the $ SN_f $ curve at SNSL points denoted by dots. The inset shows a qualitative representation of the region around the gluing bifuraction. $ SN_f $ - saddle-node bifurcation, $ SN_p $- saddle-node of periodics, H - Hopf bifurcation, GH - generalized Hopf, HC - small homoclinic bifurcation, BHC - big homoclinic bifurcation, SNSL - saddle-node separatrix loop bifurcation
Figure 10. One parameter bifurcation diagram of (6) with respect to $ \beta_2 $ with other parameters as in (8) and $ \alpha = 0.75 $. Filled green (open blue) circles represent maximum and minimum values of $ x $ in stable (unstable) limit cycles. H - Hopf bifurcation, PD- period-doubling bifurcation, TR - torus bifurcation
Figure 12. Time series for the $ x $-coordinate of the orbits of system (6) for $ \beta_2 = 0.005 $ and varying $ \alpha $. The other parameter values are as in (8). Note the change in profile of the MMO patterns. Top-left panel: MMOs spending prolonged time near $ \mathcal{F}^+ $ with SAOs near $ \mathcal{F}^0 $; top-right and bottom-left panels: MMOs spending prolonged time near $ \mathcal{F}^0 $ with SAOs near $ \mathcal{F}^0 $; bottom-right panel: mixed-type torus canard orbit
Figure 13. (A) Overlay of the projection of trajectory $ \Gamma $ of (6) onto the $ (z, x) $ - plane with the bifurcation diagram of the fast subsystem (15) for $ \beta_2 = 0.005 $. (A) $ \alpha = 0.425 $ (B) $ \alpha = 0.6 $ (C) $ \alpha = 0.68 $ (D) $ \alpha = 0.742 $. Shown are the fold curves $ \mathcal{F}^{\pm}, \mathcal{F}^0 $ (yellow), the z-curves $ \mathcal{Z}^{\pm}, \mathcal{Z}^{0} $ (red), folded node (grey dot), equilibrium point $ E^* $ (cyan dot), delayed Hopf points (magenta dots) and degenerate nodes (black dots) of (15). Note that the trajectory exhibits delayed Hopf bifurcation on the lower branch of $ \mathcal{Z} $ in panels (B)-(D). FN - folded node, SH - subcritical Hopf, SNP - saddle-node bifurcation of periodic orbit, HC - homoclinic bifurcation, $ \mathcal{Z} $ - the superslow curve. Dashed lines denote instability. Open blue circles represent maximum and minimum values of $ x $ in unstable limit cycles
Figure 14. (A) Mixed-mode oscillations exhibited by system (6) for parameter values $ \beta_2 = 0.01 $ and $ \alpha = 0.75 $. (B) Zoomed view of the dynamics near the lower fold $ \mathcal{F}^0 $ on the $ xz $-plane. The trajectory enters into the singular funnel on $ \mathcal{S}^{-a} $ shown by the shaded region and filters through the the folded-node singularity (FN) while passing close to the delayed-Hopf bifurcation point (DHB). The local vector field of the equilibrium $ E^* $ further influences its dynamics before it jumps to $ S^{+a} $
Figure 15. Time series for the $ x $-coordinate of the orbits of system (6) for $ \beta_2 = 0.0245 $ and varying $ \alpha $. Note the transition from MMO patterns to bursting oscillations. Top-left panel: MMOs with SAOs near $ \mathcal{F}^+ $; top-right and bottom-left panels: large-amplitude oscillations featuring two-timescales; bottom-right panel: subcritical elliptic bursting
Figure 16. (A) Overlay of the projection of the trajectory $ \Gamma $ of (6) onto the $ (z, x) $ - plane with the bifurcation diagram of the fast subsystem (15) for $ \beta_2 = 0.0245 $ and varying $ \alpha $. Note the transition from spiking to bursting patterns in the system. (A) $ \alpha = 0.4645 $ (B) $ \alpha = 0.526 $ (C) $ \alpha = 0.6 $ (D) $ \alpha = 0.8 $. $ SN_p $ - saddle-node of periodics, BHC - big homoclinic loop, remaining labels and curves are as in figure 13. Filled green (open blue) circles represent maximum and minimum values of $ x $ in stable (unstable) limit cycles
Figure 17. (A) Mixed-mode oscillations exhibited by system (6) for parameter values $ \beta_2 = 0.0245 $ and $ \alpha = 0.4645 $. (B) Zoomed view of the dynamics near the upper fold $ \mathcal{F}^+ $. The trajectory enters into the singular funnel on $ \mathcal{S}^{+a} $ shown by the shaded region and filters through the the folded-node singularity (FN) while making its way to the delayed-Hopf bifurcation point (DHB). The local vector field of the equilibrium $ E^* $ further influences its dynamics before it jumps to $ S^{-a} $
| [1] |
S. Ai and S. Sadhu, The entry-exit theorem and relaxation oscillations in slow-fast planar systems, Journal of Diff. Eq., 268 (2020), 7220-7249.
doi: 10.1016/j.jde.2019.11.067.
|
| [2] |
S. Ai and Y. Yingfei, Relaxation oscillations in predator-prey systems, Journal of Dynamics and Differential Equations, (2021), 1-28.
|
| [3] |
S. M. Baer and T. Erneux, Singular Hopf bifurcation to relaxation oscillations, SIAM J. Appl. Math., 46 (1986), 721-739.
doi: 10.1137/0146047.
|
| [4] |
E. Baspinar, D. Avitabile and M. Desroches, Canonical models for torus canards in elliptic bursters, Chaos, 31 (2021), 063129.
doi: 10.1063/5.0037204.
|
| [5] |
A. D. Bazykin, Nonlinear Dynamics of Interacting Populations, Series A: Monographs and Treatise; World Scientific Series on Nonlinear Science: Singapore, 1998.
doi: 10.1142/9789812798725.
|
| [6] |
N. Bolohan, V. LeBlanc and F. Lutscher, Seasonal dynamics of a generalist and a specialist predator on a single prey, Math. Appl. Sc. and Engr., 2 (2021), 72-148.
|
| [7] |
H. Broer, T. J. kaper and M. Krupa, Geometric desingularization of a cusp singularity in slow-fast systems with applications to Zeeman's examples, Journal of Dynamics and Differential Equations, Springer Verlag, 2013.
doi: 10.1007/s10884-013-9322-5.
|
| [8] |
M. Brøns, T. J. Kaper and H. G. Rotstein, Introduction to focus issue: Mixed mode oscillations: Experiment, computation, and analysis, Chaos, 18 (2008), 015101.
|
| [9] |
M. Brøns and R. Kaasen, Canards and mixed-mode oscillations in a forest pest model, Theoretical Population Biology, 77 (2010), 238-242.
|
| [10] |
B. M. Brøns, M. Krupa and M. Wechselberger, Mixed mode oscillations due to the generalized canard phenomenon, Fields Institute Communications, 49 (2006), 39-63.
|
| [11] |
J. Burke, M. Desroches, A. M. Barry, T. J. Kaper and M. A. Kramer, A showcase of torus canards in neuronal bursters, J. Math. Neurosci., 2 (2012), 2-30.
doi: 10.1186/2190-8567-2-3.
|
| [12] |
P. T. Cardin and M. A. Teixeira, Fenichel theory for multiple time scale singular perturbation problems, SIAM J. Appld. Dyn. Syst., 16 (2017), 1425-1452.
doi: 10.1137/16M1067202.
|
| [13] |
P. De Maesschalck, E. Kutafina and N. Popovic, Sector-delayed-Hopf-type mixed-mode oscillations in a prototypical three-time-scale model, Appl. Math. Comput., 273 (2016), 337-352.
doi: 10.1016/j.amc.2015.09.083.
|
| [14] |
B. Deng, Food chain chaos due to junction-fold point, Chaos, 11 (2001), 514-525.
doi: 10.1063/1.1576531.
|
| [15] |
B. Deng and G. Hines, Food chain chaos due to Shilnikov's orbit, Chaos, 12 (2002, ) 533-538.
doi: 10.1063/1.1482255.
|
| [16] |
B. Deng and G. Hines, Food chain chaos due to transcritical point, Chaos, 13 (2003), 578-585.
doi: 10.1063/1.1576531.
|
| [17] |
B. Deng, Food chain chaos with canard explosion, Chaos, 14 (2004), 1083-1092.
doi: 10.1063/1.1814191.
|
| [18] |
M. Desroches, J. Burke, T. J. Kaper and M. A. Kramer, Canards of mixed type in a neural burster, Phy. Review E, 85 (2012), 021920.
|
| [19] |
M. Desroches, J. Guckenheimer, B. Krauskopf, C. Kuehn, H. M. Osinga and M. Wechselberger, Mixed-mode oscillations with multiple time scales, SIAM Review, 54 (2012), 211-288.
doi: 10.1137/100791233.
|
| [20] |
L. Duan, D. Zhai and Q. Lu, Bifurcation and bursting in Morris-Lecar model for class I and class II excitability, Disc. Cont. Dynm. Syst (S), (2011), 391-399.
|
| [21] |
A. Erbach, F. Lutscher and G. Seo, Bistability and limit cycles in generalist predator-prey dynamics, Ecol. Complexity, 14 (2013), 48-55.
|
| [22] |
N. Fenichel, Geometric singular perturbation theory for ordinary differential equations, J. Diff. Eq., 31 (1979), 53-98.
doi: 10.1016/0022-0396(79)90152-9.
|
| [23] |
J. Guckenheimer, J. H. Tien and A. R. Willms, Bifurcations in the fast dynamics of neurons: Implications for bursting, in Bursting: The Genesis of Rhythm in the Nervous System, World Scientific, (2005), 89-122.
|
| [24] |
V. Grotan, R. Lande, S. Engen, B. E. Saether and P. J. DeVries, Seasonal cycles of species diversity and similarity in a tropical butterfly community, J. of Animal Ecol., 81 (2012), 714-723.
|
| [25] |
I. Hanski, L. Hansson and H. Henttonen, Specialist Predators, Generalist Predators, and the Microtine Rodent Cycle, J. Animal Ecol., 60 (1991), 353-367.
|
| [26] |
M. P. Hassell and R. M. May, Generalist and Specialist Natural Enemies in Insect Predator-Prey Interactions, J. of Animal Ecol., 55 (1986), 923-940.
|
| [27] |
E. M. Izhikevich, Neural excitability, spiking and bursting, Internat. J. Bifur. Chaos Appl. Sci. Engrg., 10 (2000), 1171-1266.
doi: 10.1142/S0218127400000840.
|
| [28] |
S. Jelbert, S.-V. Kuntz and C. Kuehn, Geometric blow-up for folded limit cycle manifolds in three time-scale systems, arXiv: 2208.01361.
|
| [29] |
P. Kaklamanos and N. Popović, Complex oscillatory dynamics in a three-timescale El Niño Southern Oscillation model, arXiv: 2207.03230.
|
| [30] |
P. Kaklamanos, N. Popović and K. U. Kristiansen, Bifurcations of mixed-mode oscillations in three-timescale systems: An extended prototypical example, Chaos, 32 (2022), 013108.
doi: 10.1063/5.0073353.
|
| [31] |
Y. Kang and L. Wedekin, Dynamics of a intraguild predation model with generalist or specialist predator, J. Math. Biol., 67 (2013), 1227-1259.
doi: 10.1007/s00285-012-0584-z.
|
| [32] |
C. Kuehn, Multiple Time Scale Dynamics, , Springer, 2015.
doi: 10.1007/978-3-319-12316-5.
|
| [33] |
M. Krupa, N. Popović and N. Kopell, Mixed-mode oscillations in three time-scale systems: A prototypical example, SIAM J, Appl. Dyn. Syst., 7 (2008), 361-420.
doi: 10.1137/070688912.
|
| [34] |
M. Kuwamura and H. Chiba, Mixed-mode oscillations and chaos in a prey-predator system with dormancy of predators, Chaos, 19 (2009), 1-10.
doi: 10.1063/1.3270262.
|
| [35] |
Y. A. Kuznetsov and S. Rinaldi, Remarks on food chain dynamics, Math. Biosci., 133 (1996), 1-33.
doi: 10.1016/0025-5564(95)00104-2.
|
| [36] |
B. Letson, J. Rubin and T. Vo, Analysis of interacting local oscillation mechanisms in three-timescale systems, SIAM J. Appl. Math., 77 (2017), 1020-1946.
doi: 10.1137/16M1088429.
|
| [37] |
W. Liu, D. Xiao and Y. Yi, Relaxation oscillations in a class of predator-prey systems, J. Diff. Equ., 188 (2003), 306-331.
doi: 10.1016/S0022-0396(02)00076-1.
|
| [38] |
F. Molleman, Moving beyond phenology: New directions in the study of temporal dynamics of tropical insect communities, Current Science, 114 (2018).
|
| [39] |
S. Muratori and S. Rinaldi, Remarks on competitive coexistence, SIAM J. Applied Math., 49 (1989), 1462-1472.
doi: 10.1137/0149088.
|
| [40] |
S. Muratori and S. Rinaldi, Low- and high-frequency oscillations in three-dimensional food chain system, J. Appl. Math., 52 (1992), 1688-1706.
doi: 10.1137/0152097.
|
| [41] |
A. I. Neishtadt, Persistence of stability loss for dynamical bifurcations I, Differ. Equ., 24 (1987), 1385-1391.
|
| [42] |
A. I. Neishtadt, Persistence of stability loss for dynamical bifurcations II, Differ. Equ., 24 (1988), 171-176.
|
| [43] |
W. A. Nelson, O. N. Bjornstad and T. Yamanaka, Recurrent insect outbreaks caused by temperature-driven changes in system stability, Science, 314 (2013), 796-799.
|
| [44] |
J. C. Poggiale, C. Aldebert, B. Girardot and B. W. Kooi, Analysis of a predator-prey model with specific time scales: a geometrical approach proving the occurrence of canard solutions, J. of Math. Bio., 80 (2020), 39-60.
doi: 10.1007/s00285-019-01337-4.
|
| [45] |
S. Rinaldi and S. Muratori, Limit cycles in slow-fast forest-pest models, Theor. Popul. Biol., 41 (1992), 26-43.
doi: 10.1016/0040-5809(92)90048-X.
|
| [46] |
S. Sadhu and S. Chakraborty Thakur, Uncertainty and predictability in population dynamics of a two-trophic ecological model: Mixed-mode oscillations, bistability and sensitivity to parameters, Ecological Complexity, 32 (2017), 196-208.
|
| [47] |
S. Sadhu, Complex oscillatory patterns near singular Hopf bifurcation in a two-timescale ecosystem, Discrete Continuous Dynamical Systems - B, 26 (2021), 5251-5279.
doi: 10.3934/dcdsb.2020342.
|
| [48] |
S. Sadhu, Analysis of the onset of a regime shift and detecting early warning signs of major population changes in a two-trophic three species predator-prey model with long-term transients, J. Math. Biol., 85 (2022), 1-33.
doi: 10.1007/s00285-022-01805-4.
|
| [49] |
G. Seo and G. Wolkowicz, Pest control by generalist parasitoids: A bifurcation theory approach, DCDS-S, 13 (2020), 3157-3187.
doi: 10.3934/dcdss.2020163.
|
| [50] |
G. R. Singleton, P. R. Brown, R. P. Pech, J. Jacob, G. J. Mutze and C. J. Krebs, One hundred years of eruptions of house mice in Australia - a natural biological curio, Biol. J. of Linnean Soc., 84 (2005), 617-627.
|
| [51] |
N. C. Stenseth, Population cycles in voles and lemmings: Density dependence and phase dependence in a stochastic world, Oikos, 87 (1999), 427-461.
|
| [52] |
R. Straube, D. Flockerzi and M. J. B. Hauser, Sub-Hopf/fold-cycle bursting and its relation to (quasi-)periodic oscillations, J. Phys.: Conf. Ser., 55 (2006).
|
| [53] |
W. Teka, J. Tabak, T. Vo, M. Wechselberger and R. Bertram, The dynamics underlying pseudo-plateau bursting in a pituitary cell model, J. Math. Neuro. Sc., 1 (2011).
doi: 10.1186/2190-8567-1-12.
|
| [54] |
R. Tyson and F. Lutscher, Seasonally varying predation behaviour and climate shifts are predicted to affect predator-prey cycles, Am. Nat., 188 (2016), 539-553.
|
| [55] |
T. Vo, R. Bertram and M. Wechselberger, Multiple geometric viewpoints of mixed mode dynamics associated with pseudo-plateau bursting, SIAM J. Appl. Dyn. Syst., 12 (2013), 789-830.
doi: 10.1137/120892842.
|
| [56] |
T. Vo, Generic torus canards, Physica D: Nonlinear Phenomena, 356 (2017), 37-64.
doi: 10.1016/j.physd.2017.06.005.
|
| [57] |
C. Wang and X. Zhang, Canards, heteroclinic and homoclinic orbits for a slow-fast predator-prey model of generalized Holling type Ⅲ, J. Diff. Eqns., 267 (2019), 3397-3441.
doi: 10.1016/j.jde.2019.04.008.
|
| [58] |
M. Wechselberger, Existence and bifurcation of canards in $\mathbb{R}^3$ in the case of a folded node, SIAM J. Appl. Dyn. Syst., 4 (2005), 101-139.
doi: 10.1137/030601995.
|
Two-parameter bifurcation diagram of (5) showing transitions between different dynamical regimes for all parameter values as in (8). See text for details. HB: Hopf bifurcation, TR: torus bifurcation
Mixed-mode time series of (6) in intermediate time for
Bursting patterns in system (6) for
Geometric structure and stable periodic orbits
Shape of the fold curve
Geometric structure of the surface
Two-parameter bifurcation structure of the desingularized system (14). The FSN Ⅰ (a) and FSN Ⅰ (b) curves represent folded saddle-node bifurcations of type Ⅰ, while FSN Ⅱ (a) and FSN Ⅱ (b) curves represent folded saddle-node bifurcations of type Ⅱ. See text for details
Real parts of eigenvalues of the layer problem (15) for
Two parameter bifurcation diagram of (15) in
One parameter bifurcation diagram of (6) with respect to
Time profiles of the
Time series for the
(A) Overlay of the projection of trajectory
(A) Mixed-mode oscillations exhibited by system (6) for parameter values
Time series for the
(A) Overlay of the projection of the trajectory
(A) Mixed-mode oscillations exhibited by system (6) for parameter values