Abstract
In this paper, we revisit a predator–prey model with specialist and generalist predators proposed by Hanski et al. (J Anim Ecol 60:353–367, 1991) , where the density of generalist predators is assumed to be a constant. It is shown that the model admits a nilpotent cusp of codimension 4 or a nilpotent focus of codimension 3 for different parameter values. As the parameters vary, the model can undergo cusp type (or focus type) degenerate Bogdanov–Takens bifurcations of codimension 4 (or 3). Our results indicate that generalist predation can induce more complex dynamical behaviors and bifurcation phenomena, such as three small-amplitude limit cycles enclosing one equilibrium, one or two large-amplitude limit cycles enclosing one or three equilibria, three limit cycles appearing in a Hopf bifurcation of codimension 3 and dying in a homoclinic bifurcation of codimension 3. In addition, we show that generalist predation stabilizes the limit cycle driven by specialist predators to a stable equilibrium, which clearly explains the famous Fennoscandia phenomenon.
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References
Cai L, Chen G, Xiao D (2013) Multiparametric bifurcations of an epidemiological model with strong Allee effect. J Math Biol 67:185–215
Chen X, Zhang W (2009) Decomposition of algebraic sets and applications to weak centers of cubic systems. J Comput Appl Math 232:565–581
Chow SN, Li C, Wang D (1994) Normal forms and bifurcation of planar vector fields. Cambridge University Press, Cambridge
Dumortier F, Roussarie R, Sotomayor J, Żoładek H (1991) Bifurcation of planar vector fields, nilpotent singularities and abelian integrals, vol 1480. Lecture Notes in Mathematics. Springer, Berlin
Gasull A, Kooij RE, Torregrosa J (1997) Limit cycles in the Holling–Tanner model. Publ Mat 41:149–167
Gelfand IM, Kapranov MM, Zelevinsky AV (1994) Discriminants, resultants and multidimensional determinants, Birkhäuser Boston, Inc., Boston, MA
Hanski I, Hansson L, Henttonen H (1991) Specialist predators, generalist predators, and the microtine rodent cycle. J Anim Ecol 60:353–367
Hsu SB, Huang TW (1999) Hopf bifurcation analysis for a predator-prey system of Holling and Leslie type. Taiwanese J Math 3:35–53
Lamontagne Y, Coutu C, Rousseau C (2008) Bifurcation analysis of a predator-prey system with generalised Holling type III functional response. J Dyn Differ Equ 20:535–571
Li C, Rousseau C (1989) A system with three limit cycles appearing in a Hopf bifurcation and dying in a homoclinic bifurcation: the cusp of order 4. J Differ Equ 79:132–167
Lindström T (1993) Qualitative analysis of a predator-prey system with limit cycles. J Math Biol 31:541–561
Lu M, Huang J (2021) Global analysis in Bazykin’s model with Holling II functional response and predator competition. J Differ Equ 280:99–138
Lu M, Huang J, Wang H (2023) An organizing center of codimension four in a predator–prey model with generalist predator: from tristability and quadristability to transients in a nonlinear environmental change. SIAM J Appl Dyn Syst (in press)
Sáez E, González-Olivares E (1999) Dynamics of a predator–prey model. SIAM J Appl Math 59:1867–1878
Xiang C, Huang J, Ruan S, Xiao D (2020) Bifurcation analysis in a host-generalist parasitoid model with Holling II functional response. J Differ Equ 268:4618–4662
Xiang C, Huang J, Wang H (2022) Linking bifurcation analysis of Holling–Tanner model with generalist predator to a changing environment. Stud Appl Math 149:124–163
Xiang C, Huang J, Wang H (2023) Bifurcations in Holling–Tanner model with generalist predator and prey refuge. J Differ Equ 343:495–529
Xiang C, Lu M, Huang J (2022) Degenerate Bogdanov–Takens bifurcation of codimension 4 in Holling–Tanner model with harvesting. J Differ Equ 314:370–417
Xiao D, Zhang KF (2007) Multiple bifurcations of a predator–prey system. Discrete Contin Dyn Syst Ser B 8:417–433
Yang L (1999) Recent advances on determining the number of real roots of parametric polynomials. J Symbolic Comput 28:225–242
Zhang Z, Ding T, Huang W, Dong Z (1992) Qualitative theory of differential equations. Transl Math Monogr 101, American Mathematical Society, Providence, RI
Acknowledgements
Jicai Huang’s research is partially supported by NSFC (No. 11871235 and No. 12231008). Hao Wang’s research is partially supported by NSERC (RGPIN-2020-03911 and RGPAS-2020-00090).
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Appendices
Appendix A: The proof of Theorem 1
In this Appendix, we first show that \(E_*\) is a cusp of codimension at most 4 in Theorem 1. We need to show that
where \(\Omega _{24}\) is shown in (3.21), and the other notations are shown in Theorem 1.
We denote “\(\textrm{V}(f_1,\,f_2,...,f_n)\)” as the set of common zeros of \(f_1,\,f_2,...,f_n\), “\(\textrm{res}(f_1,\,f_2,\,x)\)” as the resultant of \(f_1\) and \(f_2\) with respect to x, “\(\textrm{prem}(f_1,\,f_2,\,x)\)” as the pseudo-remainder of \(f_1\) divided by \(f_2\) with respect to x, and “\(\textrm{lcoeff}(f_1,\,x)\)” as the leading coefficient of \(f_1\) with respect to x.
Step 1. Simplify the algebraic variety \(\textrm{V}(M, M_1, M_2)\cap \Omega _{24}\).
By eliminating variables in the order \(\eta \prec \alpha \), we have
where \(r_{11}\) and \(r_{12}\) are polynomials of \((u_*, \alpha )\), \(r_{21}\) and \(r_{22}\) are 17- and 54-order polynomials of \(u_*\), and we omit their complicated expressions.
Notice that, \(\textrm{lcoeff}(M, \eta )=\alpha (\alpha +3 u_*)> 0\) and \(\textrm{lcoeff}(r_{11}, \alpha )=27 u_*^4+86 u_*^3-112 u_*^2-8 u_*-8< 0\) in \(\Omega _{24}\). Similarly, from Theorem 1 in Chen and Zhang (2009) we can get that
Step 2. Simplify the algebraic variety \(\textrm{V}(r_{11},\,r_{12},\,r_{21}r_{22})\cap \Omega _{24}\).
First, by using the Maple command \(``\textrm{realroot}''\), we know that there exist one root \(u_{*0}\in I_0\) for \(\textrm{realroot}(r_{21}, 1/10^{10})\), and four roots \(u_{*i}\in I_i\) (i=1,...,4) for \(\textrm{realroot}(r_{22}, 1/10^{10})\) in \(\Omega _{24}\), such that \(r_{21}\mid _{u_{*}=u_{*0}}=r_{22}\mid _{u_{*}=u_{*i}}=0\) (i=1,...,4), where
and
Second, by using pseudo-division we can get nine pseudo-remainders
where \(w_i\) (i=1,...,9) are (10-i)-order functions of \(\alpha \), and the coefficients are functions of \(u_*\).
Notice that, \(\textrm{lcoeff}(r_{11},\, \alpha )\ne 0\), and by using Sturm’s theorem we can get that \(\textrm{lcoeff}(w_i,\, \alpha )\ne 0\), (i=1,...,9) when \(u_{*}\in \cup _{j=0}^{4}I_j\).
Combining above analysis, we can get that in \( \cup _{j=0}^{4}I_j\), \(\textrm{V}(r_{12},\,r_{11})\ne \emptyset \) and
From (A6), we know that \(w_9\) is a linear function of \(\alpha \) which has a unique root, and from \(w_9=0\) we can get that \(\alpha =\alpha _0(u_*)\). By using Sturm’s theorem again, we can show that \(\alpha _0(u_*)\) is a well defined monotone function and has no roots in \(I_i\) (i=0,...,4). Therefore, we can obtain that
which show that when \(u_*=u_{*4}\), we have \(\alpha _0(u_*)<0\), that is \(\textrm{V}(w_9,\,r_{22})\cap \Omega _{24}=\emptyset \). So that, we only need to consider the cases \(u_*=u_{*i}\) (i=0,...,3).
Notice that, \((u_*, \alpha _0(u_*))\in \Omega _{21}\). In the following, we first consider the monotonicity of \(\alpha _1\) and \(\alpha _2\) respect to \(u_*\) where \(u_*\in (0, \frac{3-\sqrt{6}}{12} ]\). Through simple calculation, we can get that
moreover, we have \(\alpha _{11}^{2}-\alpha _{12}^{2}(48 u_*^2-24 u_*+1)<0\ (=0,\ \textrm{or}\ >0)\) if \(0<u_*<0.035607\ (u_*=0.035607,\ \textrm{or}\ 0.035607<u_*\le \frac{3-\sqrt{6}}{12})\), that is \(\textrm{sign}\frac{d\alpha _1}{du_*}=\textrm{sign}\alpha _{12}\ (\frac{\textrm{sign}\alpha _{11}+\textrm{sign}\alpha _{12}}{2},\ \textrm{or}\ \textrm{sign}\alpha _{11}) \textrm{if} 0<u_*<0.035607\ (u_*=0.035607,\ \textrm{or}\ 0.035607<u_*\le \frac{3-\sqrt{6}}{12})\). Further, we have \(\alpha _{11}>0\) when \(0.035607\le u_*\le \frac{3-\sqrt{6}}{12}\), and \(\alpha _{12}>0\) when \(0<u_*\le 0.035607\). Therefore, we can get that \(\frac{d\alpha _1}{du_*}>0\) when \(u_*\in (0, \frac{3-\sqrt{6}}{12} ]\). Similarly, we can show that \(\frac{d\alpha _2}{du_*}>0\ (=0,\ <0)\) when \(0<u_*<0.035607\ (u_*=0.035607,\ \textrm{or}\ 0.035607<u_*\le \frac{3-\sqrt{6}}{12})\).
Therefore, from the above analysis, we know that \(\alpha _1\) and \(\alpha _2\) are monotone when \(u_*\in \cup _{j=0}^{2}I_j\), and we have
Combining (A4), (A8) and (A9), when \(u_*=u_{*i}\) (i=0, 2, 4), we have \((u_{*i}, \alpha _0(u_{*i}))\notin \Omega _{21}\), that is \(\textrm{V}(w_9,\,r_{21}r_{22})\cap \Omega _{24}=\emptyset \), i.e., \(\textrm{V}(r_{11},\,r_{12},\,r_{21}r_{22})\cap \Omega _{24}=\emptyset \). When \(u_*=u_{*i}\) (i=1, 3), we have \((u_{*i}, \alpha _0(u_{*i}))\in \Omega _{21}\), that is \(\textrm{V}(w_9,\,r_{21}r_{22})\cap \Omega _{24}=\textrm{V}(w_9,\,r_{22})\cap \Omega _{24}=\{(u_*, \alpha ):\ u_*=u_{*i}\ \textrm{and}\ \alpha =\alpha _0(u_{*i}),\ i=1,\ 3\}\ne \emptyset \), then we can get that
where \(u_{*i}\) and \(\alpha _0(u_{*i})\) are shown in (A4) and (A8), respectively.
Summarizing the above analysis and from (A3) we can get that
where
Step 3. Prove that \(\textrm{V}_1=\emptyset \) in (A11).
First, we simplify the algebraic variety \(\textrm{V}(M,\,M_1)\) in (A11). By using pseudo-division again, we can get one pseudo-remainder
where \(w_{10}\) is 1-order function of \(\eta \), and the coefficients are functions of \((u_*,\,\alpha )\). Then we can get that
From (A13), we know that \(w_{10}\) is a linear function of \(\eta \) which has a unique root, and from \(w_{10}=0\) we can get that \(\eta =\eta _{00}(u_*,\,\alpha )\), where
and notice that \(\big (u_{*1},\,\alpha _0(u_{*1}),\,\eta _{00}\big (u_{*1},\,\alpha _0(u_{*1})\big )\big )\in \Omega _{24}\).
Denote \(\eta _{00}(u_*,\,\alpha )\triangleq \frac{\eta _{001}}{\eta _{002}}\), where \(\eta _{001}\) and \(\eta _{002}\) are polynomials of \((u_*,\,\alpha )\), and we know that \((u_{*1},\,\alpha _0(u_{*1}))\in I_1\times [\frac{2220321857376}{10^{14}}, \frac{2220332345838}{10^{14}}]\triangleq [u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\), where \(I_1\) is shown in (A5).
We first prove that \(\eta _{002}\ne 0\) in the region \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\) by the following 2 substeps.
Step 3.1. Prove that \(\eta _{002}\) has no critical point in the interior of the region.
The corresponding first-order partial derivatives of \(\eta _{002}\) as \(\frac{\partial \eta _{002}}{\partial u_*}\) and \(\frac{\partial \eta _{002}}{\partial \alpha }\), where we omit the detailed expressions. By using Sturm’s theorem, we can get that \(\textrm{res}\big (\frac{\partial \eta _{002}}{\partial u_*},\, \frac{\partial \eta _{002}}{\partial \alpha },\, \alpha \big )\) has no root in \([u_{00}, u_{01}]\), and we have \(\textrm{lcoeff}(\frac{\partial \eta _{002}}{\partial u_*},\,\alpha )=81{u_*}^{2}-476u_*+584 \ne 0\). Then, we can get that
in \([u_{00}, u_{01}]\), which shows that \(\eta _{002}\) has no critical point in the interior of \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\), moreover, the maximum and minimum of \(\eta _{002}\) can only be achieved on the boundary of this region.
Step 3.2. Prove that \(\eta _{002}\) is monotone and rootless at the boundary of the region.
By using Sturm’s theorem again, we can get that
have no roots in \([\alpha _{00}, \alpha _{01}]\), and
have no roots in \([u_{00}, u_{01}]\).
Hence, by calculating the values of four vertices of rectangular field \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\), we can get that \(\eta _{002}\in [-1.719412\times 10^{-16}, -1.676935\times 10^{-16}]\) in \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\), which shows that \(\eta _{002}\) is well defined in this region.
Second, by using the same techniques, we analyse the value range of \(\eta _{00}(u_*,\,\alpha )\) in \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\). Through a series of calculations, we have
where \(\eta _{003}\) and \(\eta _{004}\) are polynomials of \((u_*,\,\alpha )\), and \(\textrm{lcoeff}(\eta _{003},\,\alpha )= 729{u}^{6}-9207{u}^{5}+45790{u}^{4}-109538{u}^{3}+131768{u}^{2}-74136u+13408\ne 0\) in \([u_{00}, u_{01}]\). By using Sturm’s theorem again, we can get that \(\textrm{res}(\eta _{003},\,\eta _{004},\,\alpha )\) has no real roots in \([u_{00}, u_{01}]\).
Combining the above analysis, we have
in \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\), which shows that \(\eta _{00}(u_*,\,\alpha )\) has no critical point in the interior of \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\). Therefore, the maximum and minimum of \(\eta _{00}(u_*,\,\alpha )\) can only be achieved on the boundary of this region.
By using Sturm’s theorem, we can get that
have no roots in \([\alpha _{00}, \alpha _{01}]\), moreover, \(\frac{d(\eta _{00}(u_{00},\,\alpha ))}{d\alpha }<0\ \textrm{and}\ \frac{d(\eta _{00}(u_{01},\,\alpha ))}{d\alpha }<0\) in \([\alpha _{00}, \alpha _{01}]\). Similarly,
have no roots in \([u_{00}, u_{01}]\), moreover, \(\frac{d(\eta _{00}(u_*,\,\alpha _{00}))}{du_*}>0\ \textrm{and}\ \frac{d(\eta _{00}(u_*,\,\alpha _{01}))}{du_*}>0\) in \([u_{00}, u_{01}]\).
Hence, the minimum and maximum of \(\eta _{00}(u_*,\,\alpha )\) in \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\) are \(\eta _{00}(u_{00},\,\alpha _{01})\doteq 0.000920\) and \(\eta _{00}(u_{01},\,\alpha _{00})\doteq 0.000943\), respectively. Therefore, we have \(\eta _{00}(u_{*1},\,\alpha _0(u_{*1}))\in [0.000920, 0.000943]\). Similarly, we have \(\eta _0\doteq 0.000285\) in \([u_{00}, u_{01}]\). Therefore, we can get that \(\eta _{00}(u_{*1},\,\alpha _0(u_{*1}))>\eta _0\). That is, \(\big (u_{*1},\,\alpha _0(u_{*1}),\,\eta _{00}\big (u_{*1},\,\alpha _0(u_{*1})\big )\big )\notin \Omega _{24}\).
Summarizing the above analysis, we can obtain that
Step 4. Prove that \(\textrm{V}_3=\emptyset \) in (A11).
In the following, we use the same method as Step 3 to analyse \(\textrm{V}_3\). We first simplify the algebraic variety \(\textrm{V}(M,\,M_1)\) in (A11), from (A13) and (A14) we know that
We next prove that \(\big (u_{*3},\,\alpha _0(u_{*3}),\,\eta _{00}\big (u_{*3},\,\alpha _0(u_{*3})\big )\big )\in \Omega _{24}\). In fact, we have \((u_{*3},\,\alpha _0(u_{*3}))\in I_3\times [\frac{48549816647411}{10^{15}}, \frac{48549816647417}{10^{15}}]\triangleq [u_{10}, u_{11}]\times [\alpha _{10}, \alpha _{11}]\), where \(I_3\) is shown in (A5).
Similarly, we can also show that \(\eta _{00}(u_*,\,\alpha )\) is well defined and
in \([u_{10}, u_{11}]\times [\alpha _{10}, \alpha _{11}]\), and \(\eta _0\doteq 0.004283\) in \([u_{10}, u_{11}]\). Hence, we can get that \(\eta _{00}(u_{*3},\,\alpha _0(u_{*3}))<\eta _0\). That is, \(\big (u_{*3},\,\alpha _0(u_{*3}),\,\eta _{00}\big (u_{*3},\,\alpha _0(u_{*3})\big )\big )\in \Omega _{24}\).
Therefore, we can obtain that
then we can get that \(E_*\) is a cusp of codimension at least 4.
Second, we continue to analyze the algebraic variety \(\textrm{V}(M,\,M_2)\) in (A11), by using pseudo-division once again, we can get one pseudo-remainder,
where \(w_{11}\) is 1-order function of \(\eta \), and the coefficients are functions of \((u_*,\,\alpha )\). Then we have
and from \(w_{11}=0\) we can get a unique root \(\eta =\eta _{01}(u_*,\,\alpha )\).
Similarly, we can also show that \(\eta _{01}(u_*,\,\alpha )\) is well defined and \(\eta _{01}(u_{*3},\,\alpha _0(u_{*3}))\in [0.014132345, 0.014132346]\) in \([u_{10}, u_{11}]\times [\alpha _{10}, \alpha _{11}]\). Hence, we can get that \(\eta _{01}(u_{*3},\,\alpha _0(u_{*3}))>\eta _0\), that is, \(\big (u_{*3},\,\alpha _0(u_{*3}),\,\eta _{01}\big (u_{*3},\,\alpha _0(u_{*3})\big )\big )\notin \Omega _{24}\).
Therefore, we can obtain that
Combining the above analysis and from (A3) and (A11), we can get that
which shows that \(E_*\) is a cusp of codimension exactly 4.
It is easy to know that \(\Omega _{25}\subseteq \textrm{V}(M,\,M_1)\cap \Omega _{24}\), where \(\Omega _{25}\) is shown in (1.5), and when \((u_*,\,\alpha ,\,\eta )\in \Omega _{25}\) we have \(M=M_1=0\) and \(M_2\ne 0\), that is \(E_*\) is a cusp of codimension exactly 4.
Appendix B: The proof of \(\overline{d}_{20}<0\) and \(\overline{d}_{41}<0\) in (3.26) of Theorem 2
From (3.26), when \(\mu =0\) we have
where
and we omit the expression of \(\overline{d_{410}}\). Notice that, \((u_*,\, \alpha ,\, \eta )\in \Omega _{25}\), \(\Omega _{25}\) is shown in (1.5), that is \((u_*, \alpha )=(u_{*3},\,\alpha _0(u_{*3}))\in I_3\times [\frac{48549816647411}{10^{15}}, \frac{48549816647417}{10^{15}}]\triangleq [u_{10}, u_{11}]\times [\alpha _{10}, \alpha _{11}]\), and \(\eta =\eta _{00}(u_*,\, \alpha )\), where \(\eta _{00}(u_*,\, \alpha )\) is shown in (A15).
We first calculate \(\textrm{sign}\overline{d_{200}}\). Substitute the parameter \(\eta =\eta _{00}(u_*,\, \alpha )\) into \(\overline{d_{200}}\), we can get that \(\overline{d_{200}}\mid _{\eta =\eta _{00}(u_*,\, \alpha )}=\frac{4u_*^3\alpha (u_*+\alpha )^2\overline{d_{201}}}{\eta _{002}^2}\), where \(\overline{d_{201}}\) is a polynomial of \((u_*,\, \alpha )\), we omit the detailed expression. Therefore, we have
By using the same method in Steps 3.1–3.2 of Appendix 1, we can get that \(\overline{d_{201}}\in [4.7852072357\) \(861\times 10^{-16},\,4.7852072357899\times 10^{-16}]\) in \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\). Moreover, combining condition (3.5), we know that \(\overline{d_{20}}<0\) for small \(\mu \).
Similarly, we can get that \(\overline{d_{410}}\mid _{\eta =\eta _{00}(u_*,\, \alpha )}=\frac{-16384{u_*}^{21}\alpha \left( \alpha +u_* \right) ^{11}\overline{d_{411}}}{\eta _{002}^{12}}\), where \(\overline{d_{411}}\) is a polynomial of \((u_*,\, \alpha )\), and we omit the detailed expression. Therefore, we have
moreover, we can get that \(\overline{d_{411}}\in [2.113486566436\times 10^{-96},\,2.113486566447\times 10^{-96}]\) in \([u_{00}, u_{01}]\times [\alpha _{00}, \alpha _{01}]\), which shows that \(\overline{d_{41}}<0\) for small \(\mu \).
Appendix C: The proof of nondegeneracy condition (3.29) of Theorem 2
In order to show that the nondegeneracy condition (3.29) holds when \((u_*,\, \alpha ,\, \eta )\in \Omega _{25}\), where \(\Omega _{25}\) is shown in (1.5), we just need to show that \(\overline{f}_{1}\overline{f}_{21}\overline{f}_{22}\ne 0\). Notice that, \(\Omega _{25}\subseteq \textrm{V}(M,\,M_1,\,r_{11},\,r_{12},\) \(r_{22})\cap \Omega _{24}\), so that we first prove \(\textrm{V}(M,\,M_1,\,r_{11},\,r_{12},\,r_{22},\,\overline{f}_{1})\cap \Omega _{24}=\emptyset \).
By eliminating variables in the order \(\eta \prec \alpha \prec u_*\), we have
where \(C_0\) is a positive constant, \(r_{31}\) and \(r_{32}\) are polynomials of \((u_*, \alpha )\) and \(u_*\), respectively, and we omit their complicated expressions.
From Appendix 1Step 1, we know that \(\textrm{lcoeff}(M, \eta )> 0\) and \(\textrm{lcoeff}(r_{11}, \alpha )< 0\) in \(\Omega _{25}\). Similarly, from Theorem 1 in Chen and Zhang (2009) we can get that
which shows that \(\overline{f}_{1}\ne 0\) in \(\Omega _{25}\).
Second, we prove \(\textrm{V}(M,\,M_1,\,r_{11},\,r_{12},\,r_{22},\,\overline{f}_{21} \overline{f}_{22})\cap \Omega _{24}=\emptyset \). Similarly, by eliminating variables in the order \(\eta \prec \alpha \prec u_*\), we have
where \(r_{41}\) and \(r_{42}\) are polynomials of \((u_*, \alpha )\) and \(u_*\), respectively, and we omit their complicated expressions. From \(\textrm{V}(r_{11},\,r_{41})\ne \emptyset \), we have
By using pseudo-division, we can get that
and from (3.29) we have \(\textrm{lcoeff}(\overline{f}_{21},\,\eta )=\alpha \ne 0\), then we can get that
Moreover, from (A17), when \(u_*\in [u_{10}, u_{11}]\) we have \(\frac{u_*^2}{3}\doteq 0.00166258>\eta _{00}(u_{*3},\,\alpha _0(u_{*3}))\), which shows that
By using pseudo-division again, we can get that
where \(w_{12}\) is 1-order function of \(\eta \), and the coefficients are functions of \((u_*,\,\alpha )\). Then we can get that
and from \(w_{12}=0\) we can get a unique root \(\eta =\eta _{02}(u_*,\,\alpha )\).
By using the method in Appendix 1 Steps 3.1–3.2, we can also show that \(\eta _{02}(u_*,\,\alpha )\) is well defined and \(\eta _{02}(u_{*3},\,\alpha _0(u_{*3}))\in [0.01413234,\) 0.01413235] in \([u_{10}, u_{11}]\times [\alpha _{10}, \alpha _{11}]\). Hence, we have \(\eta _{02}(u_{*3},\,\alpha _0(u_{*3}))>\eta _0\), that is, \(\big (u_{*3},\,\alpha _0(u_{*3}),\,\eta _{02}\big (u_{*3},\,\alpha _0 (u_{*3})\big )\big )\notin \Omega _{24}\).
Therefore, we can get that
From (C1), (C3) and (C4), we can obtain that \(\overline{f}_{1}\overline{f}_{21}\overline{f}_{22}\ne 0\) when \((u_*,\, \alpha ,\, \eta )\in \Omega _{25}\), that is the nondegeneracy condition (3.29) holds.
Appendix D: The proof of \(0<\hat{b}_{11}\mid _{\lambda =0}<2\sqrt{2}\) in (3.55)
Notice that, in (3.55) we have
First, we prove \(\hat{b}_{11}\mid _{\lambda =0}>0\). Substituting \(\eta =\overline{\eta }\) in (1.7) into \(\widehat{d}_1\), we have
where
Through simple calculation, we have
and \(\hat{d}_{12}<0\), which show that \(\widehat{d}_1>0\).
Similarly, we can prove that \(\widehat{d}_2<0\). Combining the condition \(\eta =\overline{\eta }\) and \(0<u^*<\frac{1}{3}\), we have \(\hat{b}_{11}\mid _{\lambda =0}>0\).
Second, we prove that \(\hat{b}_{11}\mid _{\lambda =0}<2\sqrt{2}\), from (3.55), which is equivalent to proving that
where
and \(\hat{d}_{22}\) is also a polynomial of \((\alpha ,\,u^*)\), we omit its expression, and \(\hat{d}_1\) and \(\hat{d}_2\) are showing in (D1).
By simple calculation, we have
where
Notice that, \(\hat{d}_{21}\) is a 17-order polynomial of \(\alpha \) with coefficients are functions of \(u^*\), where the coefficients of (0–13)-order terms are positive, (15-17)-order terms are negative and 14-order term is uncertain when \(0<u^*<\frac{1}{3}\). By using the Descartes’ rule of signs, we know that \(\hat{d}_{21}\) has a unique positive real root \(\widehat{\alpha }_{1}\), and
Similarly, \(\hat{d}_{22}\) (or \(\hat{d}_{23}\)) also has a unique positive real root \(\widehat{\alpha }_{2}\) (or \(\widehat{\alpha }_{3}\)), and
Hence, by using the same method as in the analysis of \(\textrm{sign}(J)\) in proof (II) of Theorem 3, we can get that \(\widehat{\alpha }_2<\widehat{\alpha }_3<\widehat{\alpha }_1\). Therefore, we can get that \( \hat{d}_{21}+\hat{d}_{22} \sqrt{(u^*)^2 ((3 \alpha ^2+\alpha (u^*+3) u^*+3 (u^*)^3)^2-4 \alpha u^* (\alpha ^2+3 \alpha (u^*)^2+(u^*)^3))}>0, \) then we have \(72 \hat{d}_1^3 u^* (\alpha +u^*) (\eta +2 (u^*)^3-(u^*)^2)+\hat{d}_2^2 (\eta +(u^*)^2)^2<0\), that is \(\hat{b}_{11}\mid _{\lambda =0}<2\sqrt{2}\). We finish the proof.
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Zhang, Y., Huang, J. & Wang, H. Bifurcations driven by generalist and specialist predation: mathematical interpretation of Fennoscandia phenomenon. J. Math. Biol. 86, 94 (2023). https://doi.org/10.1007/s00285-023-01929-1
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DOI: https://doi.org/10.1007/s00285-023-01929-1
Keywords
- Predator–prey model
- Specialist predator
- Generalist predator
- Nilpotent cusp of codimension 4
- Nilpotent focus of codimension 3
- Degenerate Bogdanov–Takens bifurcation