<?xml version="1.0" encoding="UTF-8"?><!DOCTYPE article  PUBLIC "-//NLM//DTD Journal Publishing DTD v3.0 20080202//EN" "http://dtd.nlm.nih.gov/publishing/3.0/journalpublishing3.dtd"><article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" dtd-version="3.0" xml:lang="en" article-type="research article"><front><journal-meta><journal-id journal-id-type="publisher-id">IJMNTA</journal-id><journal-title-group><journal-title>International Journal of Modern Nonlinear Theory and Application</journal-title></journal-title-group><issn pub-type="epub">2167-9479</issn><publisher><publisher-name>Scientific Research Publishing</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.4236/ijmnta.2013.24028</article-id><article-id pub-id-type="publisher-id">IJMNTA-40275</article-id><article-categories><subj-group subj-group-type="heading"><subject>Articles</subject></subj-group><subj-group subj-group-type="Discipline-v2"><subject>Engineering</subject><subject> Physics&amp;Mathematics</subject></subj-group></article-categories><title-group><article-title>
 
 
  Dynamics of a Hyperparasitic System with Prolonged Diapause for Host*
 
</article-title></title-group><contrib-group><contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>imin</surname><given-names>Zhang</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="corresp" rid="cor1"><sup>*</sup></xref></contrib><contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Chaofeng</surname><given-names>Zhang</given-names></name><xref ref-type="aff" rid="aff2"><sup>2</sup></xref></contrib></contrib-group><aff id="aff2"><addr-line>School of International Trade and Economics, University of International Business and Economics, Beijing, China</addr-line></aff><aff id="aff1"><addr-line>Department of Mathematics and Finance-Economics, Sichuan University of Arts and Science, Dazhou, China</addr-line></aff><author-notes><corresp id="cor1">* E-mail:<email>lmzhang2000@163.com(IZ)</email>;</corresp></author-notes><pub-date pub-type="epub"><day>29</day><month>11</month><year>2013</year></pub-date><volume>02</volume><issue>04</issue><fpage>201</fpage><lpage>208</lpage><history><date date-type="received"><day>October</day>	<month>9,</month>	<year>2013</year></date><date date-type="rev-recd"><day>November</day>	<month>9,</month>	<year>2013</year>	</date><date date-type="accepted"><day>November</day>	<month>17,</month>	<year>2013</year></date></history><permissions><copyright-statement>&#169; Copyright  2014 by authors and Scientific Research Publishing Inc. </copyright-statement><copyright-year>2014</copyright-year><license><license-p>This work is licensed under the Creative Commons Attribution International License (CC BY). http://creativecommons.org/licenses/by/4.0/</license-p></license></permissions><abstract><p>
 
 
   A hyperparasitic system with prolonged diapause for host is investigated. It is assumed that host prolonged diapause occur at larval stage, and parasitoid attack is limited to egg stage before the initiation of host diapause. Such behavior has been reported for many ichneumons. Hyperparasite only attacks the parasitoids that parasitize the hosts. Hyperparasitic system is often used in biological control. The existence and stability of nonnegative fixed points are explored. Numerical simulations are carried out to explore the global dynamics of the system, which demonstrate appropriate prolonged diapause rate and appropriate intrinsic growth rate can stabilize the system. The reasons are explained according to the ecological perspective. Furthermore, many other complexities which include quasi-periodicity, period-doubling bifurcations leading to chaos, chaotic attractor, intermittent and supertransients are observed.
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</p></abstract><kwd-group><kwd>Hyperparasitic System; Prolonged Diapause; Dynamic Complexities</kwd></kwd-group></article-meta></front><body><sec id="s1"><title>1. Introduction</title><p>Hosts and parasitoids are mostly univoltine and have no overlap between successive generations. Therefore, their interactions can be modeled by discrete differences. An early work by Beddington et al. [<xref ref-type="bibr" rid="scirp.40275-ref1">1</xref>] showed that discrete host-parasitoid models can produce a richer set of dynamic patterns than those observed in continuous-time models. More recently, many researchers [2-12] have reported that discrete host-parasitoid models can have very complex dynamics.</p><p>However, few studies have explored hyperparasitic systems using mathematical approaches and difference equations. Actually, hyperparasite can play a crucial role in the control of a host-parasitoid interaction if they are successfully established in the community. Furthermore, few works on complex dynamics in parasitic system have considered diapause. In the natural world, many insects which inhabit unpredictable environments display diapause for one year or more, which can be described as prolonged or extra-long diapause [<xref ref-type="bibr" rid="scirp.40275-ref13">13</xref>]. As observed in numerous laboratory and field experiments, diapause is induced by changing responses to temperature, photoperiod, humidity, hormonal treatment and other factors [14- 16]. Thus, incorporating hyperparasite and diapause in parasitic system is more realistic and more practical significance.</p><p>In the paper, a hyperparasitic system with prolonged diapause for host is investigated. In the system, we assume that host prolonged diapause occur at larval stage, and parasitoid attack is limited to egg stage before the initiation of host diapause. The parasitoids are physiological “regulators” [<xref ref-type="bibr" rid="scirp.40275-ref17">17</xref>]. In this case, the parasitoid can potentially attack all hosts, but do not undergo prolonged diapause itself. Such behavior has been reported for many ichneumons [<xref ref-type="bibr" rid="scirp.40275-ref18">18</xref>]. Hyperparasite only attacks the parasitoids that parasitize the hosts [<xref ref-type="bibr" rid="scirp.40275-ref19">19</xref>]. Based on the above considerations, the model can be represented by the following difference equation:</p><disp-formula id="scirp.40275-formula55563"><label>, (1)</label><graphic position="anchor" xlink:href="2-2340096\fdc94d00-5cc0-44b9-b242-cf079af1fceb.jpg"  xlink:type="simple"/></disp-formula><p>where <img src="2-2340096\d0bfd0cd-f721-4da6-ab67-44085768cf22.jpg" /> denote the densities of the host, parasitoid, hyperparasite respectively at generation<img src="2-2340096\2ce9a85e-50b7-4345-807f-5315707f0505.jpg" />. In the absence of parasitism, the host adapts to Moran-Ricker model [20,21], that is to say<img src="2-2340096\16a4c299-d165-4c47-b87a-b2771ab8378a.jpg" />, where <img src="2-2340096\cf9efb66-3585-474d-b809-fc72a2315725.jpg" /> is the intrinsic growth rate and <img src="2-2340096\3f433df1-5420-487f-b736-cc4782b1f16b.jpg" /> is the carrying capacity. The function <img src="2-2340096\3c3e4d49-69c4-4e0f-bcf0-05c450a852fa.jpg" /> given by Poisson distribution proposed by Nicholson and Bailey [<xref ref-type="bibr" rid="scirp.40275-ref22">22</xref>] stands for the probability that a host escapes parasitism，where <img src="2-2340096\e8323652-be78-4c8c-bd47-ebe45ba4e6bb.jpg" /> is the parasitoid searching efficiency. By analogy with the above, the probability that a parasitoid escapes hyperparasites is<img src="2-2340096\28c86dd3-2889-471e-b894-74af92733098.jpg" />. Accordingly, <img src="2-2340096\f7666bbb-4e07-4d5e-82cb-80319bb4cb25.jpg" />is the hyperparasite searching efficiency. The parameters <img src="2-2340096\48489763-3833-44e0-a7a5-4b7ad07cdd82.jpg" /> and <img src="2-2340096\8d8a0070-f114-41f9-bd30-6e1c19d0aba9.jpg" /> represent diapause rate and survival rate of the host, respectively. According to their biological meaning, <img src="2-2340096\6bb9c359-2fed-4a4a-bff9-fd14c3b6a0ae.jpg" />and <img src="2-2340096\f3162e95-ffd1-44e3-9d93-d8e6f9860b0b.jpg" /> are non-negative and less than 1.</p></sec><sec id="s2"><title>2. Stability Analysis</title><p>In this section, the existence and asymptotic stability analysis of the non-negative equilibrium points of system (1) are investigated. The system has four non-negative equilibrium points which are given by the following statements:</p><p>a) The equilibrium point <img src="2-2340096\d19ff839-ee94-4ee8-927d-4e0cc705603a.jpg" /> always exists.</p><p>b) The positive equilibrium point <img src="2-2340096\aa26832f-67e5-4c47-aec7-0a32add3688d.jpg" /> is given by <img src="2-2340096\c6a4fae4-dbcb-4f7c-9a89-505fcc268835.jpg" /> which exists if and only if</p><disp-formula id="scirp.40275-formula55564"><label>(2)</label><graphic position="anchor" xlink:href="2-2340096\11a1559b-1ebf-4c49-ad29-17e9d6209bf3.jpg"  xlink:type="simple"/></disp-formula><p>The positive equilibrium point <img src="2-2340096\84f8f252-1d99-432e-85de-160753c50b8a.jpg" /> is given by</p><disp-formula id="scirp.40275-formula55565"><label>(3)</label><graphic position="anchor" xlink:href="2-2340096\8b5c2e4d-5c3e-438c-b848-1059e75b2cb7.jpg"  xlink:type="simple"/></disp-formula><p>which exists if and only if</p><disp-formula id="scirp.40275-formula55566"><label>(4)</label><graphic position="anchor" xlink:href="2-2340096\51668199-d97b-4f34-a89f-0bd9e951913f.jpg"  xlink:type="simple"/></disp-formula><p>The positive equilibrium point <img src="2-2340096\284d417c-c5c8-4c6c-ac40-779e36c44d91.jpg" /> is given by</p><disp-formula id="scirp.40275-formula55567"><label>(5)</label><graphic position="anchor" xlink:href="2-2340096\274fd663-5ccb-4556-a5ae-6b1ee6cfb9ea.jpg"  xlink:type="simple"/></disp-formula><p>which exists if and only if</p><disp-formula id="scirp.40275-formula55568"><label>(6)</label><graphic position="anchor" xlink:href="2-2340096\68f8d0a1-f649-421b-a35e-2b7e263f5f83.jpg"  xlink:type="simple"/></disp-formula><p>Analysis of the stability of the system (1) close to the above equilibrium points requires that the system is fully specified in terms of densities at time <img src="2-2340096\49183824-15c1-4648-9bec-9be57dd64dcd.jpg" /> and<img src="2-2340096\041f9a6e-8ffe-4c62-a972-4767df7ed1f4.jpg" />. For this, we introduce two variables, <img src="2-2340096\4252966b-323d-4050-b70b-777fd5161ab8.jpg" />and<img src="2-2340096\2b0dc03b-6c92-4242-9747-317aea22eb09.jpg" />, corresponding to the densities of hosts and parasitoids at time <img src="2-2340096\9985ac74-6e4f-49db-a08d-3278fc659c08.jpg" /> respectively. Then the system (1) corresponds to the following form:</p><disp-formula id="scirp.40275-formula55569"><label>(7)</label><graphic position="anchor" xlink:href="2-2340096\4296cd7e-1718-4202-ad5c-39fac6450b5e.jpg"  xlink:type="simple"/></disp-formula><p>Accordingly, the four equilibrium points of the system (1) corresponds to the following forms respectively:</p><p><img src="2-2340096\ddb4f0e4-4981-4340-bb9b-f867286b1863.jpg" /></p><p>The stabilities of equilibrium points<img src="2-2340096\6aae5eb4-40d3-4f60-b15e-3b543e36c18c.jpg" />, <img src="2-2340096\ad048749-02ea-4b85-b213-6759da1f0512.jpg" />, <img src="2-2340096\9c53827d-116c-48f5-8b80-0356e7b0ff90.jpg" />and <img src="2-2340096\4ce10633-7488-498a-b8fa-6cc9f0eb156a.jpg" /> are as the same as these points <img src="2-2340096\3ac3b4f9-673d-4de5-9d6f-43496f29cc62.jpg" /> <img src="2-2340096\640564ac-6ed8-4a4d-b3e3-5a9ac3e491b2.jpg" /> <img src="2-2340096\640ccae3-a7de-48c5-a990-d87ae351b4e0.jpg" /> and <img src="2-2340096\3b53334e-fc40-44fa-ae28-1586c1bffb0a.jpg" /> respectively. Now we study the linear stability of fixed points in the system (7). The Jacobian matrix at an arbitrary <img src="2-2340096\873f40e0-3af2-42f1-b547-fa134fd680c4.jpg" /> is given by</p><p><img src="2-2340096\465e3b44-0703-4bc8-a3d0-9a13e6712d8d.jpg" />where</p><p><img src="2-2340096\8a0afd9f-2436-4db0-a61f-9bb240714757.jpg" /></p><p>The characteristic equation of <img src="2-2340096\6950dacd-9ea2-4463-bc1e-b9f61a210bd0.jpg" /> is</p><disp-formula id="scirp.40275-formula55570"><label>, (8)</label><graphic position="anchor" xlink:href="2-2340096\0cbb02f2-4aed-4ef4-ad13-1ea89e853631.jpg"  xlink:type="simple"/></disp-formula><p>where</p><p><img src="2-2340096\8fb89fef-fc3a-4e3a-8543-aa963c206cfb.jpg" /></p><p><img src="2-2340096\c49c58e1-29db-484d-9f6f-d5e749cd1a49.jpg" /></p><p>Moreover, an application of the local stability analysis of the system (7), gives the following results:</p><p>(1) Substituting the fixed point <img src="2-2340096\aa577031-5136-4d64-a709-3e99f4cb957e.jpg" /> into the Equation (8), we get</p><disp-formula id="scirp.40275-formula55571"><label>. (9)</label><graphic position="anchor" xlink:href="2-2340096\87a65668-cb01-484e-bfdd-842536db656d.jpg"  xlink:type="simple"/></disp-formula><p>The roots of the Equation (9) are <img src="2-2340096\c29c2ebd-6eb9-4743-9254-6edaa29ce80c.jpg" /></p><p><img src="2-2340096\7dfcc428-f994-4ab1-872d-50b6dcce1d25.jpg" /></p><p>The modulus of <img src="2-2340096\6faf8b20-c549-418f-99a9-17a47aec48e8.jpg" /> is less than one. Then <img src="2-2340096\6db3a173-8854-4b18-9b13-b3b45d83c081.jpg" /> is local stability if and only if</p><p><img src="2-2340096\4be2f46f-463c-4336-843a-bf1f6f6ffe35.jpg" />which yields</p><disp-formula id="scirp.40275-formula55572"><label>(10)</label><graphic position="anchor" xlink:href="2-2340096\24ce10ac-3980-4fa6-a989-3e6c6aafda40.jpg"  xlink:type="simple"/></disp-formula><p>(2) Substituting the fixed point <img src="2-2340096\952e932f-efd5-478d-9068-5097f7b0b128.jpg" /> into the equation (8), we get</p><disp-formula id="scirp.40275-formula55573"><label>, (11)</label><graphic position="anchor" xlink:href="2-2340096\99ed6fb3-e44e-41c4-928f-9f3efa136c7e.jpg"  xlink:type="simple"/></disp-formula><p>where</p><p><img src="2-2340096\28db7913-201b-407b-9420-384b2561b477.jpg" />.</p><p>Several roots of the Equation (11) are <img src="2-2340096\41369d91-3d9f-45b9-8301-cdbfc7544399.jpg" /> <img src="2-2340096\fc3e762f-ffbb-49da-8bac-21d520c4c847.jpg" />. Obviously, the modulus of <img src="2-2340096\192adc2d-e4e8-4693-ae7f-092ac58f880b.jpg" /> is less than one. The modulus of <img src="2-2340096\a9a1684d-6d53-41fd-89e2-625dc08a7242.jpg" /> is less than one if and only if</p><disp-formula id="scirp.40275-formula55574"><label>(12)</label><graphic position="anchor" xlink:href="2-2340096\99f7e8c8-dcbc-44f0-a75e-4b297139ba41.jpg"  xlink:type="simple"/></disp-formula><p>Under the conditions of (2) and (12), the stability of <img src="2-2340096\e72284ce-1ba2-4fa0-9a16-4c4a12838809.jpg" /> is identified by the equation</p><disp-formula id="scirp.40275-formula55575"><label>. (13)</label><graphic position="anchor" xlink:href="2-2340096\a4f61e22-96b5-49e5-acd6-0d3e572f7250.jpg"  xlink:type="simple"/></disp-formula><p>It follows from the well-known Schur-cohn criterion [<xref ref-type="bibr" rid="scirp.40275-ref23">23</xref>] that the modulus of all roots of the Equation (13) is less than one if and only if</p><disp-formula id="scirp.40275-formula55576"><label>. (14)</label><graphic position="anchor" xlink:href="2-2340096\e7554e67-9073-4941-9f46-63a557e94926.jpg"  xlink:type="simple"/></disp-formula><p>From the inequalities (12) and (14), we obtain the following conditions for the stability of<img src="2-2340096\9e1f1b28-0f4d-4bc8-9111-fbc56b50f150.jpg" />:</p><p>Proposition 1. The equilibrium point <img src="2-2340096\062cd05b-8be5-4e6c-885b-08fa2cc1f668.jpg" /> is locally stable if and only if the following conditions hold:</p><disp-formula id="scirp.40275-formula55577"><label>. (15)</label><graphic position="anchor" xlink:href="2-2340096\c63ff9b5-49df-44b0-93b4-f7427f528a12.jpg"  xlink:type="simple"/></disp-formula><p>Proof. From the conditions (15), we can know the inequalities (2) and (12) obviously hold. Let's study the inequalities (14).</p><p>1)<img src="2-2340096\a501aaa3-78b8-4d43-ad0e-d5227b87cb32.jpg" />.</p><p>2)<img src="2-2340096\706a44a8-f14f-4cac-b9ce-94853be3f15d.jpg" />.</p><p>When<img src="2-2340096\14d27a79-2aad-4070-a0b9-fc0d96790889.jpg" />, according to the conditions (15)we obtain<img src="2-2340096\f358af93-0c3b-4ec5-ab88-bf1124f14712.jpg" />.</p><p>When<img src="2-2340096\a2438499-12f4-49a0-80d0-5b6d75f92332.jpg" />, according to the conditions (15), we obtain</p><p><img src="2-2340096\84dbb6a1-606d-4d35-b2e6-5fbe57398e6e.jpg" />.</p><p>3)<img src="2-2340096\2cd67855-6334-45ab-9e5d-b31464c91521.jpg" />.</p><p>Obviously,</p><p><img src="2-2340096\91379ce7-9c05-47d6-b14c-0e7eb8b30a58.jpg" /></p><p>Therefore, if the conditions (15) are satisfied, the equilibrium point <img src="2-2340096\e2c128ec-4717-48f8-800f-ef1c3e7e160e.jpg" /> is locally stable.</p><p>(3) Substituting the fixed point <img src="2-2340096\da4f6504-9996-4882-9c29-f3f3a999d529.jpg" /> into the Equation (8), we get</p><disp-formula id="scirp.40275-formula55578"><label>(16)</label><graphic position="anchor" xlink:href="2-2340096\0553f96b-f6bd-4de5-90ba-c2338e56f1fd.jpg"  xlink:type="simple"/></disp-formula><p>where</p><p><img src="2-2340096\6ac3128f-2d62-48f9-80b2-2ded2f6707e5.jpg" /></p><p><img src="2-2340096\61316530-97a1-47d3-b475-33212273ab3c.jpg" /></p><p>Two roots of the Equation (16) are <img src="2-2340096\8f511439-894e-4de0-b787-b2a6efdb6f34.jpg" /> <img src="2-2340096\21a3a796-9391-4250-a3b9-2c7deefc4df3.jpg" />. Obviously, the modulus of <img src="2-2340096\5dd227b5-11a0-41f8-923f-8aa0b1f183f0.jpg" /> is less than one. The modulus of <img src="2-2340096\1010b582-b3bd-4857-9f83-5f15d5afd798.jpg" /> is less than one if and only if</p><disp-formula id="scirp.40275-formula55579"><label>. (17)</label><graphic position="anchor" xlink:href="2-2340096\a6eead43-3386-4eeb-b53a-0acdb3b7c980.jpg"  xlink:type="simple"/></disp-formula><p>Under the conditions of (4) and (17), the stability of <img src="2-2340096\38c64075-5633-4ee5-b74f-38c3fbfb673d.jpg" /> is identified by the equation</p><disp-formula id="scirp.40275-formula55580"><label>. (18)</label><graphic position="anchor" xlink:href="2-2340096\6e18ef7f-9709-431c-bcd6-4e2ec5cd5bd9.jpg"  xlink:type="simple"/></disp-formula><p>By analogy with the above, the modulus of all roots of the Equation (18) is less than one if and only if</p><disp-formula id="scirp.40275-formula55581"><label>(19)</label><graphic position="anchor" xlink:href="2-2340096\6a3a7f07-c0dc-463f-b2da-103a427d09f1.jpg"  xlink:type="simple"/></disp-formula><p>are satisfied. Based on the above analysis, we obtain the following sufficient conditions for the stability of<img src="2-2340096\124be38c-a154-4224-9114-54eae6c2e1a7.jpg" />.</p><p>Proposition 2. The equilibrium point <img src="2-2340096\071c161d-8b13-4af0-a418-b0cc6f8b41a0.jpg" /> is locally stable if the following conditions hold:</p><p><img src="2-2340096\8b4373fd-dedd-4ae6-9e78-b843b559935c.jpg" /></p><p><img src="2-2340096\fa87a8a6-eac7-475a-a163-226df5695c19.jpg" /></p><p><img src="2-2340096\313f5591-acee-40b0-a67a-e3876ff7aa88.jpg" /></p><p>Proof. We only need verify the inequalities (19). According to the condition<img src="2-2340096\0ab5aeb2-f9c8-4c1c-b38c-4cbc789e25f7.jpg" />, we obtain<img src="2-2340096\7b3e8e1c-6514-4aff-a7f4-208e42339e9e.jpg" />.</p><p>Substitute the signs of <img src="2-2340096\a84ddbe2-e793-4d58-a982-3643943b8a41.jpg" /> and<img src="2-2340096\010d5430-dd8f-4946-b1d4-4a68a17ca23b.jpg" />, we get<img src="2-2340096\f704a44c-a001-451b-8418-9c0ae796d7df.jpg" />,<img src="2-2340096\176b22ef-962a-4245-aedd-26edda31e6da.jpg" />. Substituting the values of<img src="2-2340096\1d90e0f4-3e67-41a9-a7b3-b0639d797cc1.jpg" />, <img src="2-2340096\071534ee-f93b-4c22-b659-58dda6a630ed.jpg" />for <img src="2-2340096\aa897c0a-d5b9-4db1-bdbf-06111576a06a.jpg" /> and rearranging the term we get</p><p><img src="2-2340096\b35e5608-7195-4c9e-9d86-e66f2c927fc6.jpg" /></p><p>According to the condition<img src="2-2340096\62f2f3e3-411c-4657-9e07-d4527d0ff048.jpg" />, we obtain<img src="2-2340096\98a778f1-3040-44a0-85a4-b122e62e1855.jpg" />. By analogy with<img src="2-2340096\c1c55374-e371-4a79-b5f4-2b102821cb89.jpg" />, we get</p><p><img src="2-2340096\c0201a05-9dd8-4319-a330-828c4c5b40ba.jpg" />.</p><p>According to the signs of<img src="2-2340096\2a6d1824-7fa6-4363-b8c3-fb4954efb31b.jpg" />, we obtain<img src="2-2340096\1d8a1e5b-87b8-4437-81e4-96017c87095f.jpg" />. Now, we prove the fourth inequality of (19).</p><p><img src="2-2340096\dc01c062-dd4f-4c1f-b6ec-c65db68a3ace.jpg" />.</p><p>According to the conditions <img src="2-2340096\b6cce67f-3d22-442a-b216-b2cbdc42725f.jpg" /> and<img src="2-2340096\a447d413-ac1c-4c81-9f90-98ca1ce082dd.jpg" />, we obtain<img src="2-2340096\e74400bc-c89d-487b-8436-db1135ed023a.jpg" />. That is to say<img src="2-2340096\c47fee64-2a0f-40d8-a5e3-e9b5f0bdd163.jpg" />. At the same time,<img src="2-2340096\7fbd5a2b-e926-4c72-bcbd-afba2801d452.jpg" />. The proof is complete.</p><p>(4) Substituting the fixed point <img src="2-2340096\8bd02b6a-038b-4584-997b-ff14340afc7e.jpg" /> into the equation (8), we get</p><disp-formula id="scirp.40275-formula55582"><label>, (20)</label><graphic position="anchor" xlink:href="2-2340096\06dd8b66-922d-4cfe-b7d6-36a0e6a81b37.jpg"  xlink:type="simple"/></disp-formula><p>where</p><p><img src="2-2340096\93916742-7f80-4234-aa2c-7642a2cb124a.jpg" /></p><p>Under the condition of (6), the stability of <img src="2-2340096\05e4a3be-89d0-401e-a88f-b0db3cf47a67.jpg" /> is identified by the following equation</p><disp-formula id="scirp.40275-formula55583"><label>(21)</label><graphic position="anchor" xlink:href="2-2340096\b623411f-3598-4c6e-a647-b9a5794128d1.jpg"  xlink:type="simple"/></disp-formula><p>According to Schur-cohn criterion [<xref ref-type="bibr" rid="scirp.40275-ref23">23</xref>], the modulus of all roots of the Equation (21) is less than one if and only if</p><disp-formula id="scirp.40275-formula55584"><label>(22)</label><graphic position="anchor" xlink:href="2-2340096\9af126b3-86b9-431f-9232-a4441cfeecbb.jpg"  xlink:type="simple"/></disp-formula><p>are satisfied. Based on the above analysis, we obtain the following sufficient conditions for the stability of<img src="2-2340096\922be15d-4c5f-4541-a6f4-4695b3522aba.jpg" />.</p><p>Proposition 3. Under the condition of (6), the equilibrium point <img src="2-2340096\2eb93dbf-e548-4371-bd17-89dd3b2a8915.jpg" /> is locally stable if the following conditions hold:</p><p><img src="2-2340096\f0225d50-a72e-4197-b09b-62e02b9fd6d3.jpg" /></p><p>where</p><p><img src="2-2340096\b95822ff-9618-4f95-8126-2ea4b7915f0c.jpg" /></p><p>Proof. According to the condition<img src="2-2340096\88dee28d-03ab-4a61-b811-ae1ea1476a98.jpg" />, we obtain<img src="2-2340096\927c3737-e93c-44f1-be5f-9f56c5088c26.jpg" />. Substitute the signs of<img src="2-2340096\0ffe1688-a976-46ec-b1d5-b6d65c619eb5.jpg" />, <img src="2-2340096\0b7e1184-13f0-4698-bb35-e468edf9b9f1.jpg" />and<img src="2-2340096\55940e7f-823b-4aa5-bf9b-d1d2be4a8831.jpg" />, we get<img src="2-2340096\9c759a40-77c1-4209-b0e9-05235c625086.jpg" />,<img src="2-2340096\0bcc884c-94ea-4f53-bd02-7516bb58daf0.jpg" />. Substitute the values of<img src="2-2340096\dc5e6824-6430-4728-b2cf-d9008a55af36.jpg" />, <img src="2-2340096\c034561f-a6e7-4b38-8ae9-dc9b121c3f92.jpg" />and<img src="2-2340096\2247462f-8072-4cc7-aba1-dd95902238c0.jpg" />, we get</p><p><img src="2-2340096\1cc5f1ba-d947-41ca-bd64-db88097c9da1.jpg" />By the conditions <img src="2-2340096\e1242b1a-a098-42a9-9729-202966f41ade.jpg" /> and<img src="2-2340096\0794a7ed-f055-4f74-8f46-42a93bd281ff.jpg" />, we obtain<img src="2-2340096\9f8bc6bc-37b9-4436-96d4-e2c7c5714b4e.jpg" />, <img src="2-2340096\100a29a2-4c46-49cb-8588-75f20ffd5f0c.jpg" />and<img src="2-2340096\9b0c6dfa-1aea-45f1-95a0-0812e1b44f80.jpg" />. Then, it is easy to verify<img src="2-2340096\e1856b77-8fb2-495e-a03e-3e2a7d494b8f.jpg" />. And.</p><p><img src="2-2340096\b2401987-d31c-4c73-a05c-75a84f488834.jpg" /></p><p>According to the condition<img src="2-2340096\6cac2f2a-883d-4550-8b7e-200ae4f6342b.jpg" />, we obtain<img src="2-2340096\79371c1e-d030-49a4-bd82-6e1254f411b3.jpg" />. That is to say<img src="2-2340096\a42e7659-7f30-4980-95e2-f98079ef4688.jpg" />. According to the conclusion<img src="2-2340096\0f607ecd-c375-4bb1-a44b-c1d6ba06a3bc.jpg" />, we obtain<img src="2-2340096\84425ad0-a220-4762-814c-b678c6afe1bb.jpg" />. Now, we prove the fifth inequality of (22).</p><p><img src="2-2340096\1d17e206-cc4e-4d64-8877-93b328a7c3b8.jpg" /></p><p>By the condition<img src="2-2340096\7fba51e6-d586-4c84-8b10-b1cfdb7a8def.jpg" />, we obtain</p><p><img src="2-2340096\79b5b383-b9d1-40bf-ad6a-7e5069750220.jpg" /></p><p>The proof is complete.</p></sec><sec id="s3"><title>3. Numerical Simulations</title><p>In this section, we use the bifurcation diagrams, the Maximum Lyapunov exponents, phase portraits and so on to explore the possibilities of dynamical behaviors for system (1).</p><sec id="s3_1"><title>3.1. Bifurcation Analysis</title><p>In the section, a one-dimensional bifurcation analysis is carried out to investigate the overall dynamic behavior of the system. One-dimensional bifurcation diagrams give information about the dependence of the dynamics on a certain parameter. The analysis is expected to reveal the type of attractor to which the dynamics will ultimately settle down after passing an initial transient phase and within which the trajectory will then remain forever [<xref ref-type="bibr" rid="scirp.40275-ref2">2</xref>]. The bifurcation parameters are considered in the following two cases:</p><p>1) Varying <img src="2-2340096\65b136c8-552c-4563-bd13-1d2d4fc7e52e.jpg" /> in the range<img src="2-2340096\7fa45be2-ca07-4305-835e-e0ce205ee92d.jpg" />, and keeping other parameters fixed as below:</p><p><img src="2-2340096\2603bbfa-09a2-48f3-9a78-18fb21134a3f.jpg" />, <img src="2-2340096\4f9e7986-b634-486d-8203-a33ded85ba3c.jpg" />, <img src="2-2340096\afd2f52c-ff17-4af2-8dd7-33ecc4b28eec.jpg" />, <img src="2-2340096\99959ce4-538b-42af-b5f1-cac6d266f483.jpg" />,<img src="2-2340096\91a06d55-9cc4-4467-bd68-9273b37b466b.jpg" />. (23)</p><p>2) Varying <img src="2-2340096\5c30405d-cf24-454c-81ea-529193571791.jpg" /> in the range <img src="2-2340096\534a343d-8879-4d60-853b-7d93098e27a3.jpg" />, and keeping other parameters fixed as below:</p><p><img src="2-2340096\03a21b33-096a-4270-bd6c-c38b60aaf8fc.jpg" />, <img src="2-2340096\d25a32fc-e6cd-4ae2-901f-c0c7d90ccbeb.jpg" />, <img src="2-2340096\d1348882-2d95-4c65-8ace-9beddb6a0d42.jpg" />, <img src="2-2340096\97d30c5e-3824-4765-8d2f-43c391007c5d.jpg" />,<img src="2-2340096\8e781758-97d2-46e9-8e9b-5416237d8121.jpg" />. (24)</p><p><xref ref-type="fig" rid="fig1">Figure 1</xref>(a)&quot; target=&quot;_self&quot;&gt;<xref ref-type="fig" rid="fig1">Figure 1</xref>(a) shows the bifurcation diagram in the <img src="2-2340096\02b43826-63f6-4492-af87-22a89fb83f40.jpg" /> space with the parameters given by case 1). As <img src="2-2340096\ec6a4da8-f7d0-497a-9641-6e6b38f4811a.jpg" /> increases from <img src="2-2340096\531f1d69-b368-4ec2-a809-d65b9858f67e.jpg" /> to<img src="2-2340096\b074b6b1-536d-4814-9e15-c616c4e21899.jpg" />, the system is chaotic. Subsequently the chaotic attractor abruptly disappears and a period-4 attractor appears which constitute a type of attractor crisis. In the range <img src="2-2340096\adcb7014-eb10-408a-8aec-b82792ad2c5e.jpg" /> , the system passes through a quasi-periodic band with frequency-lockings and tangent bifurcations. As <img src="2-2340096\3414aeac-0b85-4431-bcda-392111b3a66c.jpg" />further increases, a period-2 attractor appears. When <img src="2-2340096\318e0f35-ff72-47ad-9cb3-621a3fdd634f.jpg" /> increases from <img src="2-2340096\c245db9f-90f1-46b0-a784-b4d9b683b6e6.jpg" /> to<img src="2-2340096\b4a9248b-767a-47be-98a5-bc1a78d1d4f2.jpg" />, the system goes through a quasi-periodic band with frequency-locking and tangent bifurcation. As <img src="2-2340096\56643d31-070c-4b75-8e5d-119da94ee98c.jpg" /> is slightly beyond<img src="2-2340096\c5935967-f47f-4111-b4ad-b7d0b2a9c1bc.jpg" />, a stable coexistence of the system is observed. When <img src="2-2340096\055694f7-261b-4a6d-9bac-5d92c5a5914b.jpg" />increases beyond<img src="2-2340096\26b15ee3-14df-4b57-b60d-46797706d3f1.jpg" />, the system crosses a chaotic band. When <img src="2-2340096\b26cc041-4c49-4ad5-bef6-f9858e58fe51.jpg" /> is slightly increased beyond<img src="2-2340096\0bff56f9-335c-479b-b3fb-e4f2ccd95314.jpg" />, the hyperparasite population is extinct, while the parasitoid population enters another chaotic band with period windows. <xref ref-type="fig" rid="fig1">Figure 1</xref>(b)&quot; target=&quot;_self&quot;&gt;<xref ref-type="fig" rid="fig1">Figure 1</xref>(b) is the local amplifications of <xref ref-type="fig" rid="fig1">Figure 1</xref>(a)&quot; target=&quot;_self&quot;&gt;<xref ref-type="fig" rid="fig1">Figure 1</xref>(a) with<img src="2-2340096\3533d6ca-2b9a-4163-8628-6a65cd284b18.jpg" />.</p><p>The Maximum Lyapunov exponents have been proved to be the most useful dynamic diagnostic tool for chaotic systems. It is the average exponential rate of divergence or convergence of nearby orbits in phase space [<xref ref-type="bibr" rid="scirp.40275-ref24">24</xref>]. The Maximum Lyapunov exponents corresponding to <xref ref-type="fig" rid="fig1">Figure 1</xref>(a)&quot; target=&quot;_self&quot;&gt;<xref ref-type="fig" rid="fig1">Figure 1</xref>(a) are given in <xref ref-type="fig" rid="fig1">Figure 1</xref>(c)&quot; target=&quot;_self&quot;&gt;<xref ref-type="fig" rid="fig1">Figure 1</xref>(c), which are in agreement with the bifurcation diagram. When<img src="2-2340096\96e23072-ac46-4e2e-a7e4-3839e34da23a.jpg" />, the Maximum Lyapunov exponents change from positive to negative, which corresponds with the system changing from chaos to period. In the range<img src="2-2340096\29655be2-4819-4596-a56a-ab5447e3f286.jpg" />, the Lyapunov exponents fluctuate around 0 with very small</p><p>amplitude standing for quasi-periodicity, which are the same as in the range<img src="2-2340096\5a670692-4fee-4194-a25a-3ecb6a8ec457.jpg" />. As <img src="2-2340096\065e8726-bd89-4329-9ee8-9aad47321ed5.jpg" /> increases from <img src="2-2340096\aa7ff739-0888-4d45-8a85-2580362ccbb2.jpg" /> to<img src="2-2340096\2918d4a3-1cce-4af3-be21-ae6ba668e476.jpg" />, the Maximum Lyapunov exponents are negative, corresponding to a stable coexistence of the system. When <img src="2-2340096\0c772d02-8ef6-4775-8f60-a299ffd5fdd5.jpg" /> is slightly increased beyond<img src="2-2340096\481c190a-39d6-4ce7-8b32-cc233c1d70a4.jpg" />, Most of the Maximum Lyapunov exponents are positive and few are negative. So there exist period windows in the chaotic band.</p><p>As can be seen from <xref ref-type="fig" rid="fig1">Figure 1</xref>, the behaviors of the system are very complicated, including stable coexistence, chaotic bands with period windows, quasi-periodicity with frequency-locking. Furthermore, from an ecological point of view, it is apparent that appropriate prolonged diapause rate can moderate coexistence. The reason is that appropriate diapause rate helps the fraction hosts to escape parasitism, but high diapause goes against the parasitoid growth.</p><p><xref ref-type="fig" rid="fig2">Figure 2</xref>(a)&quot; target=&quot;_self&quot;&gt;<xref ref-type="fig" rid="fig2">Figure 2</xref>(a) shows the bifurcation diagram in the <img src="2-2340096\52747682-69d1-4368-9095-5379509ce37a.jpg" /> plane with the parameters given by case (Ⅱ). As the parameter <img src="2-2340096\247c612a-2cba-44a8-b7e2-aca4196b6e41.jpg" /> increases from <img src="2-2340096\b70d5fa6-755a-48fe-a189-164dc4efdc53.jpg" /> to<img src="2-2340096\1abab5f7-edc5-4d64-a76a-fcd1fc6a400d.jpg" />, a stable coexistence of the system is observed. As <img src="2-2340096\f6ea2ac0-0e58-4909-b1f9-6aeebb93393c.jpg" /> further increases, a Hopf bifurcation occurs at<img src="2-2340096\547de165-e855-48a2-ad5f-24273fed7274.jpg" />. Then the system enters quasi-periodicity, including frequency-lockings and tangent bifurcations. When<img src="2-2340096\386c6ab3-a7b2-447e-9c0c-9a2f4168a9d4.jpg" />, the quasi-periodicity attractor abruptly disappears. In the range<img src="2-2340096\de0bc375-e0d3-40e0-b359-15166b5b1ac6.jpg" />, there is a cascade of period-doubling bifurcations leading to chaos, which is</p><p>the same as in the range<img src="2-2340096\3581daa2-49a7-4659-97d1-6c0a4161cf81.jpg" />. A typical chaotic attractor is presented in <xref ref-type="fig" rid="fig2">Figure 2</xref>(b)&quot; target=&quot;_self&quot;&gt;<xref ref-type="fig" rid="fig2">Figure 2</xref>(b) at<img src="2-2340096\d5c4fd3f-1ebb-4967-8099-8374e6751ff3.jpg" />. Subsequently the chaotic attractor abruptly disappears and a period-2 attractor appears. When <img src="2-2340096\48e80661-6b02-4bfe-b3f6-aacd8f096eee.jpg" /> increases from 4.145 to 4.5, the system enters a chaotic band again. <xref ref-type="fig" rid="fig2">Figure 2</xref>(c)&quot; target=&quot;_self&quot;&gt;<xref ref-type="fig" rid="fig2">Figure 2</xref>(c) is the local amplifications of <xref ref-type="fig" rid="fig2">Figure 2</xref>(a)&quot; target=&quot;_self&quot;&gt;<xref ref-type="fig" rid="fig2">Figure 2</xref>(a) with<img src="2-2340096\aac1e487-a1c4-4895-86c4-ede5185127f4.jpg" />.</p><p>From <xref ref-type="fig" rid="fig2">Figure 2</xref>, we can know that appropriate intrinsic growth rate <img src="2-2340096\d3461d90-4a16-46f8-a2e7-44542f396c51.jpg" /> can stabilize the system, but the high intrinsic growth rate may destabilize the stable dynamics into more complex dynamic. The reason is that the population would increase over carrying capacity with high intrinsic growth rate and then lose its stability.</p></sec><sec id="s3_2"><title>3.2. Intermittent Chaos and Supertransients</title><p>Intermittency as illustrated in <xref ref-type="fig" rid="fig3">Figure 3</xref>(a) is characterized by switches between apparently regular and chaotic behaviors even though all the control parameters are constant and no external noise is present [<xref ref-type="bibr" rid="scirp.40275-ref25">25</xref>]. The switching seems random although the dynamic model is deterministic, and the behavior is completely aperiodic and chaotic.</p><p><xref ref-type="fig" rid="fig3">Figure 3</xref>(b) shows an example of supertransients, which are used to denote an unusually long convergence to an attractor. These transient dynamics are considerably longer than the timescale of significant environmental perturbations [<xref ref-type="bibr" rid="scirp.40275-ref26">26</xref>], because the timescale of ecological</p><p>interest is tens or hundreds of generations while supertransients can persist thousands of generations or even longer. In <xref ref-type="fig" rid="fig3">Figure 3</xref>(b), the hyperparasite population size suddenly stabilizes into a 4-periodic attractor after about 720 generations of complicated fluctuations.</p></sec></sec><sec id="s4"><title>4. Conclusion</title><p>In this paper, we have proposed and investigated the host-parasitoid-hyperparasite system with prolonged diapause for host. The existence and stability of the nonnegative fixed points are explored. Subsequently, numerical simulations are carried out to exhibit other complex dynamics including stable coexistence, quasi-periodicity, period-doubling bifurcations, and chaotic bands with periodic windows, quasiperiodic attractor and non-unique attractor, intermittent chaos and supertransients and so on. Furthermore, these simulated results are explained according to ecological perspective. From <xref ref-type="fig" rid="fig1">Figure 1</xref>, we can know that the system coexists with<img src="2-2340096\409d5944-a0bf-4d5e-9542-a110591bbc72.jpg" />. That is to say, appropriate diapause rate is better for the stability of the system. Low diapause rate makes the host population suffer from high parasitism risk. High diapause rate goes against the parasitoids growth. These two cases destabilize the system. From <xref ref-type="fig" rid="fig2">Figure 2</xref>, we can know that the system is stable with<img src="2-2340096\d70a1110-7be1-464c-90dc-9ba33d24e7cc.jpg" />, but the high intrinsic growth rate may destabilize the stable dynamics into more complex dynamic. The host population would increase over carrying capacity with high intrinsic growth rate and then make the whole system lose its stability.</p></sec><sec id="s5"><title>REFERENCES</title></sec><sec id="s6"><title>NOTES</title></sec></body><back><ref-list><title>References</title><ref id="scirp.40275-ref1"><label>1</label><mixed-citation publication-type="other" xlink:type="simple">J. R. Beddington, C. A. Free and J. H. Lawton, “Dynamic Complexity in Predator-Prey Models Framed in Difference Equations,” Nature, Vol. 255, No. 5503, 1975, pp. 58-60. http://dx.doi.org/10.1038/255058a0</mixed-citation></ref><ref id="scirp.40275-ref2"><label>2</label><mixed-citation publication-type="other" xlink:type="simple">S. Y. Tang and L. S. 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