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Introduction to the special issue “ : A Network Perspective and Retrospective”

ARTICLE in ECOLOGICAL MODELLING · DECEMBER 2014 Impact Factor: 2.32 · DOI: 10.1016/j.ecolmodel.2014.10.005

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Stuart R Borrett Brian D Fath University of North Carolina at Wilmington Towson University

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Ecological Modelling

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Introduction to the special issue “Systems Ecology: A Network Perspective and Retrospective”

“Systems ecology can be broadly defined as the study of the details of systems design by which the overall behavior is pro- development, dynamics, and disruption of ” duced from separate parts and mechanisms” Odum (1994, p. ix) Van Dyne (1966, p. 10) “Systems ecology focuses on the of ecosystems and “Systems ecology is the study of whole ecosystems and includes tries to reveal them by the use of a systems approach” measurements of overall performance as well as a study of the Jørgensen (2012, p. 1)

Fig. 1. Academic tree of Bernard C. Patten. Main branches of tree indicate completed graduate students (PhD or MS) primarily advised by Dr. Patten during his career. Secondary twigs indicate graduate students subsequently mentored by Patten’s students. Leaves denote doctoral students, stars indicate masters students, and flowers identify postdoctoral associates. The tree only shows two generations, though it could continue in some places. Not shown are graduate students not finished or that did not finish their degree with Dr. Patten including Jack Waide, Anita Caudle, Susan Durham, Jill Schulze, Debbie Stinner, Joe Schubauer, Lee Graham, Jianguo Liu, nor are Dr. Patten’s postdoctoral advisees including John Walsh, Richard Dame, Tarzan Legovic, Masahiko Higashi, Margret Cochran, and Oksana Buzhdygan. Original drawing by Andrea Dingeldein, 2013. http://dx.doi.org/10.1016/j.ecolmodel.2014.10.005 0304-3800/© 2014 Published by Elsevier B.V. 2 Introduction to the special issue “Systems Ecology: A Network Perspective and Retrospective” / Ecological Modelling 293 (2014) 1–3

1. Introduction and Giampietro (this volume) juxtapose hierarchy theory and the systems ecology expressed in network to show Systems ecology emerged as a subdiscipline of ecology in their similarities as well as emphasize the distinct utility of the the late 1960s (Golley, 1993) with the application of ideas and two approaches. Finally, Fath (this volume) brings together systems tools from mathematics, computer science, engineering, operations thinking ideas from ecology, sociology, and architecture to better research, , and general systems (Shugart and O’Neil, identify what makes a sustainable. Together these papers 1979; Halfon, 1979; Van Dyne, 1966). The goals of the science are show the breadth and utility of systems thinking for ecology. generally (1) to understand ecosystem organization, function, and Modeling, simulation, and are essential ele- dynamics, and (2) to use this understanding to solve applied prob- ments of systems ecology, and this second set of papers show a lems. While mathematical modeling has a long history in ecology range of modeling approaches and applications. To start, Taub and (e.g., Gompertz, 1825; Lotka, 1925, see Kingsland, 1995 for more McLaskey (this volume) revisit the empirical approach of closed detail), systems ecology can be distinguished from the more general ecological systems and remind us that not all ecological models mathematical or by its focus on whole ecolog- are mathematical. Lin and Webster (this volume) contrast three ical systems and its use of what Meadows (2008) calls systems alternative dynamic models of stream to thinking (see also Fath this issue). determine which best explained the observed dynamics in Hugh Dr. Bernard C. Patten has been a central contributor to the devel- White Creek, NC. Burns et al. (this volume) then present a sys- opment of systems ecology since its inception. His strengths in field tems analysis technique to quantify the direct and indirect effects ecology and natural history merged with simulation, computation, of perturbations in dynamic ecosystem models. In the next pair of and systems thinking led to unique new insights, methods, and papers, Koo and colleagues (this volume) use a simulation model approaches for science. His contributions have taken many forms to forecast how the growth of red spruce growth may be including fundamental research in modeling, simulation, and sys- impacted by climate change, and then couple this growth model tems analysis of ecological systems with over 142 peer reviewed to a model to predict the suitability of articles and 68 book chapters or proceedings papers, including areas in the southern Appalachian Mountains (USA) for red spruce. many highly cited articles (Patten, 1962, 1978, 1982; Patten and The last paper in this section, Shuman et al. (this volume) review Odum, 1981). He also served as the editor for eight books, includ- methods of testing an individual based model of forest dynamics, ing the landmark four volume series Systems Analysis and Simulation and then use the methods to test the success of a model in predict- in Ecology in the early 1970s that collected and codified many early ing forest dynamics in both China and Russia. Collectively, these systems ecology contributions. Patten’s involvement with three papers illustrate a myriad of modeling approaches and applications working groups is notable: (1) the System Ecology Research Group encompassed by systems ecology. at Oak Ridge National Laboratory (with G. Van Dyne, J. Olsen, S. The third group of papers is centered on network ecology, Auerbach), (2) he chaired SCOPE/ICSU’s Scientific Advisory Com- which was a focus of the symposium. This concentration was in mittee on Wetlands and Shallow Continental Bodies, and part because of Patten and Witcamp’s (1967) early recognition (3) the Okefenokee Swamp Long Term Ecological Research site. He that “to understand ecosystems ultimately will be to understand was also instrumental in developing the curricular tools for training networks”. This led Patten and his colleagues to be early movers young systems ecologists (e.g., Patten, 1975), and taught systems in this domain. Borrett et al.’s paper (this volume) opens this col- ecology and ecological modeling courses around the world. Patten lection and documents the rapid increase in the use of network has also been an important mentor for graduate students and post- approaches in ecology and documents the breadth of types of net- doctoral associates – directly and indirectly. Fig. 1 illustrates part work models and analyses used in the field. web studies of Patten’s academic tree with main branches indicating his direct are a quintessential example of network ecology, and Buzhdygan graduate students and the leaves showing these student’s gradu- et al. (this volume) provide a current example of this work by ate students. In part because of the relevance of systems ecology comparing structural statistics of seven mountain pastoral trophic to applied problems, several of Patten’s graduate students went networks and investigating their relation to other ecosystem prop- to work outside of the academy. Thus, the influence of Patten’s erties. Then, Finn et al. (this volume) illustrate how network models systems ecology mentoring likely extends much further than the can be created from fish telemetry data and analyzed to compare traditional academic tree can show. the of the organisms and infer structure. The articles included in this special issue of Ecological Modelling The remaining four papers in the network group present were presented at a symposium, April 12–14, 2012, at the Uni- advances in the realm of ecosystem network ecology. Small et al. versity of Georgia, in honor of Dr. Patten’s career titled “Systems (this volume) used inverse linear modeling to construct network Ecology: A Network Perspective and Retrospective”. They reflect models of nitrogen cycling in the Laurentian Great Lakes and use part of the influence Dr. Patten’s work has had on the field and ecosystem network analysis (ENA) to show that 4% of the nitrogen a diversity of topics that gather under the umbrella of systems exported from Lake Erie entered into the system in Lake Supe- ecology. rior. Hines and Borrett (this volume) show that results of ENA at three levels of analysis (whole network, neighborhood, and node) produce different system insights into models of N cycling in the 2. Overview Cape Fear River Estuary. Whipple et al. (this volume) extend their previous work on comparative network environ analysis (Whipple The papers in this special issue of Ecological Modelling can be et al., 2007) to show the insights of comparing the storage ver- gathered around three main themes: systems thinking, modeling sion of network environ analysis among models of N cycling in & simulation, and network ecology. the Neuse River Estuary. The contribution by Tuominen et al. (this Systems thinking broadly applied to ecological and environ- volume) extends our understanding of the utility analysis of ENA mental science is the first main theme of this special issue. This by showing how model throughflow maps into the integral utility group starts with Patten’s (this volume) clarion call for ecologists relationships. In the final contribution, Jorgensen and Nielsen (this to “get the science right” by using a systems perspective and holis- volume) explicitly link the thermodynamic and network branches tic approach. Ulanowicz (this volume) responds to Patten’s call for of systems ecology by showing how ENA measures can be modified building a better science by highlighting the importance of cap- to account for eco- such that the analyses also account for turing the effects of the non-existent in our science. Further, Allen quality, including that embedded in biological information. Introduction to the special issue “Systems Ecology: A Network Perspective and Retrospective” / Ecological Modelling 293 (2014) 1–3 3

While we have organized the special issue around these three Patten, B.C., 1978. Systems-approach to concept of environment. Ohio J. Sci. 78, main themes, each paper contributes to all three areas to some 206–222. Patten, B.C., Odum, E.P., 1981. The cybernetic of ecosystems. Am. Nat. 118, extent. Together, these contributions illustrate the diversity of 886–895. work in systems ecology in part inspired by the career of work by Patten, B.C., 1982. Environs – relativistic elementary particles for ecology. Am. Nat. Dr. Patten. 119, 179–219. Patten, B.C., Witkamp, M., 1967. Systems analysis of 134Cesium kinetics in terrestrial microcosms. Ecology 48, 813–824. Acknowledgements Shugart, H.H., O’Neil, R.V. (Eds.), 1979. Systems Ecology. Dowden, Hutchinson & Ross, Inc., Strodusburg, PA. Van Dyne, G.M., 1966. Ecosystems, Systems Ecology, and Systems Ecologists. ORNL- We thank the symposium participants a stimulating, fun, and 3975. Oak Ridge National Laboratory, Oak Ridge, TN, pp. 1–40. lively discussion. We thank the International Society for Ecologi- Whipple, S.J., Borrett, S.R., Patten, B.C., Gattie, D.K., Schramski, J.R., Bata, S.A., cal Modelling, Elsevier, and the University of Georgia for providing 2007. Indirect effects and distributed control in ecosystems: comparative funding to support this symposium, and the staff of the Odum network environ analysis of a seven-compartment model of nitrogen flow in the Neuse River Estuary: Time series analysis. Ecol. Model. 206, 1– School of Ecology for their logistical support. In particular, we 17. would like to thank Beth Gavrilles, Terry Camp, and Brenda Mat- a,b, tox for their conference assistance. We would also like to think Stuart R. Borrett ∗ Lee Snelling for her help securing funds for the event, and the a Department of and Marine Biology, UGA Complex Carbohydrate Research Center for providing space University of North Carolina Wilmington, for the symposium. We also thank Andrea Dingeldein for drawing Wilmington, NC 28403, United States the academic tree. b Duke Network Analysis Center, Social Science Research Institute, Duke University, Durham, NC References 27708, United States

a,b Golley, F.B., 1993. A History of the Ecosystem Concept in Ecology. Yale University Brian D. Fath Press, New Haven. a Department of Biological , Towson Gompertz, B., 1825. On the nature of the function expressive of the law of human University, Towson, MD 21252, United States mortality and on a new mode of determining contingencies. Philos. Trans. b R. Soc. Lond. 115, 513–585. Advanced Systems Analysis Program, International Halfon, E. (Ed.), 1979. Theoretical Systems Ecology: Advances and Case Studies. Institute for Applied Systems Analysis, Laxenburg, Academic Press, Inc., New York. Austria Jørgensen, S.E., 2012. Introduction to Systems Ecology. CRC Press. Kingsland, S.E., 1995. Modeling Nature: Episodes in the History of Population Ecol- Stuart J. Whipple ogy, 2nd ed. University of Chicago Press, Chicago. Lotka, A.J., 1925. Element of Physical Biology. Williams & Wilkins Company, Balti- Odum School of Ecology, University of Georgia, more. Athens, GA 30602, United States Meadows, D.H., 2008. Thinking in Systems: A Primer. Chelsea Green Publishing. Odum, H.T., 1994. Ecological and General Systems: An Introduction to Systems Ecol- ogy (Revised). University Press of Colorado, Niwot, Colorado. ∗ Corresponding author at: Department of Biology Patten, B.C., 1962. in net phytoplankton of Raritan Bay. J. Mar. Res. and Marine Biology, University of North Carolina 20, 57–75. Wilmington, Wilmington, NC 28403, United States. Patten, B.C., 1975. Ecosystem as a coevolutionary unit: a theme for teaching systems Tel.: +1 650 723 1684; fax: +1 650 725 2166. ecology. In: Innis, G.S. (Ed.), New Directions in the Analysis of Ecological Systems, Part 1. Simulation Councils Proceedings Series, vol. 5(1). , pp. 1–8. E-mail address: [email protected] (S.R. Borrett)