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Serpentine: The Evolution and Ecology of a Model System

Susan Harrison
Nishanta Rajakaruna
Copyright Date: 2011
Edition: 1
Pages: 464
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  • Book Info
    Book Description:

    Serpentine soils have long fascinated biologists for the specialized floras they support and the challenges they pose to plant survival and growth. This volume focuses on what scientists have learned about major questions in earth history, evolution, ecology, conservation, and restoration from the study of serpentine areas, especially in California. Results from molecular studies offer insight into evolutionary patterns, while new ecological research examines both species and communities.Serpentinehighlights research whose breadth provides context and fresh insights into the evolution and ecology of stressful environments.

    eISBN: 978-0-520-94845-7
    Subjects: Ecology & Evolutionary Biology

Table of Contents

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  1. Front Matter
    (pp. I-IV)
  2. Table of Contents
    (pp. V-VI)
    (pp. VII-X)
    (pp. XI-XII)
    (pp. XIII-XIV)

    Terrestrial life, perched on the Earth’s continental crust, has evolved on soils formed from relatively low-density rocks such as granite that are rich in silica, calcium, potassium, and phosphorous. The chemistry of these soils is usually amenable to plant growth almost by definition. Deeper in the Earth, forming its mantle and most of its oceanic crust, are darker and denser ultramafic (high-iron and -magnesium) rocks and minerals. Near the surface they may become serpentinized—altered in contact with water. These submarine rocks are seldom seen on land but occasionally become stranded on the edges of continents during the process of...


    • 1 Serpentinites and Other Ultramafic Rocks: Why They Are Important for Earth’s History and Possibly for Its Future
      (pp. 3-28)
      Eldridge M. Moores

      Geology is a historical science, one of the “storytelling sciences,” not simply a laboratory science. As such, geologists try to not only understand basic and timeless principles related to the rocks being studied but also give an account of what has happened in the past, and when possible, use this past history to forecast future events (Primack and Abrams, 2006: 17).

      Serpentine, strictly speaking, is a mineral. Rocks formed mostly of serpentine are called serpentinites. Serpentine forms chiefly by the alteration (hydration) of the minerals olivine and pyroxene, found mostly in rocks called peridotites, a type of ultramafic rock. The...

    • 2 Microbes in Extreme Environments: Implications for Life on the Early Earth and Other Planets
      (pp. 29-48)
      Dawn Cardace and Tori M. Hoehler

      Delineating the boundaries of the biosphere is an area of exciting research, integrating work in biology, geology, and chemistry. Of particular interest is how microorganisms adapt to survive under stress (e.g., extreme acidity or alkalinity, temperature fluctuations, changing activity of water) and persist as communities, perhaps over geologically meaningful time scales. Considering how terrestrial environments support and/or challenge microbial life in Earth’s most extreme settings also fuels scientific investigation as we recognize habitable zones (where conditions appear appropriate for life as we know it) in our solar system and around stars other than our own Sun. In this chapter, we...

    • 3 Phylogenetic Patterns of Endemism and Diversity
      (pp. 49-70)
      Brian L. Anacker

      Every species in nature uses a subset of the habitats available to it, but those with narrow ranges and unique adaptations have traditionally captured the attention of naturalists and ecologists (Futuyma and Moreno, 1988; Stevens, 1989; Brown, 1995; Berenbaum, 1996; Losos et al., 1998; Gaston and Blackburn, 2000; Schluter, 2002; Fine et al., 2005; Grant and Grant, 2007). In plants, endemism to a particular soil type is an especially common and important form of habitat specialization. As Kruckeberg pointed out, “Endemism is the hallmark of specialized edaphic habitats” (Kruckeberg, 2002). Edaphic endemism contributes to species diversity by promoting spatial turnover...

    • 4 Plant Speciation
      (pp. 71-96)
      Kathleen M. Kay, Kimiora L. Ward, Lorna R. Watt and Douglas W. Schemske

      The world’s serpentine regions are known for their striking levels of endemism and the distinctive flora they possess relative to surrounding areas. Although much work has been done to catalog the plant diversity on serpentine, including taxonomic, morphological, and physiological diversity, relatively little has been done to understand the evolutionary origins of serpentine species. Yet serpentine species present an excellent system to study the general processes of plant speciation. We critically evaluate the theory and evidence for the mechanisms of plant speciation on serpentine. We highlight the contributions that studies of serpentine plants have made to the general understanding of...

    • 5 Intraspecific Variation, Adaptation, and Evolution
      (pp. 97-138)
      Ryan E. O’Dell and Nishanta Rajakaruna

      Intraspecific variation refers to the genotypic (genetic) and resulting phenotypic (morphological and physiological) variation found within a species. Variation within a species is crucial for adaptation and evolution by natural selection. Over time, selection can result in genetically distinct populations of a species that are adapted to specific environmental conditions. Such populations are referred to as ecotypes. The study of distinct climatic, elevational, latitudinal, geographic, and edaphic ecotypes has provided much insight into the process of evolution by natural selection (Briggs and Walters, 1997; Levin, 2000; Silvertown and Charlesworth, 2001).

      Edaphic islands, characterized by distinct soil characteristics, provide unique settings...

    • 6 Genomic Approaches to Understanding Adaptation
      (pp. 139-154)
      Eric J. von Wettberg and Jessica Wright

      Advances in molecular biology technologies allow us to ask ecological and evolutionary questions in more elaborate and detailed ways in any species—not just a narrow set of model organisms. These technological advances allow us to probe the mechanistic and genetic basis for expressed traits, explore patterns of genetic variation in organisms for signs of selection and evidence of past population processes, and identify cryptic members of communities, such as soil microbes that are critical to both plant success and ecosystem processes. Here we illustrate the power of these tools with examples from serpentine systems.

      Next-generation sequencing technologies (Shendure and...

    • 7 Local Adaptation in Heterogeneous Landscapes: Reciprocal Transplant Experiments and Beyond
      (pp. 155-180)
      Jessica W. Wright and Maureen L. Stanton

      Adaptation to different selection pressures (i.e., adaptive differentiation) is thought to be a major driver of diversification and accordingly has been a focus of evolutionary biology since the early work of Turesson (1922) and Clausen, Keck, and Hiesey (1940, 1948). The classic work of Clausen’s team revealed the power of reciprocal transplant experiments to determine whether populations are locally adapted to their home environment, characterize phenotypes that differentiate locally adapted ecotypes, and, if sufficiently well replicated, identify axes of environmental variation along which adaptive differentiation has occurred. Their extensive experiments and observations set a new standard for characterizing ecologically relevant...

    • 8 Herbivory and Other Cross-Kingdom Interactions on Harsh Soils
      (pp. 181-200)
      Sharon Y. Strauss and Robert S. Boyd

      Edaphically stressful substrates, like serpentine, present plants with challenges that differ from other substrates. Stressful substrates often require plant adaptations to toxicity stressors like heavy metals, as well as to low nutrient concentrations or abnormal ratios of necessary elements (Proctor and Woodell, 1975; Ellis and Weis, 2006). Pressures from enemies may also be greater on edaphically stressful substrates than on normal soils (Fine et al., 2005). On the other hand, substrates with high concentrations of heavy metals (like serpentine) may provide plants with opportunities for elemental defense, such as heavy metal accumulation (Martens and Boyd, 1994; Mroz, 2008). We describe...

    • 9 Invasions and the Evolution of Range Limits
      (pp. 201-220)
      Diane M. Thomson, Brooke S. Baythavong and Kevin J. Rice

      Understanding how species are sometimes able to expand their range boundaries into a novel environment is key to managing biological invasions, a major component of global change and threat to biodiversity (Hastings et al., 2005). Paradoxically, although invasions are a critical conservation problem, they also can potentially contribute to the development of solutions by creating opportunities to study how and why range limits form or change. Ecology and evolution are rich in theory and models explaining the development and maintenance of range limits, so there is no lack of hypotheses about these processes (Lenormand, 2002; Kawecki, 2008; Gaston, 2009). However,...


    • 10 Plant Competition and Facilitation in Systems with Strong Environmental Gradients
      (pp. 223-236)
      Kara A. Moore and Sarah C. Elmendorf

      The distinctive plant communities and environmental gradients found in serpentine ecosystems provide a rich palette for testing fundamental questions about species interactions. Plant–plant interactions on serpentine soils differ from those on nonserpentine soils in the identity, abundance, and diversity of interacting plants. Interactions on serpentine also are distinct due to the high-stress, low-resource, and often extremely heterogeneous environment. One of the most striking elements of serpentine plant communities is the frequent dominance of native species in an otherwise exotic-dominated landscape. Serpentine soils are a significant refuge for native species diversity globally and characterized by high levels of species endemism...

    • 11 Community Invasibility: Spatial Heterogeneity, Spatial Scale, and Productivity
      (pp. 237-248)
      Kendi F. Davies

      Invasive species are one of the most significant threats to native species diversity, and identifying the factors that make places more or less invasible has been one of the most important issues in the study of invasions (Wilcove et al., 1998; Pimentel et al., 2000). From a theoretical perspective, the reasons some communities are more invasible than others is a question that continues to intrigue ecologists (Levine et al., 2004; Davies et al., 2005; Fridley et al., 2007; Stohlgren et al., 2008; Cadotte et al., 2009) because it underlies fundamental concepts in community ecology: species coexistence and assembly (Chesson, 2000;...

    • 12 Disturbance and Diversity in Low-Productivity Ecosystems
      (pp. 249-274)
      Hugh D. Safford and Chris R. Mallek

      Ecological theory predicts that the amount of resources available in an ecosystem should affect its response and sensitivity to ecological disturbances like fire, herbivory, and soil disturbance. Plant stature and life form, biomass, rates of growth, and plant palatability are all influenced by habitat productivity, and these factors play key roles in determining disturbance frequencies and intensities (Pickett and White, 1985; Bond and van Wilgen, 1996; Grime, 2001; Table 12.1). Theory and empirical investigations find that site quality is often related to rates of competitive displacement, with relatively unproductive, less competitive environments less reliant on disturbance for diversity regulation (Huston,...

    • 13 Plant-Pollinator Interactions in Naturally Fragmented Habitats
      (pp. 275-296)
      Amy T. Wolf and Robbin Thorp

      Serpentine landscapes appear to provide ideal systems for studying the effects of habitat isolation on species interactions, including the relationship between flowering plants and their pollinators. Serpentine plant assemblages are generally discrete (Harrison et al., 2006), highly endemic (Skinner and Pavlik, 1994; Safford et al., 2005), and physiologically specialized (Brooks, 1987). Maps from California (Kruckeberg, 1984), Cuba (Reeves et al., 1999), New Caledonia (Dawson, 1981), South Africa (Williamson and Balkwill, 2006), and elsewhere illustrate the highly insular nature of serpentine substrates, a pattern that is repeated at multiple geographic scales (Harrison et al., 2006). In this chapter, we explore the...

    • 14 Spatial Ecology: The Effects of Habitat Patch Size, Shape, and Isolation on Ecological Processes
      (pp. 297-308)
      Susan Harrison

      Serpentine outcrops, with their distinctive floras strongly contrasting with those of surrounding landscapes, bear as strong a resemblance to islands as any habitat on land. As Kruckeberg (1991) first suggested, it seems intuitive that the diversity of life on serpentine outcrops may be shaped by the same spatial attributes that are so important in islands—areas, distances from one another and from a mainland, and the qualities of the “matrix” through which organisms must move to reach them. This chapter reviews all the available studies that address effects of the size and isolation of serpentine outcrops on ecology of their...

    • 15 Systematic Conservation Planning: Protecting Rarity, Representation, and Connectivity in Regional Landscapes
      (pp. 309-328)
      James H. Thorne, Patrick R. Huber and Susan Harrison

      Ecologists have long recognized the need for a strategy of land-based conservation aimed at protecting the full spectrum of biological diversity. The first effort at land protection for conservation in the United States, the 1872 creation of the National Park system, was motivated mainly by appreciation of beauty and outdoor recreation; later, lands were set aside for timber supply (the Forest Reserve Act of 1891) and huntable wildlife (the National Wildlife Refuge system, 1903). By the mid-twentieth century, it was clear that many species and natural communities were unprotected and at risk. The Nature Conservancy formed in 1946 as an...

    • 16 Biodiversity, Ecosystem Functioning, and Global Change
      (pp. 329-358)
      David Hooper

      Environmental changes, including elevated CO₂, climate change, enhanced nutrient deposition, change in land use type and intensity, and species invasions, are contributing to worldwide loss of biodiversity (Vitousek, 1994). Ecosystem responses may either buffer or exacerbate these changes through various feedback loops, with important implications for the services ecosystems provide to humanity (Daily, 1997; Millennium Ecosystem Assessment, 2005; Suding et al., 2008). Whole-ecosystem studies provide a complement to modeling approaches and smaller scale studies to test whether our understanding of the underlying mechanisms scale up, and if not, which other processes need more consideration (Field et al., 1996; Vitousek, 2004)....

    • 17 Climate Change and Plant Communities on Unusual Soils
      (pp. 359-382)
      Ellen I. Damschen, Susan Harrison, Barbara M. Going and Brian L. Anacker

      Human-caused climate change is altering patterns of temperature, precipitation, and net primary productivity across the globe (Nemani et al., 2003; IPCC, 2007), and many ecological responses have already been detected (Parmesan and Yohe, 2003). One of the greatest challenges facing conservation biologists today is predicting the fates of species and communities under altered climates, which is difficult for at least three reasons. First, the direction and magnitude of climatic changes vary across regions (Nemani et al., 2003; IPCC, 2007). Second, species responses are highly variable (IPCC, 2007). Third, climate change can have complex indirect effects through altering species interactions, such...

    • 18 Restoration and Revegetation of Harsh Soils
      (pp. 383-414)
      Ryan E. O’Dell and Victor P. Claassen

      Human activities such as fire regime alteration, introduction of non-native plant and animal species, livestock grazing, agriculture, logging, pollution, motorized recreation, construction, and mining can dramatically alter ecosystem structure and function. Adverse impacts to ecosystems from human activities may include alteration of plant, animal, and microbial community cover/abundance, structure, composition, and diversity; soil removal and/or erosion; and landscape instability (mass wasting/landslide).

      Restoration, reclamation, rehabilitation, and revegetation of degraded ecosystems are undertaken for the benefit of human health, environment, or both. Restoration is the process of returning an ecosystem to a condition very similar in ecological structure and function to what...


    • 19 What Have We Learned from Serpentine in Evolution, Ecology, and Other Sciences?
      (pp. 417-428)
      Susan Harrison and Nishanta Rajakaruna

      We conclude this book by briefly reviewing some of the most provocative conclusions from the foregoing chapters. We hope these examples, though not exhaustive, illustrate the wealth of general scientific understanding that has come from studying serpentine ecosystems. If this book provides inspiration for future collaborations between disparate workers—earth and life scientists, evolutionary biologists and gene-free ecologists, naturalists and theoreticians, basic scientists and those who aim to save the world—then serpentine will have served as a good model system for the excitement and synergy that comes from crossing sharp boundaries.

      Prior to the plate tectonics revolution, geologists believed...

    (pp. 429-434)
    (pp. 435-446)