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Provided for Non-Commercial Research and Educational Use Provided for non-commercial research and educational use. Not for reproduction, distribution or commercial use. This article was originally published in the Encyclopedia of Ecology, Volumes 1-5 published by Elsevier, and the attached copy is provided by Elsevier for the author’s benefit and for the benefit of the author’s institution, for non-commercial research and educational use including without limitation use in instruction at your institution, sending it to specific colleagues who you know, and providing a copy to your institution’s administrator. All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution’s website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier’s permissions site at: http://www.elsevier.com/locate/permissionusematerial M Scotti. Development Capacity. In Sven Erik Jørgensen and Brian D. Fath (Editor-in-Chief), Ecological Indicators. Vol. [2] of Encyclopedia of Ecology, 5 vols. pp. [911-920] Oxford: Elsevier. Author's personal copy Ecological Indicators | Development Capacity 911 animals that appear on and in dung and the processes they Cross WF, Benstead JP, Frost PC, and Thomas SA (2005) Ecological stoichiometry in freshwater benthic systems: Recent progress and initiate are highly predictable, but in detail depend on the perspectives. Freshwater Biology 50: 1895–1912. habitat and climate under investigation. Specialized copro- Findlay S and Sinsabaugh R (eds.) (2000) Dissolved Organic Matter in philous fungi, similarly, exhibit a clear sequence of Aquatic Ecosystems. San Diego: Academic Press. Flindt MR, Pardal MA, Lillebø AI, Martins I, and Marques JC (1999) utilization of their habitat, spores of early stages already Nutrient cycling and plant dynamics in estuaries: A brief review. Acta being present in the dung when it is deposited by the Oecologica 20: 237–248. herbivore. Apparently, users of dung are highly specialized, Grac¸ a MAS (2001) The role of invertebrates on leaf litter decomposition in streams – A review. International Review of Hydrobiology and their community is significantly different from what 86: 383–393. usually is referred to as decomposers or detritivores. Grac¸ a MAS, Ba¨ rlocher F, and Gessner MO (2005) Methods to Study Depending on climate, dung patches may become comple- Litter Decomposition: A Practical Guide. Dordrecht: Springer. Kirkman H and Kendrick GA (1997) Ecological significance and tely decomposed in less than 2 months but may also last for commercial harvesting of drifting and beach-cast macro-algae and more than 2 years. Thus, as for most types of detritus, seagrasses in Australia: A review. Journal of Applied Phycology 9: 311–326. nutrient release is not sudden but extended over a longer Mackensen J, Bauhus J, and Webber E (2003) Decomposition rates of time interval, so that detritus provides a long-term store coarse woody debris – A review with particular emphasis on and source of nutrients at the basis of ecosystems. Australian tree species. Australian Journal of Botany 51: 27–37. Moore JC, Berlow EL, Coleman DC, et al. (2004) Detritus, trophic dynamics and biodiversity. Ecology Letters 7: 584–600. Orr M, Zimmer M, Jelinski DE, and Mews M (2005) Wrack deposition on See also: Decomposition and Mineralization; Freshwater different beach types: Spatial and temporal variation in the pattern of subsidy. Ecology 86: 1496–1507. Lakes; Freshwater Marshes; Habitat. Schaefer M (1991) Secondary production and decomposition. In: Ro¨ hrig E and Ulrich B (eds.) Ecosystems of the World – 7: Temperate Deciduous Forests, pp. 175–218. Amsterdam: Elsevier. Sinsabaugh RL, Osgood MP, and Findlay S (1994) Enzymatic models Further Reading for estimating decomposition rates of particulate detritus. Journal of the North American Benthological Society 13: 160–171. Attiwill PM and Adams MA (1993) Nutrient cycling in forests. New Swift MJ, Heal OW, and Anderson JM (1979) Decomposition in Phytologist 124: 561–582. Terrestrial Ecosystems. Berkeley: University of California Press. Coleman DC, Crossley DA, and Hendrix PF (2004) Fundamentals in Soil Wetzel RG (1995) Death, detritus, and energy flow in aquatic Ecology. Amsterdam: Elsevier. ecosystems. Freshwater Biology 33: 83–89. Development Capacity M Scotti, University of Parma, Parma, Italy ª 2008 Elsevier B.V. All rights reserved. Introduction System Overhead and Its Constitutive Terms Ecosystem Network Analysis Number of Roles and Development Capacity Total System Throughput and Average Mutual Ecological Applications Information Further Reading Ascendency and Development Capacity Introduction Ascendency is estimated as the product of average mutual information (AMI) by total system throughput The ever-increasing interest toward ecosystem status and (TST). It measures the ability of a system to prevail performance stimulates the application of tools for whole- over alternative configurations. AMI evaluates how the system assessment; a prominent collection of quantitative system is developed, defining flow articulation, and TST methods to achieve this aim consists of ecosystem net- expresses the size of an ecosystem as the sum of all work analysis (ENA). the flows. The apparatus of ENA comprises indices, derived Since growth and development are natural processes, from information theory, that quantify global attributes, they possess finite limits: the minimum value for ascen- and, in particular, ascendency (A) measures growth (size) dency is 0, while its upper boundary, defined as and development (organization of flows) of ecosystems. development capacity (C), is affected by system topology. Author's personal copy 912 Ecological Indicators | Development Capacity As a consequence, with a given number of nodes and 255 TST, the highest development capacity is assigned to a wholly connected network (with all flows of equal inten- Bacteria 5205 sity), while a balanced and completely articulated 75 1600 3275 topology (linear and closed chain) shows the minimum 2309 development capacity, corresponding, only in this case, to 11 184 8881 Detritus Plants Detritus the system ascendency. 300 200 feeders The remaining fraction preventing ascendency 860 2003 635 3109 1814 from reaching its theoretical maximum is called total 167 370 overhead (È), and it is calculated as the difference Carnivores between development capacity (C) and system ascen- dency (A). 203 The article briefly sketches on information theory Figure 1 Cone Spring ecosystem is composed of five principles and probabilistic approach applied to the compartments (plants, bacteria, detritus feeders, carnivores, and study of ecological networks, introducing an explana- detritus) and it shows: two imports (green), three exports (red), tion of development capacity and associated overhead five dissipations (blue), and eight intercompartmental exchanges (black). Flows are expressed in kcal mÀ2 yrÀ1. definitions. Then, it shows an alternative equation to define development capacity, described as a function of the 0 1 2 n n + 1 n + 2 effective number of ecosystem roles (a measure of the 0 0 Imports 00 degree to which the system is differentiated into distinct functions). 1 0 Finally, through the comparison between ascendency, 2 overhead, and development capacity, examples of appli- Respirat n Expo cations to real ecosystems are introduced, strengthening elements n + the important part ascendency plays as a whole-system Intercompartmental 3 eleme 3 rts index. exchanges ions nts Ecosystem Network Analysis n elements n 0 Network analysis depicts the ecosystem as a collection of n + 1 0 0 0 0 0 nodes (species or trophospecies) connected by arrowhead n + 2 0 0 0 0 0 arcs portraying trophic relations (exiting the prey items and entering the predator). n + 3 elements Trophic exchanges are measured as energy flows (i.e., kcal mÀ2 yrÀ1) or nutrient transfers of different currencies Figure 2 Graphical example of T matrix for a generic (i.e., carbon – mg C mÀ2 yrÀ1; nitrogen – mg N mÀ2 n-compartment ecosystem, with size ¼ (n þ 3)  (n þ 3). Imports À1 À2 À1 to system nodes are summarized in the first row, exports and day ; phosphorus – g P m yr ). dissipations are set in the last two columns, and the internal n  n Figure 1 depicts the Cone Spring ecosystem with five matrix shows transfers between system compartments. This compartments (n ¼ 5) and eighteen flows distinguished in model of T matrix is adopted for Cone Spring. two imports from outside (to plants and detritus), three exports to outside (from plants, bacteria, and detritus), 2 3 0 11184 0 0 0 635 0 0 five respirations (a ground symbol on each node), and 6 7 6 7 eight intercompartmental exchanges. 6 0 0 0 0 0 8881 300 2003 7 6 7 A graphical scheme can likewise be represented as an 6 7 6 0 0 0 75 0 1600 255 3275 7 6 7 (n þ 3)  (n þ 3) square matrix of transfers [T], with tij 6 7 6 0 0 0 0 370 200 0 1814 7 elements describing outflows from row compartment i to 6 7 T ¼ 6 7 ½1 column node j (see Figure 2). First row stands for imports 6 0 0 0 0 0 167 0 203 7 6 7 6 7 from outside to the column compartments, and the last 6 0 0 5205 2309 0 0 860 3109 7 6 7 two columns list exports and respirations from any row 6 7 compartment. The internal n  n submatrix reports inter- 4 00 0 000005 nal exchanges. 00 0 00000 Author's personal copy Ecological Indicators | Development Capacity 913 Total System Throughput and Average yielding to Mutual Information ÀÁ Xnþ2
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