The Regulation of Chemical Budgets Over the Course of Terrestrial Ecosystem Succession Author(S): Eville Gorham, Peter M
Total Page:16
File Type:pdf, Size:1020Kb
The Regulation of Chemical Budgets Over the Course of Terrestrial Ecosystem Succession Author(s): Eville Gorham, Peter M. Vitousek and William A. Reiners Source: Annual Review of Ecology and Systematics, Vol. 10 (1979), pp. 53-84 Published by: Annual Reviews Stable URL: http://www.jstor.org/stable/2096785 . Accessed: 14/12/2013 10:19 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Annual Reviews is collaborating with JSTOR to digitize, preserve and extend access to Annual Review of Ecology and Systematics. http://www.jstor.org This content downloaded from 66.77.17.54 on Sat, 14 Dec 2013 10:19:00 AM All use subject to JSTOR Terms and Conditions Ann. Rev. Ecol. Syst. 1979. 10:53-84 Copyright 0 1979 by Annual Reviews Inc. All rights reserved THE REGULATION *4155 OF CHEMICAL BUDGETS OVER THE COURSE OF TERRESTRIAL ECOSYSTEM SUCCESSION Eville Gorham Department of Ecology and Behavioral Biology, University of Minnesota, Minneapolis, Minnesota 55455 Peter M. Vitousek Department of Biology, Indiana University, Bloomington, Indiana 47401 William A. Reiners Department of Biological Sciences, Dartmouth College, Hanover, New Hampshire 03755 INTRODUCTION Vitousek & Reiners (158) have suggested that change in net ecosystem production is a major determinant of the balance between inputs and out- puts of elements in terrestrial ecosystems. They argued that in the course of primary succession element outputs are initially relatively high (approx- imating inputs), that they then drop to a minimum because of element accumulation in biomass and detritus when net ecosystem production is highest, and that eventually output rates rise again approximately to equal inputs in late succession when net ecosystem production approaches zero (Figure 1). In most cases of secondary succession, net ecosystem production is negative immediately following disturbance, and in such cases output rates can exceed input rates. The probable importance of change in net ecosystem production is supported in particular by observations of higher element outputs from later successional ecosystems (22, 90, 157-159) and 53 0066-4162/79/1120-0053$01.00 This content downloaded from 66.77.17.54 on Sat, 14 Dec 2013 10:19:00 AM All use subject to JSTOR Terms and Conditions 54 GORHAM,VITOUSEK & REINERS also by the recorded increases in element outputs following disturbance (17, 86, 91, 150, 170). Other processes can also affect chemical budgets systematically in the course of terrestrial ecosystem succession. These processes may lessen or obscure the importance of net ecosystem production in particularinstances, or they may cause the level of inputs and outputs, though still in balance, to be higher or lower at the end of succession than at the beginning. The objective of this paper is to identify and discuss the diverse processes that can influence inputs and outputs in the course of terrestrial succession. The processes examined are those that affect element inputs to systems, includ- ing rock and soil weathering, nitrogen fixation, particle impaction, and gas A. - M4RYa / / 0 \ NET / SECONDARY.-'> ECOSYSTEM SUCCESSION ' PRODUCT/ON0 jfB SECONDIRY SUCCESS/ON ELEMENT , 1 INPUT RATE OUTPUT I RATE PRIMARY SUCS/ONV 0 SUCCESSiONALTIME - Figure I A. Patterns of change in net ecosystem production (NEP) during primary and secondary succession of terrestrial ecosystems. The starting time for primary succession is immediately following exposure of a new site; for secondary succession it is immediately following a destructive disturbance. Note the scale break for the pulse of negative NEP following disturbance. This negative pulse may be accentuated by combustion of biomass, accelerated decomposition, removal of forest products, erosion, and other factors. The inte- grated area under a properly scaled negative pulse should approximate the integrated area under the positive NEP part of the curve. See Vitousek & Reiners (158) for further explanation. B. Patterns of output rates for a limiting element in primary and secondary succession, assuming that changes in storage are controlled principally by NEP and that input rates are constant. Note the break in scale for the pulse of high output rates following disturbance in secondary succession. The integrated area under this pulse above the line for input rate should approximate the integrated area for outputs below the line for input rate. This content downloaded from 66.77.17.54 on Sat, 14 Dec 2013 10:19:00 AM All use subject to JSTOR Terms and Conditions CHEMICALBUDGETS DURING SUCCESSION 55 absorption;those that change with alterations in hydrology, including losses of dissolved substances, erosion, and oxidation-reduction reactions; and those biological processes that directly or indirectly affect the balance be- tween inputs and outputs, including net ecosystem production, element mobilization or immobilization, cation/anion balance, the production of allelochemic substances, and changes in element utilization by the biota. Each of these processes varies systematically in the course of succession, though the magnitude of each change may be difficult to predict in the course of any particular succession. We seek in this review to establish an organizational framework that will stimulate research upon the mechanisms controlling ecosystem inputs and outputs, and thus to facilitate the development of general mechanistic mod- els for biogeochemical cycling in terrestrial ecosystems. We cannot reduce the complexity of nature to simplicity, but we can provide a clearer appreci- ation and understandingof the diverse interacting mechanisms at work over the course of terrestrialsuccession. The new insights that will lead to a fuller understanding of the chemical budgets of ecosystems can come only from further detailed examination of these mechanisms. Before we analyze in detail the processes controlling chemical budgets in ecosystems, we define precisely what types of ecosystems we are considering and explain what we mean by succession and by element inputs and outputs. Succession Succession is a venerable and central concept in ecological thought (21, 26, 27, 50), yet today it is undergoing intense reconsideration (24, 36, 38, 39, 75, 76, 167). Much of this reconsideration addresses the regularity and directionality of processes and the causal mechanisms underlying change in community structure. In this reexamination new generalizations, para- digms, and terminology are being tested as replacements for the old. We wish to examine how some ecosystem properties vary as the biota changes in composition and mass with time. Until a better term is introduced, we shall use the term "succession" to describe ecosystem change following exposure of new sites or disturbance of previously occupied sites. The Watershed-EcosystemApproach For convenience and clarity, we consider as our conceptual model a water- shed ecosystem occupying a self-contained basin not subject to appreciable groundwater input. In such a situation, the only hydrologic transfers of elements into an ecosystem are in precipitation, and the only hydrologic transfers of elements out are loss to groundwater and stream water (10). Under these conditions, hydrologic fluxes of elements can be measured quite rigorously, and the effects of any perturbations of these fluxes can be assessed (10). However, in this paper we are also concerned with other kinds This content downloaded from 66.77.17.54 on Sat, 14 Dec 2013 10:19:00 AM All use subject to JSTOR Terms and Conditions 56 GORHAM, VITOUSEK & REINERS of fluxes, and for some systems and some elements the watershed model is not particularly germane (171). Definition of Inputs By convention, chemical budgets for watersheds or catchment areas are usually recorded as inputs in bulk precipitation minus outputs in stream water (46, 72, 86, 95). Inputs by gaseous absorption and particulate impac- tion are sometimes included, but their treatment is inconsistent. In some cases they are calculated by difference and included as inputs [(98) for sulfur.] In other cases they are calculated by difference but not included in inputs [(11, 98) for nitrogen]. Gaseous outputs (e.g. of nitrogen and sulfur) may also occur and add to hydrologic outputs. Weathering is not usually included as an input because it is generally calculated by difference (85). Consequently, elements derived primarily from weathering will most often have negative budgetary balances. Although it is convenient to calculate element budgets for ecosystems in terms of hydrologic transfers alone, and there is some conceptual justifica- tion for considering weathering as an internal transformation, we believe that this approach can lead to some conceptual and practical problems. First, ions derived from weathering of primary minerals in the solum can represent inputs in the same sense as ions derived from the atmosphere. Weathering of primary minerals is not a form of recycling-by definition (100) there is no chance for elements to return to primary forms within the system. Second, regarding weathering as an input can aid in understanding the processes