<!DOCTYPE html PUBLIC "-//W3C//DTD XHTML 1.0 Transitional//EN" "http://www.w3.org/TR/xhtml1/DTD/xhtml1-transitional.dtd"> <html> <head> <title>UTas ePrints - Mortality and predation in ecosystem models: is it important how these are expressed?</title> <script type="text/javascript" src="http://eprints.utas.edu.au/javascript/auto.js"><!-- padder --></script> <style type="text/css" media="screen">@import url(http://eprints.utas.edu.au/style/auto.css);</style> <style type="text/css" media="print">@import url(http://eprints.utas.edu.au/style/print.css);</style> <link rel="icon" href="/images/eprints/favicon.ico" type="image/x-icon" /> <link rel="shortcut icon" href="/images/eprints/favicon.ico" type="image/x-icon" /> <link rel="Top" href="http://eprints.utas.edu.au/" /> <link rel="Search" href="http://eprints.utas.edu.au/cgi/search" /> <meta content="Fulton, Elizabeth A." name="eprints.creators_name" /> <meta content="Smith, Anthony D.M." name="eprints.creators_name" /> <meta content="Johnson, Craig R." name="eprints.creators_name" /> <meta content="beth.fulton@csiro.au" name="eprints.creators_id" /> <meta name="eprints.creators_id" /> <meta content="Craig.Johnson@utas.edu.au" name="eprints.creators_id" /> <meta content="article" name="eprints.type" /> <meta content="2007-05-31" name="eprints.datestamp" /> <meta content="2008-02-04T05:27:15Z" name="eprints.lastmod" /> <meta content="show" name="eprints.metadata_visibility" /> <meta content="Mortality and predation in ecosystem models: is it important how these are expressed?" name="eprints.title" /> <meta content="pub" name="eprints.ispublished" /> <meta content="270702" name="eprints.subjects" /> <meta content="restricted" name="eprints.full_text_status" /> <meta content="Functional response; Mortality; Model closure; Biogeochemical; Ecosystem; Model" name="eprints.keywords" /> <meta content="Definitive version available online at http://www.elsevier.com/locate/issn/03043800" name="eprints.note" /> <meta content="The effects of the form of the grazing and mortality terms used in plankton models are well known. The same cannot be said for ecosystem models. As ecosystem models become more popular more needs to be known about the effects of model formulation on model behaviour and performance. The impact of the form of the grazing response function and mortality terms used in a biogeochemical ecosystem model are considered here. We show that in the large and inter-linked webs used in ecosystem models, model behaviour is far more sensitive to the form of the grazing term than to that of the mortality terms that close the modelled food web. When using biogeochemical ecosystem models in shallow marine ecosystems, the most dynamic and sophisticated functional responses describing grazing require more parameters and validation than the simpler Holling disk equation, but usually still lead to the same general conclusions about the system state and the effects of changes in forcing functions. Thus, the use of more complex functional responses is not necessarily warranted in many cases. Similarly, the extra effort and data required to explicitly represent the top predators (sharks, mammals and birds) is not necessary if they are not the focus of the study. A quadratic mortality term applied to intermediate predators (such as piscivores) is sufficient to achieve plausible model behaviour. It should be noted, however, that some degree of sophistication is required in the grazing and mortality terms. Use of simple linear functional responses and mortality terms is unsuitable for models used to consider a range of nutrient loading or harvesting scenarios." name="eprints.abstract" /> <meta content="2003-11" name="eprints.date" /> <meta content="published" name="eprints.date_type" /> <meta content="Ecological Modelling" name="eprints.publication" /> <meta content="169" name="eprints.volume" /> <meta content="1" name="eprints.number" /> <meta content="157-178" name="eprints.pagerange" /> <meta content="10.1016/S0304-3800(03)00268-0" name="eprints.id_number" /> <meta content="UNSPECIFIED" name="eprints.thesis_type" /> <meta content="TRUE" name="eprints.refereed" /> <meta content="0304-3800" name="eprints.issn" /> <meta content="http://dx.doi.org/10.1016/S0304-3800(03)00268-0" name="eprints.official_url" /> <meta content="Abrams, P.A., 1993. Why predation rate should not be proportional to predator density. Ecology 74, 726-733. Abrams, P.A., 1994. The fallacies of ''ratio-dependent'' predation. Ecology 75, 1842-1850. Abrams, P.A., Ginzburg, L.R., 2000. The nature of predation: prey dependent, ratio dependent or neither? TREE 15, 337-341. Arditi, R., Ginzburg, L.R., 1989. Coupling in predator-prey dynamics: ratio-dependence. J. Theor. Biol. 139, 311-326. Arditi, R., Saiah, H., 1992. Empirical evidence of the role of heterogeneity in ratio-dependent consumption. Ecology 73, 1544-1551. Baretta, J.W., Ebenhoh, W., Ruardij, P., 1995. The European Regional Seas Ecosystem Model, a complex marine ecosystem model. Neth. J. Sea Res. 33, 233-246. Begon, M., Mortimer, M., 1986. Population Ecology: A Unified Study of Animals and Plants. Blackwell Science Publications, London, Melbourne, Boston. Berryman, A.A., 1992. The origins and evolution of predator-prey theory. Ecology 73, 1530-1535. Brostrom, G., 1998. A note on the C/N and C/P ratio of the biological production of the Nordic seas. Tellus 50B, 93-109. 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A model for trophic interaction. Ecology 56, 881-892. Ebenhoh, W., Kohlmeier, C., Radford, P.J., 1995. The benthic biological submodel in the European Regional Seas Ecosystem Model. Neth. J. Sea Res. 33, 423-452. Edwards, A.M., Brindley, J., 1999. Zooplankton mortality and the dynamical behaviour of plankton population models. Bull. Math. Biol. 61, 303-339. Edwards, A.M., Yool, A., 2000. The role of higher predation in plankton population models. J. Plankton Res. 22, 1085-1112. Fransz, H.G., Mommaerts, J.P., Radach, G., 1991. Ecological modelling of the North Sea. Neth. J. Sea Res. 28, 67-140. Frost, B.W., 1975. A threshold feeding behaviour in Calanus pacificus. Limnol. Oceanogr. 20, 263-266. Fulton, E.A., 2001. The effects of model structure and complexity on the behaviour and performance of marine ecosystem models. Ph.D. thesis, School of Zoology, University of Tasmania, Hobart, Tasmania. Fulton, E.A., Parslow, J.S., Smith, A.D.M., Johnson, C.R., in press. Biogeochemical Marine Ecosystem Models. II. The effect of physiological detail on model performance. Ecol. Model. Fulton, E.A., Smith, A.D.M., Johnson, C.R., in press. Biogeochemical Marine Ecosystem Models. I. A model of Marine Bay Ecosystems. Ecol. Model. Fulton, E.A., Smith, A.D.M., Johnson, C.R., in press. Effects of spatial resolution on the performance and interpretation of marine ecosystem models. Ecol. Model. Gao, H., Wei, H., Sun, W., Zhai, X., 2000. Functions used in biological models and their influences on simulations. Ind. J. Mar. Sci. 29, 230-237. Gecek, S., Legovic, T., 2001. Nutrients and grazing in modeling the deep chlorophyll maximum. Ecol. Model. 138, 143-152. Ginzburg, L.R., Akcakaya, H.R., 1992. Consequences of ratiodependent predation for steady-state properties of ecosystems. Ecology 73, 1536-1543. Gleeson, S.K., 1994. Density dependence is better than ratio dependence. Ecology 75, 1834-1835. Gutierrez, A.P., 1992. Physiological basis of ratio-dependent predator-prey theory: the metabolic pool model as a paradigm. Ecology 73, 1552-1563. Hairstone, N.G., Smith, F.E., Slobodkin, L.B., 1960. Community structure, population control and competition. Am. Nat. 94, 421-425. Hassell, M.P., 1978. The Dynamics of Arthropod Predator-Prey Systems. Monographs in Population Biology, vol. 13. Princeton University Press, Princeton. Hassell, M.P., Commins, H.N., 1978. Sigmoid functional responses and population stability. Theor. Popul. Biol. 14, 62-67. Holling, C.S., 1959. Some characteristics of simple types of predation and parasitism. Can. Entomol. 91, 385-398. Holling, C.S., 1966. The functional response of invertebrate predators to prey density. Mem. Entomol. Soc. Can. 48, 1-86. Hollowed, A.B., Bax, N., Beamish, R., Collie, J., Fogarty, M., Livingston, P., Pope, J., Rice, J.C., 2000. Are multispecies models an improvement on single-species models for measuring fishing impacts on marine ecosystems? ICES J. Mar. Res. 57, 707-719. Jorgensen, S.E., 1994. Fundamentals of Ecological Modelling, 2nd ed. Developments in Environmental Modelling 19. Elsevier, Amsterdam, London, New York. Legaspi, B.C., Carruthers, R.I., Morales-Ramos, J.A., 1996. Functional response as a component of dynamic simulation models in biological control: the Catolaccus-boll weevil system. Ecol. Model. 89, 43-57. Legovic, T., 1989. Predation in food webs. Ecol. Model. 48, 267- 276. Lima, I.D., Olson, D.B., Doney, S.C., 2002. Intrinsic dynamics and stability properties of size-structured pelagic ecosystem models. J. Plankton Res. 24, 533-556. Luckyanov, N.K., Cropper Jr., W.P., Harwell, M.A., 1995. State analysis of ecological models: model reactions to parameter change. Ecol. Model. 82, 99-104. Matson, P., Berryman, A.A., 1992. Special feature: ratio-dependent predator-prey theory. Ecology 73, 1529-1566. May, R.M., 1976. Theoretical Ecology.WB Saunders, Philadelphia. McGillicuddy, D.J., McCarthy, J.J., Robinson, A.R., 1995. Coupled physical and biological modelling of the spring bloom in the North Atlantic (I): model formulation and one dimensional bloom processes. Deep Sea Res. 42, 1313-1357. McNair, J.N., 1986. The effects of refuges on predator-prey interactions: a reconsideration. Theor. Popul. Biol. 29, 38-63. Murray, A., Parslow, J., 1997. Port Phillip Bay Integrated Model: Final Report. Technical Report No. 44. Port Phillip Bay Environmental Study. CSIRO, Canberra, Australia. Murray, A.G., Parslow, J.S., 1999. The analysis of alternative formulations in a simple model of a coastal ecosystem. Ecol. Model. 119, 149-166. Myerscough, M.R., Darwen, M.J., Hogarth, W.L., 1996. Stability, persistence and structural stability in a classical predator-prey model. Ecol. Model. 89, 31-42. Radach, J., Moll, A., 1993. Estimation of the variability of production by simulating annual cycles of phytoplankton in the central North Sea. Prog. Oceanogr. 31, 339-419. Radford, P.J., 1996. The Modules of the European Regional Seas Ecosystem Model II, 1993-1996. Plymouth Marine Laboratory, Plymouth, UK. Sainsbury, K., Punt, A.E., Smith, A.D.M., 2000. Design of operational management strategies for achieving fishery ecosystem objectives. ICES J. Mar. Res. 57, 731-741. Sarnelle, O., 1994. Inferring process from pattern: trophic level abundances and imbedded interactions. Ecology 75, 1835-1841. Slobodkin, L.B., 1992. A summary of the special feature and comments on its theoretical context and importance. Ecology 73, 1564-1566. Steele, J.H., Henderson, E.W., 1992. The role of predation in plankton models. J. Plankton Res. 14, 157-172. Steele, J.H., Henderson, E.W., 1995. Predation control of plankton demography. ICES J. Mar. Res. 52, 565-573. Tett, P., Wilson, H., 2000. From biogeochemical to ecological models of marine microplankton. J. Mar. Syst. 25, 431-446. Varriale, M.C., Gomes, A.A., 1998. A study of a three species food chain. Ecol. Model. 110, 119-133. Walters, C., 1986. Adaptive Management of Renewable Resources. MacMillan, New York. Walters, C., Kitchell, J.F., 2001. Cultivation/depensation effects on juvenile survival and recruitment: implications for the theory of fishing. Can. J. Fish. Aquat. Sci. 58, 39-50. Walters, C., Christensen, V., Pauly, D., 1997. Structuring dynamic models of exploited ecosystems from trophic mass-balance assessments. Rev. Fish. Biol. Fish. 7, 139-172. Walters, C., Pauly, D., Christensen, V., Kitchell, J.F., 2000. Representing density dependent consequences of life history strategies in an ecosystem model: ECOSIM II. Ecosystems 3, 70-83. Wiedenmann, R.N., Smith Jr., J.W., 1993. Functional response of the parasite Cotesia flavipes (Hymenoptera: Braconidae) at low densities of the host Diatraea saccharalis (Lepidoptera: Pyralidae). Environ. Entomol. 22, 849-858." name="eprints.referencetext" /> <meta content="Fulton, Elizabeth A. and Smith, Anthony D.M. and Johnson, Craig R. (2003) Mortality and predation in ecosystem models: is it important how these are expressed? Ecological Modelling, 169 (1). pp. 157-178. ISSN 0304-3800" name="eprints.citation" /> <meta content="http://eprints.utas.edu.au/1121/1/2003_Fulton%2C_Smith_%26_Johnson_Ecol_Mod.pdf" name="eprints.document_url" /> <link rel="schema.DC" href="http://purl.org/DC/elements/1.0/" /> <meta content="Mortality and predation in ecosystem models: is it important how these are expressed?" name="DC.title" /> <meta content="Fulton, Elizabeth A." name="DC.creator" /> <meta content="Smith, Anthony D.M." name="DC.creator" /> <meta content="Johnson, Craig R." name="DC.creator" /> <meta content="270702 Marine and Estuarine Ecology (incl. Marine Ichthyology)" name="DC.subject" /> <meta content="The effects of the form of the grazing and mortality terms used in plankton models are well known. The same cannot be said for ecosystem models. As ecosystem models become more popular more needs to be known about the effects of model formulation on model behaviour and performance. The impact of the form of the grazing response function and mortality terms used in a biogeochemical ecosystem model are considered here. We show that in the large and inter-linked webs used in ecosystem models, model behaviour is far more sensitive to the form of the grazing term than to that of the mortality terms that close the modelled food web. When using biogeochemical ecosystem models in shallow marine ecosystems, the most dynamic and sophisticated functional responses describing grazing require more parameters and validation than the simpler Holling disk equation, but usually still lead to the same general conclusions about the system state and the effects of changes in forcing functions. Thus, the use of more complex functional responses is not necessarily warranted in many cases. Similarly, the extra effort and data required to explicitly represent the top predators (sharks, mammals and birds) is not necessary if they are not the focus of the study. A quadratic mortality term applied to intermediate predators (such as piscivores) is sufficient to achieve plausible model behaviour. It should be noted, however, that some degree of sophistication is required in the grazing and mortality terms. Use of simple linear functional responses and mortality terms is unsuitable for models used to consider a range of nutrient loading or harvesting scenarios." name="DC.description" /> <meta content="2003-11" name="DC.date" /> <meta content="Article" name="DC.type" /> <meta content="PeerReviewed" name="DC.type" /> <meta content="application/pdf" name="DC.format" /> <meta content="http://eprints.utas.edu.au/1121/1/2003_Fulton%2C_Smith_%26_Johnson_Ecol_Mod.pdf" name="DC.identifier" /> <meta content="http://dx.doi.org/10.1016/S0304-3800(03)00268-0" name="DC.relation" /> <meta content="Fulton, Elizabeth A. and Smith, Anthony D.M. and Johnson, Craig R. (2003) Mortality and predation in ecosystem models: is it important how these are expressed? Ecological Modelling, 169 (1). pp. 157-178. 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border: solid 1px #ccc; padding: 3px"><tr> <td align="left"><a href="http://eprints.utas.edu.au/cgi/users/home">Login</a> | <a href="http://eprints.utas.edu.au/cgi/register">Create Account</a></td> <td align="right" style="white-space: nowrap"> <form method="get" accept-charset="utf-8" action="http://eprints.utas.edu.au/cgi/search" style="display:inline"> <input class="ep_tm_searchbarbox" size="20" type="text" name="q" /> <input class="ep_tm_searchbarbutton" value="Search" type="submit" name="_action_search" /> <input type="hidden" name="_order" value="bytitle" /> <input type="hidden" name="basic_srchtype" value="ALL" /> <input type="hidden" name="_satisfyall" value="ALL" /> </form> </td> </tr></table></td></tr> <tr> <td class="toplinks"><!-- InstanceBeginEditable name="content" --> <div align="center"> <table width="720" class="ep_tm_main"><tr><td align="left"> <h1 class="ep_tm_pagetitle">Mortality and predation in ecosystem models: is it important how these are expressed?</h1> <p style="margin-bottom: 1em" class="not_ep_block"><span class="person_name">Fulton, Elizabeth A.</span> and <span class="person_name">Smith, Anthony D.M.</span> and <span class="person_name">Johnson, Craig R.</span> (2003) <xhtml:em>Mortality and predation in ecosystem models: is it important how these are expressed?</xhtml:em> Ecological Modelling, 169 (1). pp. 157-178. ISSN 0304-3800</p><p style="margin-bottom: 1em" class="not_ep_block"></p><table style="margin-bottom: 1em" class="not_ep_block"><tr><td valign="top" style="text-align:center"><a href="http://eprints.utas.edu.au/1121/1/2003_Fulton%2C_Smith_%26_Johnson_Ecol_Mod.pdf"><img alt="[img]" src="http://eprints.utas.edu.au/style/images/fileicons/application_pdf.png" border="0" class="ep_doc_icon" /></a></td><td valign="top"><a href="http://eprints.utas.edu.au/1121/1/2003_Fulton%2C_Smith_%26_Johnson_Ecol_Mod.pdf"><span class="ep_document_citation">PDF</span></a> - Full text restricted - Requires a PDF viewer<br />383Kb</td><td><form method="get" accept-charset="utf-8" action="http://eprints.utas.edu.au/cgi/request_doc"><input value="1403" name="docid" accept-charset="utf-8" type="hidden" /><div class=""><input value="Request a copy" name="_action_null" class="ep_form_action_button" onclick="return EPJS_button_pushed( '_action_null' )" type="submit" /> </div></form></td></tr></table><p style="margin-bottom: 1em" class="not_ep_block">Official URL: <a href="http://dx.doi.org/10.1016/S0304-3800(03)00268-0">http://dx.doi.org/10.1016/S0304-3800(03)00268-0</a></p><div class="not_ep_block"><h2>Abstract</h2><p style="padding-bottom: 16px; text-align: left; margin: 1em auto 0em auto">The effects of the form of the grazing and mortality terms used in plankton models are well known. The same cannot be said for ecosystem models. As ecosystem models become more popular more needs to be known about the effects of model formulation on model behaviour and performance. The impact of the form of the grazing response function and mortality terms used in a biogeochemical ecosystem model are considered here. We show that in the large and inter-linked webs used in ecosystem models, model behaviour is far more sensitive to the form of the grazing term than to that of the mortality terms that close the modelled food web. When using biogeochemical ecosystem models in shallow marine ecosystems, the most dynamic and sophisticated functional responses describing grazing require more parameters and validation than the simpler Holling disk equation, but usually still lead to the same general conclusions about the system state and the effects of changes in forcing functions. Thus, the use of more complex functional responses is not necessarily warranted in many cases. Similarly, the extra effort and data required to explicitly represent the top predators (sharks, mammals and birds) is not necessary if they are not the focus of the study. A quadratic mortality term applied to intermediate predators (such as piscivores) is sufficient to achieve plausible model behaviour. It should be noted, however, that some degree of sophistication is required in the grazing and mortality terms. Use of simple linear functional responses and mortality terms is unsuitable for models used to consider a range of nutrient loading or harvesting scenarios.</p></div><table style="margin-bottom: 1em" border="0" cellpadding="3" class="not_ep_block"><tr><th valign="top" class="ep_row">Item Type:</th><td valign="top" class="ep_row">Article</td></tr><tr><th valign="top" class="ep_row">Additional Information:</th><td valign="top" class="ep_row">Definitive version available online at http://www.elsevier.com/locate/issn/03043800</td></tr><tr><th valign="top" class="ep_row">Keywords:</th><td valign="top" class="ep_row">Functional response; Mortality; Model closure; Biogeochemical; Ecosystem; Model</td></tr><tr><th valign="top" class="ep_row">Subjects:</th><td valign="top" class="ep_row"><a href="http://eprints.utas.edu.au/view/subjects/270702.html">270000 Biological Sciences > 270700 Ecology and Evolution > 270702 Marine and Estuarine Ecology (incl. Marine Ichthyology)</a></td></tr><tr><th valign="top" class="ep_row">Collections:</th><td valign="top" class="ep_row">UNSPECIFIED</td></tr><tr><th valign="top" class="ep_row">ID Code:</th><td valign="top" class="ep_row">1121</td></tr><tr><th valign="top" class="ep_row">Deposited By:</th><td valign="top" class="ep_row"><span class="ep_name_citation"><span class="person_name">Professor Craig R. Johnson</span></span></td></tr><tr><th valign="top" class="ep_row">Deposited On:</th><td valign="top" class="ep_row">31 May 2007</td></tr><tr><th valign="top" class="ep_row">Last Modified:</th><td valign="top" class="ep_row">04 Feb 2008 16:27</td></tr><tr><th valign="top" class="ep_row">ePrint Statistics:</th><td valign="top" class="ep_row"><a target="ePrintStats" href="/es/index.php?action=show_detail_eprint;id=1121;">View statistics for this ePrint</a></td></tr></table><p align="right">Repository Staff Only: <a href="http://eprints.utas.edu.au/cgi/users/home?screen=EPrint::View&eprintid=1121">item control page</a></p> </td></tr></table> </div> <!-- InstanceEndEditable --></td> </tr> <tr> <td><!-- #BeginLibraryItem "/Library/footer_eprints.lbi" --> <table width="795" border="0" align="left" cellpadding="0" class="footer"> <tr valign="top"> <td colspan="2"><div align="center"><a href="http://www.utas.edu.au">UTAS home</a> | <a href="http://www.utas.edu.au/library/">Library home</a> | <a href="/">ePrints home</a> | <a href="/contact.html">contact</a> | <a href="/information.html">about</a> | <a href="/view/">browse</a> | <a href="/perl/search/simple">search</a> | <a href="/perl/register">register</a> | <a href="/perl/users/home">user area</a> | <a href="/help/">help</a></div><br /></td> </tr> <tr><td colspan="2"><p><img src="/images/eprints/footerline.gif" width="100%" height="4" /></p></td></tr> <tr valign="top"> <td width="68%" class="footer">Authorised by the University Librarian<br /> © University of Tasmania ABN 30 764 374 782<br /> <a href="http://www.utas.edu.au/cricos/">CRICOS Provider Code 00586B</a> | <a href="http://www.utas.edu.au/copyright/copyright_disclaimers.html">Copyright & Disclaimers</a> | <a href="http://www.utas.edu.au/accessibility/index.html">Accessibility</a> | <a href="http://eprints.utas.edu.au/feedback/">Site Feedback</a> </td> <td width="32%"><div align="right"> <p align="right" class="NoPrint"><a href="http://www.utas.edu.au/"><img src="http://www.utas.edu.au/shared/logos/unioftasstrip.gif" alt="University of Tasmania Home Page" width="260" height="16" border="0" align="right" /></a></p> <p align="right" class="NoPrint"><a href="http://www.utas.edu.au/"><br /> </a></p> </div></td> </tr> <tr valign="top"> <td><p> </p></td> <td><div align="right"><span class="NoPrint"><a href="http://www.eprints.org/software/"><img src="/images/eprintslogo.gif" alt="ePrints logo" width="77" height="29" border="0" align="bottom" /></a></span></div></td> </tr> </table> <!-- #EndLibraryItem --> <div align="center"></div></td> </tr> </table> </body> </html>