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Origins of the Novel Ecosystems Concept Joseph Mascaro1, James A Chapter 5 Origins of The Novel Ecosystems Concept Joseph Mascaro1, James A. Harris2, Lori Lach3, Allen Thompson4, Michael P. Perring5, David M. Richardson6 and Erle C. Ellis7 1Department of Global Ecology, Carnegie Institution for Science, Stanford, California, USA 2Environmental Science and Technology Department, Cranfield University, UK 3Ecosystem Restoration and Intervention Ecology (ERIE) Research Group, School of Plant Biology and the Centre for Integrative Bee Research, University of Western Australia, Australia 4School of History, Philosophy, and Religion, Oregon State University, USA 5Ecosystem Restoration and Intervention Ecology (ERIE) Research Group, School of Plant Biology, University of Western Australia, Australia 6Centre for Invasion Biology, Department of Botany and Zoology, Stellenbosch University, South Africa 7Geography & Environmental Systems, University of Maryland, USA “In such ways anthropogenic ecosystems differ from those developed independently of man. But the essential forma- tive processes . are the same.” Arthur G. Tansley, 1935 5.1 INTRODUCTION his academic nature, variously accepting and rejecting terms that appear throughout the literature. He accepts Reading Tansley’s (1935) opus, combatively titled ‘The ‘climax’ (within which he notably allows slow gradual use and abuse of vegetational concepts and terms’ change), coins the term ‘ecosystem’ (surely his greatest nearly a century on, is illuminating in many ways. For gift to the field), then roundly rejects the idea that eco- one, the prose leaps off the page in a manner rarely seen systems are a kind of ‘complex organism’, a phrase he in ecology journals today: he moves through emotional finds too inexact to be useful. He italicizes profusely states no modern editor would allow, gleefully decon- (evidenced in the opening quotation, for which the structing the exactitude of his ‘old friend’ Fredrick emphasis is original), an effort to convey final authority; Clements’ proposition of a ‘climatic climax’, a single like Clements, Tansley exhibits ecology’s roots in botany, terminus of succession for a whole biome. He betrays where taxonomy and strict classification reigned. Novel Ecosystems: Intervening in the New Ecological World Order, First Edition. Edited by Richard J. Hobbs, Eric S. Higgs, and Carol M. Hall. © 2013 John Wiley & Sons, Ltd. Published 2013 by John Wiley & Sons, Ltd. 45 46 Novel ecosystems The lasting value of Tansley’s paper comes not in Whittaker. When ecology mourned Gleason in 1975, his concluding litany of terminology (most of which is Robert McIntosh wrote for Torrey that “Gleason’s then now altered or abandoned), but in his flashes of uncer- heretical idea is now widely recognized as part of the tainty about nature. He raises questions to which he conventional wisdom in ecology” (McIntosh 1975). does not know the answer: “Is man part of ‘nature’ or Yet controversy remained. not?” He frames the quintessential debate in early 20th Had he lived another decade, Gleason would have century ecology without making a ruling: “Many ecol- received a gift reserved for all but a few theoreticians: ogists hold that all vegetation is always changing. It irrefutable proof. In the form of sediment from North may be so: we do not know enough either to affirm or American lakes, the past was being steadily disrobed by to deny so sweeping a statement.” With statements like palynological studies (Davis 1981, 1983; Pielou 1991; these, Tansley hints at an important insight: strict defi- Delcourt 2002). These samples, which were largely nitions of ecosystem types may never be tractable from a period of glacial retreat at the close of the because there is too much spatiotemporal variation to Wisconsinan 8,000 to 14,000 years ago, painted a categorize. clear picture: in times of environmental change, tree The organizing objective of this chapter was to cat- species move as individuals and not as part of discrete egorically define a novel ecosystem. Doing so at the communities or organisms (Fig. 5.1). While students outset would imply that we were already convinced of from Madison to Cornell can rattle off tree and shrub their existence as something separate from other eco- species that ‘characterize’ the beech-maple forest asso- systems, a notion upon which our read of Tansley has ciation today, students visiting the same habitat in a cast doubt. We therefore begin not by declaring what previous interglacial would find mesic forests of quite novel ecosystems are or are not (a task which we shall different composition. Such patterns inspired one pale- come to presently) but by reviewing the foundational oecologist contributing to this volume to call himself a principles that point to the existence and importance ‘radical Gleasonian’ (Jackson 2006). From the tropics, of novel ecosystems. From these, and a brief review of Janzen (1985) concluded that the seduction of species previous formulations of the concept, we hope the idea belonging to communities was better thought of as will emerge for the reader as it has for the authors. species fitting into a habitat. We ultimately step into synthesis, and present a new The individualistic concept provides the raw biologi- framework for the novel ecosystems concept. cal material from which the novel ecosystems concept is made. Because the flora and fauna move independ- ently of one another in response to their environment, 5.2 FOUNDATIONS OF THE NOVEL communities are constantly in flux at some temporal ECOSYSTEMS CONCEPT resolution. Without this property of the biosphere, novel ecosystems would not be possible. In some areas, The novel ecosystem concept rests on three founda- such as those undergoing secondary succession, the tional principles, the first of which follows from Gleason fluctuations may be rapid and repeatable; in others, (1926): “It may be said that every species of plant is a such as those undergoing changes in temperatures or law unto itself, the distribution of which in space carbon dioxide concentrations, the ‘fluctuations’ may depends on its individual peculiarities of migration be directional and permanent. Temperature drove tree and environmental requirements.” In Gleason’s time, species to move during glacial advances and retreats ecology was populated largely by naturalists and bota- (Davis 1981), while CO2 concentrations in the ocean nists with an instinctual urge to classify. It is remark- caused shifts in foraminifera communities (Pagani able that he was able to look at the same forests and et al. 1999). Where the ranges of species expanded or grasslands as his predecessors and devise so different a contracted (whatever the cause), communities were hypothesis to explain them, one that was abjectly aller- altered. Most recently, a massive increase in the rate of gic to classification. Gleason viewed communities as species introductions has unfolded: prior to human whisperers of organization: the ephemeral overlapping arrival, Hawaii lacked reptiles, amphibians, flightless of ranges combined with a faint alignment. Decades mammals and ants, yet now it has all in abundance went by before the individualistic concept became (Ziegler 2002). more widely accepted, but Gleason did live to see it The earlier examples highlight community changes embraced by giants such as John Curtis and Robert caused both by biotic (i.e. range shifts) and abiotic Origins of the novel ecosystems concept 47 4 7 8 710 10 4 4 11 8 5 5 6 8 10 9 6 10 11 4 6 10 6 12 10 5 14 10 8 14 8 12 12 2 2 9 12 11 2 7 7 8 8 12 14 12 11 4 5 7 8 7 14 14 4 2 6 6 14 6 6 6 5 10 >14 6 6 8 10 23 12 8 10 13 10 8 21 15 11 14 Larix 15 5 12 12 Castanea Fagus laricina dentata grandi- >14 Larch >13 14 14 Chestnut folia Beech 0 400 km 0 400 km 0 400 km Figure 5.1 The Gleasonian framework of ecological communities. In the North American temperate forests shown, black bands indicate the southernmost extent for each species in time (1000 years). The varying trajectories of these ranges reflect the independent movement of each species as they coped with warming temperatures at the close of the Wisconsinan glaciation. From Davis (1983). Reproduced by permission of Missouri Botanical Garden Press. forcing (i.e. responses to environmental change) and are directional and permanent. Indeed, Chapin and this leads to the second foundational concept upon Starfield (1997) first invoked the term ‘novel ecosys- which novel ecosystems rest: the biotic and abiotic tem’ to characterize what they believed would be the characteristics of an ecosystem are tethered. They ultimate outcome of anthropogenic changes to climate, interact, following from one another and feeding back disturbance regimes and species composition in boreal on one another (Tansley 1935; Jenny 1941; Odum latitudes. Old views in terrestrial plant succession 1969; Naeem 2002). Jenny (1980) made perhaps referred to human-modified ecosystems such as farms the most humorous declaration, calling the biotic as “dis-climax” (Clements 1916). Note the root “dis” factor “. a real bugbear. Like everybody else I could see (having a negative or reversing force) and the similar- that the vegetation affects the soil and the soil affects ity to the more modern term for the same systems: the vegetation, the very circulus vitiosus I was trying ‘degraded’ (reduced in rank). Humans have unequivo- to avoid.” Twenty years later, Naeem (2002) came to cally had a negative impact on myriad species and a similarly baffling (but editorially sanitized) conclu- ecosystems, particularly by co-opting land for the pro- sion: “. biodiversity is a product of its environment, duction of resources (Vitousek et al. 1997). However, and the antithesis, that the environment is, in part, a when we are left to qualify a river that has increased in product of the organisms within it, is also correct.” trophic complexity due to non-native fishes on Hawaii Empirical studies spanning the globe confirm strong (Ziegler 2002), or an old Puerto Rican baseball field interactions between the biotic and abiotic parts of that is thick with pantropical tree species whose leaves ecosystems, from tubeworms mollifying habitat around have never before competed for sunlight (Lugo and oceanic vents (Cordes et al.
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