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Evolution of Ecosystems: Terrestrial

Scott A Elias, Royal Holloway University of London, London, UK

As life has evolved on Earth, it has been organized into biological communities of increasing complexity. These communities are temporary associations of species that shift in response to environmental disturbances.

Secondary article
Article Contents
. Introduction . Evidence from the Fossil Record . Ecosystem Stability . Ecosystem Response to Disturbance

As life on Earth evolved, the organisms began interacting with one another, rst in the seas, then on the land. These interactions became increasingly more complex as the organisms themselves became more sophisticated. The earliest biological communities on land were probably quite simple, with primitive plants being eaten by arthropods and other invertebrates. We can see in the fossil record how ecological complexity evolved alongside the organisms that formed these communities. A brief overview of the fossil record provides glimpses into these ancient ecosystems. The chronology of the fossil record on land is shown in Figure 1.

about more recent ecosystems, and progressively less about more ancient ones, as the fossil record of more recent eras is much more complete than the record of more ancient times. Nevertheless, through many years of palaeontological research, scientists have built up considerable knowledge of the history of life on land, and the ways in which that life was organized into biological communities.

Palaeozoic era
The rst evidence for life on land comes from the Ordovician period, when primitive plants invaded the land. Palaeozoic plant life did not begin to resemble modern plants until about 420 million years ago. Other than a few types of primitive arthropods, we have little evidence of the animal life on land from this period (Moullade and Nairn, 1991). Fossils from the Silurian period (440410 million years ago) provide the rst substantial evidence for the development of terrestrial ecosystems, including vascular plants and primitive

Evidence from the Fossil Record

Our knowledge of the history of ancient ecosystems is based on the fossil record. Accordingly, we know more

Era Cenozoic

Period Quaternary Tertiary Cretaceous

Age (millions of years) 2 65

Major events in ecosystem evolution on land Glaciations; extinction event at 10 000 years BP Increasing diversity of flowering plants and mammals Rise of modern insect orders, flowering plants, mammals; extinction event ends the age of dinosaurs Dinosaur diversity increases; first winged reptiles and first birds Rise of modern conifers and cydads; first dinosaurs Mass extinction event; rise of gymnosperms Tree ferns increase; ancestors of birds, mammals and reptiles

Mesozoic Jurassic Triassic Permian Carboniferous Palaeozoic

146 208 245 286

360 Devonian Silurian Ordovician 510 410 445 First trees and forests; first land vertebrates First vascular plants and arthropods on land First primitive land plants

Figure 1 Summary of geological time, showing the major events in ecosystem evolution on land.

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Evolution of Ecosystems: Terrestrial

arachnids. Silurian plants had branching stems with spore-producing bodies (sporangia) at their tips, but no leaves. Cooksonia has generally been considered the oldest known land plant. Several species of this plant are known from fossil specimens in North America, Europe, Asia and Africa. However, the lycophyte plant, Baragwanathia, is structurally more complex than Cooksonia, and Silurian fossils of this plant have been found in Australian fossil beds that are signicantly older than the beds containing Cooksonia fossils in the Northern Hemisphere. The Devonian period gave rise to two major lines of vascular plants (Banks, 1972). Early Devonian plants were less than 1 m tall, but Late Devonian vegetation included the rst trees and forests. Thus, the level of ecological complexity expanded dramatically during this period, as the development of forest vegetation led to the creation of more potential ecological niches for animal life. It was during this period that two major animal groups colonized the land: the rst primitive insects and the rst land-living vertebrates, called tetrapods. These primitive land-dwellers, labyrinthodont amphibians such as Ichthyostega, essentially sh with legs, are thought to have given rise to all later terrestrial vertebrate groups (MacDonald, 1994). By the Early Carboniferous period, they had evolved into clearly dened amphibian groups. The development of seedbearing plants added complexity to the diet of plantfeeding animals. Seeds are an excellent source of animal nutrition because they contain high concentrations of both carbohydrates and proteins. Thus a new ecological niche became available in the Devonian period. A major habitat was rst exploited by insects during the Carboniferous period: the skies. It was during this long interval of warm, humid, tropical climate that the rst winged insects appeared, and these were the only ying animals for more than 100 million years. One environmental change in the Carboniferous period may have been the trigger for an important step in vertebrate evolution: the development of the amniote egg. During later parts of the Carboniferous, the climates of the world tended to become drier. Amniote eggs have a shell to help prevent drying, and a series of membranes that surround the developing embryo. The amniote egg allowed the ancestors of birds, mammals and reptiles to reproduce on dry land, thus freeing these animals from having to lay eggs in water. The Permian period marked a turning point in the history of life on Earth. The end of the Permian (245 million years ago) saw the largest mass extinction event in Earths history. This event most greatly aected marine communities, but in some ways it was just as signicant in shaping the course of vertebrate evolution, because the extinction of existing forms allowed new groups to dominate, particularly the dinosaurs (Erwin, 1993). Modern conifers also made their rst appearance in the Permian.

Mesozoic era
The Triassic period was a time of transition in which some hold-overs from the Permian ourished briey, but new groups arose that went on to dominate the Mesozoic biota. The major winners in this biological lottery were the conifers, cycads and dinosaurs. Late Triassic fossil beds also hold evidence for the rst primitive (dicynodont) mammals (Sennikov, 1996). These were no larger than rats, and were probably nocturnal. The rst true dinosaurs appeared at about the same time as these early mammals. Giant plant-eating dinosaurs dominated the Jurassic period, eating ferns, cycads and conifers. The conifers of that period included relatives of modern trees, such as redwoods, cypresses, pines and yews. Smaller carnivorous dinosaurs were the predators in the Jurassic, and ying reptiles (pterosaurs) and the rst birds claimed their niches in the sky. The Cretaceous period was the last period in the Age of Dinosaurs, but new dinosaur species appeared right up to the end, when a mass extinction event took place (65 million years ago). There is now convincing evidence that this mass extinction was caused mainly by an asteroid that struck the Gulf of Mexico region, sending a cloud of dust, smoke and water vapour into the atmosphere, probably darkening the planet for many years (Dressler and Sharpton, 1999). In this cold, dark world, the vegetation withered and died, followed by the plant-eating dinosaurs and their predators and scavengers. But waiting in the wings were new groups of plants and animals, ready to take advantage as the old order passed away.

Cenozoic era
The Tertiary period, sometimes called the Age of Mammals, began with a veritable explosion of new mammal groups, some of which have persisted until today, but many of which have become extinct. Early Tertiary mammals were no larger than small bears of today; most species were a lot smaller. By the middle Tertiary, plate tectonic shifts were accompanied by climatic change, marked by a general cooling. Glaciers formed in Antarctica for the rst time during the Cenozoic period, and the tropical zone diminished in size as temperate woodlands and grasslands expanded. The cooling trend culminated in the Ice Ages of the Pleistocene. Many modern mammalian groups had evolved by the middle Tertiary, including the ancestors of dogs, cats, rhinoceroses, pigs, horses, camels and the rst anthropoid apes (ancestors of modern humans). Mammal body size had grown much larger by this time. By the end of the Tertiary period, most of the modern groups of animals and plants were already in place. The positions of the continents, combined with the disturbance to previous atmospheric circulation patterns brought about by the Himalayan uplift, ushered in the Quaternary period, and a series of at least 17 glaciations.

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Evolution of Ecosystems: Terrestrial

From an ecological perspective, perhaps the most signicant evolutionary event during the Quaternary period was the rise of our own species, Homo sapiens, as we have reshaped the biotic world to our own liking during the time since the last glaciation, bringing about more extinctions of other species than most of the previous mass extinction events in the geological record.

community remains relatively stable, even though some individual species suer. Again, this argues for greater stability in ecosystems with high biodiversity.

Ecosystem Response to Disturbance

One key element in ecosystem dynamics is the ability of the system to withstand disturbances, such as climatic change or large-scale res. The resistance of an ecosystem to disturbances, and the speed at which it recovers after the disturbance (resilience), are two important components of ecosystem stability. Some ecologists believe that biodiversity increases both the resilience and resistance of ecosystems. Loreau and Behera (1999) built a mathematical model to test this theory, and concluded that there is no simple relationship between diversity and an ecosystems ability to resist change or recover after a disturbance. As the fossil record has shown, large-scale disturbances have often been the catalysts for change in the biological world. If such disturbances are strong enough (such as an asteroid impact) or are essentially permanent (such as the breaking up of supercontinents), no ecosystem will be able to recover, no matter how great its level of biodiversity.

Ecosystem Stability
The ecosystems of the Quaternary period, being the most recent, are also the ones we know most about. This is partly because we can observe modern ecosystems, and partly because the Quaternary fossil record is more intact than that of the previous periods. The Quaternary fossil record is shedding considerable light on how ecosystems work over long periods of time. We now know, for instance, that the most important ecological events take place at time scales of centuries and millennia. As such, these ecological changes are not often observable by modern biologists, because their studies are too brief to include them. Largescale shifts in the ranges of plants take place over hundreds and sometimes thousands of years. At this time scale, no ecosystem is truly stable, because species ranges move about constantly, often in response to environmental changes (Johnson and Mayeux, 1992). But the associations of species in some communities can be surprisingly long lasting. What is the glue that holds these communities together? There are numerous theories in the ecological literature that address this problem. Haydon (2000) argues that stable communities require high levels of connectance, that is, a high degree of interaction between species. This conclusion was borne out in a study by Deruiter et al. (1995), who examined food webs from native and agricultural soils, and found patterns connecting the trophic levels that appear to be important to ecosystem stability. This patterning resulted directly from the way energy ows through the food webs, showing that energy ow and community structure govern ecosystem stability by imposing stabilizing patterns of interaction strengths. Yachi and Loreau (1999) focused on the biodiversity of ecosystems as a key element in their stability. According to their hypothesis, biodiversity ensures ecosystems against unravelling because the presence of many species provides a sort of ecological safety net, so that even if some species are removed, others will maintain the functioning of the ecosystem. Kaufman et al. (1998) developed an ecological model that predicted there is indeed a critical level of biodiversity, below which ecosystems are highly susceptible to extinction. Tilman (1996) studied the eects of drought on a prairie ecosystem in North America. This study showed that when climatic variations harm some species in a community, the numbers of unharmed competitors increase. Thus the overall vigour of the

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Evolution of Ecosystems: Terrestrial

the National Academy of Sciences of the United States of America 96: 14631468.

Further Reading
Beerbower R (1985) Early development of continental ecosystems. In: Tiney BH (ed.) Geological Factors and the Evolution of Plants, pp. 47 91. New Haven, Connecticut: Yale University Press. Kenrick P and Crane PR (1997) The origin and early evolution of plants on land. Nature 389: 3339.

Matthews JV (1974) Quaternary environments at Cape Deceit (Seward Peninsula, Alaska): evolution of a tundra ecosystem. Geological Society of America Bulletin 85: 13531384. Shear WA (1991) The early development of terrestrial ecosystems. Nature 351: 283289. Tschudy RH, Pilmore CL, Orth CJ, Gilmore JS and Knight T (1984) Disruption of the terrestrial plant ecosystems at the Cretaceous Tertiary boundary, western interior. Science 225: 10301032. Vitousek PM, Mooney HA, Lubchenco J and Melillo JM (1997) Human domination of Earths ecosystems. Science 277: 494499.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group /