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On the Nature and Nomenclature of Ecology's Fourth Level

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On the Nature and Nomenclature of Ecology's Fourth Level Biol. Rev. (2008), 83, pp. 71–78. 71 doi:10.1111/j.1469-185X.2007.00032.x Levels of organization in biology: on the nature and nomenclature of ecology’s fourth level William Z. Lidicker, Jr.* Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720, USA (Received 30 May 2007; revised 15 October 2007; accepted 22 October 2007) ABSTRACT Viewing the universe as being composed of hierarchically arranged systems is widely accepted as a useful model of reality. In ecology, three levels of organization are generally recognized: organisms, populations, and communities (biocoenoses). For half a century increasing numbers of ecologists have concluded that recognition of a fourth level would facilitate increased understanding of ecological phenomena. Sometimes the word ‘‘ecosystem’’ is used for this level, but this is arguably inappropriate. Since 1986, I and others have argued that the term ‘‘landscape’’ would be a suitable term for a level of organization defined as an ecological system containing more than one community type. However, ‘‘landscape’’ and ‘‘landscape level’’ continue to be used extensively by ecologists in the popular sense of a large expanse of space. I therefore now propose that the term ‘‘ecoscape’’ be used instead for this fourth level of organization. A clearly defined fourth level for ecology would focus attention on the emergent properties of this level, and maintain the spatial and temporal scale-free nature inherent in this hierarchy of organizational levels for living entities. Key words: ecoscape, landscape, ecosystem, ecological system, spatial ecology, hierarchy theory, community ecology, emergent properties, holism, spatial and temporal scales. CONTENTS I. Introduction .......................................................................................................................................... 71 II. The systems concept ............................................................................................................................. 72 III. The ecological levels ............................................................................................................................. 73 IV. What about a fourth level? ................................................................................................................... 73 V. What about a name? ............................................................................................................................ 75 VI. Discussion and conclusions ................................................................................................................... 76 VII. Acknowledgements ................................................................................................................................ 77 VIII. References ............................................................................................................................................. 77 I. INTRODUCTION organization of life. I review the current status of the ecologically relevant portion of this hierarchy and ask if Hierarchical arrangements are increasingly being appreci- a fourth level for ecology is warranted. If so, what should it ated by ecologists as meaningful conceptual schemes, both be called? Sometimes hierarchies are nested, as in the one for heuristic purposes and to guide research questions and under scrutiny here, in that each level is composed of parts protocols (Ahl & Allen, 1996; Allen & Hoekstra, 1992; Allen representing the next lower level. However, other hierar- & Starr, 1982; Lidicker, 1988; McIntosh, 1963; Odum, chies are not nested, the levels being recognized by the 1953; O’Neill, DeAngelis, Waide et al., 1986; Rowe, 1961; importance of differing constraints characteristic of each Urban, O’Neill & Shugart, 1987). In this essay, I focus on level (Ahl & Allen, 1996). Often these constraints are tied to one particular hierarchy, namely, the conventional levels of varying spatial scales and temporal frequencies that * Address for correspondence: E-mail: [email protected] Biological Reviews 83 (2008) 71–78 Ó 2007 The Author Journal compilation Ó 2007 Cambridge Philosophical Society 72 William Z. Lidicker, Jr characterize entities at each level. Examples of non-nested of organism, population, and community. This is the case hierarchies of interest to ecologists are behavioural domi- even though the need for recognizing a fourth level has been nance, trophic levels, social groupings, military rankings, felt for more than half a century. and business organizations. Recently, a provocative non- It should be emphasized that assignment of a phenome- hierarchical approach to conceptualizing emergent proper- non of interest to a level of organization is only the starting ties of ecological systems has been proposed (Ponge, 2005). point in our quest for understanding. Experience tells us, The nested levels of organization hierarchy has been however, that it is an efficacious place to begin. The history widely adopted by biologists, and in some form or other of ecology (Golley, 1993; Lidicker, 1994a, 2002a; McIntosh, appears in many introductory texts. It is sometimes called 1985) illustrates the increasing depth of understanding that the ‘‘pyramid of life’’ because, being nested, each successive has been achieved as successive levels of organization have level contains fewer and fewer instances. This widely been incorporated into disciplinary discourse (Table 1). This appreciated hierarchy, applying to all of biology, is based success alone is a signal that, imperfect as it is, the levels of on the holistic notion of systems. Each level in the hierarchy organization approach provides a useful model for material is represented by a system composed of interacting parts reality. Once this level is determined in any real situation, which are also in themselves systems (holons). As such, the next imperative is to establish the spatial and temporal a given system provides the organizational framework for dimensions of the system of interest. the functioning of the parts, as well as the context within Before addressing the nature and nomenclature of which the focal system is itself a subsystem. As Ahl & Allen a fourth level in the ecological hierarchy, I briefly review (1996) explain it: the parts determine what is possible in the relevance of general systems theory for ecology, as well principle, and the context limits which of these possibilities as the current status of the three levels of organization can in fact be realized. An important feature of this generally recognized. hierarchy, which may be viewed as a disadvantage for some purposes, is that each level is arbitrarily defined in a way that is scale independent. The real world is of course much influenced by matters of spatial and temporal scale, and this II. THE SYSTEMS CONCEPT is being increasingly appreciated (Peterson & Parker, 1998; Urban et al., 1987; Wiens, 1989). Therefore there are limits The essence of the systems concept (Bertalanffy, 1950; on the extent to which the levels of organization concept Friederichs, 1958; Koestler, 1969; Lindeman, 1942; Novikoff, can successfully model life’s real organization. On the other 1945; Odum, 1953; Smuts, 1926; Tansley, 1935) is that a hand, being liberated from complications of scale allows system consists of a three-dimensional portion of the uni- one to focus on the implications of the increasing verse over a specified time period. The boundaries of a complexity generated by successive levels in the hierarchy. system are arbitrary in principle but are most usefully The fact that ecologists widely acknowledge the utility of matched to natural discontinuities of matter and energy viewing the universe as composed of hierarchically arranged such as the planet Earth or an individual human being. In systems, implies no less than that ecology as a discipline may addition to the boundary, a system contains parts which be appropriately defined as ‘‘the study of ecologically relevant interact with each other, as well as inputs and outputs systems’’ (e.g. Evans, 1956; McIntosh, 1963; Tansley, 1935). (fluxes). Aside from these five components, the most Yet, incredibly there is no generally accepted definition of important property of a system is that as a whole it has a level of complexity above that of the three traditional ones features not possessed by its constituent components. These Table 1. Ecologically relevant levels of biological organization. Those placed in square brackets are speculative, requiring further investigation Level Biotic parts Subdiscipline Organism (multi-cellular) Organs Physiological ecology Population Two or more individuals of one kind Population ecology or species Community (biocoenosis) Populations of two or more kinds Community ecology or species ECOSCAPE Two or more community types Landscape ecology ‘‘Spatial ecology’’, ‘‘Ecosystem ecology’’ [Biome] [Ecoscapes] [ ? ] [Physiographic (topographic) region; [Biomes] [ ? ] ecoregion] [Biotic province; bioregion] [Physiographic units; ecoregions] [ ? ] [Continents, ocean basins] [Biotic provinces] [ ? ] Biosphere (ecosphere) Continental Global ecology and ocean basin biotas Biological Reviews 83 (2008) 71–78 Ó 2007 The Author Journal compilation Ó 2007 Cambridge Philosophical Society Levels of organization in ecology 73 new properties arise as a consequence of the interactions Thus it developed that ecology encompasses three levels among the parts as well as constraints that arise from the of biological organization: organism,
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