Paleoecology and Late Quaternary Environments of the Colorado Rockies

Present-day environments cannot be completely understood without knowledge of their history since the last ice age. Paleoecological studies show that the modern ecosystems did not spring full-blown onto the Rocky Mountain region within the last few centuries. Rather, they are the product of a massive reshuffling of species that was brought about by the last ice age and indeed continues to this day. Chronologically, this chapter covers the late Quaternary Period: the last 25,000 years. During this interval, ice sheets advanced southward, covering Canada and much of the northern tier of states in the United States. Glaciers crept down from mountaintops to fill high valleys in the Rockies and Sierras. The late Quaternary interval is important because it bridges the gap between the ice-age world and modern environments and biota. It was a time of great change, in both physical environments and biological communities. The Wisconsin Glaciation is called the Pinedale Glaciation in the Rocky Mountain region (after terminal moraines near the town of Pinedale, Wyoming; see chapter 4). The Pinedale Glaciation began after the last (Sangamon) Interglaciation, perhaps 110,000 radiocarbon years before present (yr BP), and included at least two major ice advances and retreats. These glacial events took different forms in different regions. The Laurentide Ice Sheet covered much of northeastern and north-central North America, and the Cordilleran Ice Sheet covered much of northwestern North America. The two ice sheets covered more than 16 million km2 and contained one third of all the ice in the world’s glaciers during this period. The history of glaciation is not as well resolved for the Colorado Front Range region as it is for regions farther north. For instance, although a chronology of three separate ice advances has been established for the Teton Range during Pinedale times, in northern Colorado we know only that there were earlier and later Pinedale ice advances. We do not know when the earlier advance (or multiple advances) took place. However, based on geologic evidence (Madole and Shroba 1979), the early Pinedale glaciation was more extensive than the late Pinedale was.

Plant-Herbivore Interactions

The alpine provides a tremendous opportunity for studying plant-herbivore interactions at the population, community, and ecosystem levels. For herbivores, variations in topography and microclimate result in a relatively large amount of spatial variation in plant communities within short distances (chapter 6). A large community of herbivores, from nematodes to grasshoppers to elk, occurs on Niwot Ridge. Furthermore, given the low rates of nutrient availability in alpine soils (Fisk and Schmidt 1995; chapter 12) combined with the slow-growing perennial habit of the vegetation, alpine plants should, in theory, invest heavily in defense against herbivores (Coley et al. 1985). The goal of this chapter is to provide: (1) a summary of the feeding behaviors of the herbivores on Niwot Ridge, (2) information on the nutritional and secondary chemistry of plants on Niwot Ridge as it relates to herbivory, and (3) a review of hypotheses on community dynamics of herbivores and plants relevant to the alpine. The ultimate objective is to provide a synthesis of information that will stimulate interest in alpine tundra as a system for studying the dynamics of plant-herbivore interactions at all levels of ecological organization. The flora of Niwot Ridge has been divided into six communities (May and Webber 1982; chapter 6). Regardless of community association, nearly all of the plant species occurring on the ridge are perennials and several are very long lived (May and Webber 1982). Communities can change across small spatial scales (meters), and community origin and maintenance are believed to be largely determined by abiotic factors (Walker et al. 1994; chapter 6). However, several studies suggest that biotic factors such as herbivory may have a significant impact on plant community dynamics (Huntly et al. 1986; Davies 1994). There is significant variation in the nutritional composition of plants on Niwot Ridge. Generally, and in the absence of plant secondary compounds, species that are high in nitrogen and low in fiber are presumed to be the most desirable as forage. Based solely on these nutritional variables, the clover Trifolium parryi is hypothesized to be one of the more-preferred forages, whereas alpine sandwort, Minuartia obtusiloba, should be one of the less-preferred food items.

Nitrogen Cycling

In this chapter, we discuss the current understanding of internal N cycling, or the flow of N through plant and soil components, in the Niwot Ridge alpine ecosystem. We consider the internal N cycle largely as the opposing processes of uptake and incorporation of N into organic form and mineralization of N from organic to inorganic form. We will outline the major organic pools in which N is stored and discuss the transfers of N into and from those pools. With a synthesis of information regarding the various N pools and relative turnover of N through them, we hope to provide greater understanding of the relative function of different components of the alpine N cycle. Because of the short growing season, cold temperatures, and water regimes tending either toward very dry or very wet extremes, the alpine tundra is not a favorable ecosystem for either production or decomposition. Water availability, temperature, and nutrient availability (N in particular) all can limit alpine plant growth (chapter 9). Cold soils also inhibit decomposition so that N remains bound in organic matter and is unavailable for plant uptake (chapter 11). Consequently, N cycling in the alpine often is presumed to be slow and conservative (Rehder 1976a, 1976b; Holzmann and Haselwandter 1988). Nonetheless, studies reveal large spatial variation in primary production and N cycling in alpine tundra across gradients of snowpack accumulation, growing season water availability, and plant species composition (May and Webber, 1982, Walker et al., 1994, Bowman, 1994, Fisk et al. 1998; chapter 9). Furthermore, evidence for relatively large N transformations under seasonal snowcover (Brooks et al., 1995a, 1998) and maintenance of high microbial biomass in frozen soils (Lipson et al. 1999a) provide a complex temporal component of N cycling on Niwot Ridge. Our discussion of N cycling on Niwot Ridge will focus on two main points: first, the spatial variation in N turnover in relation to snowpack regimes and plant community distributions; and second, the temporal variability of N transformations during both snow-free and snow-covered time periods.

Controls on Decomposition Processes in Alpine Tundra

The snowpack gradient in the alpine generates a temperature and moisture gradient that largely controls organic matter decomposition. While low temperatures constrain decomposition and mineralization (chapter 12), moisture appears to be the strongest source of landscape variation in the alpine, with surface decay rates of plant materials highest in moist and wet meadow habitats. Despite a longer snow-free season and higher surface temperatures in dry meadows, decay in these areas is substantially lower than in moist meadows. Studies of decay rates of roots within the soil indicate that decay is uniformly low in all habitats and is limited by low temperatures and perhaps by the absence of certain groups of decomposer invertebrates. As in other ecosystems, substrate quality indices such as nitrogen and lignin content can be shown to be important factors influencing the rate of decay of specific substrates. Alpine ecosystems were overlooked during the flurry of activity associated with the extensive ecosystem science programs of the 1960s and 1970s. With the few exceptions to be discussed here, decomposition studies in cold regions were conducted in arctic tundra or northern temperate and boreal forests. The need for this information in conjunction with efforts to understand carbon cycling in the alpine stimulated a substantial research effort in the 1990s. Studies have included both the effects of landscape location on decay (O’Lear and Seastedt 1994; Bryant et al. 1998), information on the importance of substrate chemistry on decomposition processes (Bryant et al. 1998), and preliminary information on some of the decomposer organisms (O'Lear and Seastedt 1994; Addington and Seastedt 1999). Niwot Ridge researchers also participated in the Long-term Intersite Decomposition Experiment Team (LIDET) study, which involved placement of a dozen different litter types in the alpine and in 27 other sites from the tropics to the arctic tundra (Harmon 1995). All but one of the plant species used in the LIDET experiments were exotic to the alpine. Collectively these studies have provided sufficient information to represent the alpine in global decomposition modeling efforts.

Geomorphic Systems of Green Lakes Valley

There are at least three justifications for the examination of the geomorphology of the area in which ecosystem studies are conducted. First, the present landscape and the materials that make it up provide the substrate on which ecosystem development occurs and may impose constraints, such as where soil resources are limited, on that development. Second, the nature of the landscape and the geomorphic processes acting on it often define a large part of the disturbance regime within which ecosystem processes occur (Swanson et al. 1988). Third, the processes of weathering, erosion, sediment transport, and deposition that define geomorphic dynamics within the landscape are themselves ecosystem processes, for example, involving the supply of resources to organisms. In this last context, it is noteworthy that drainage basins (also called watersheds or catchments) were recognized as units of scientific study during a similar time period in both geomorphology and ecology (Slaymaker and Chorley 1964; Bormann and Likens 1967; Chorley 1969). The drainage basin concept, the contention that lakes and streams act to integrate ecological and geomorphic processes, remains important in both sciences and underlies the studies in Green Lakes Valley reviewed here. Over the past 30 years, Niwot Ridge and the adjacent catchment of Green Lakes Valley have been the subject of much research in geomorphology. Building on the studies of Outcalt and MacPhail (1965), White (1968), and Benedict (1970), work has emphasized the study of present-day processes and dynamics, especially of mass wasting in alpine areas. These topics have been reviewed by Caine (1974, 1986), Ives (1980), and Thorn and Loewenherz (1987). Studies of geomorphic processes have been conducted in parallel with work on Pleistocene (3 million to 10,000 yr BP) and Holocene (10,000 yr BP to present) environments in the Colorado Front Range (Madole 1972; Benedict 1973) that have been reviewed by White (1982). This chapter is intended to update those reviews in terms that complement the presentation of ecological phenomena such as nitrogen saturation in the alpine (chapter 5) as well as to refine observations and conclusions of earlier geomorphic studies.

Plant Nutrient Relations

Alpine soils do not generally exhibit high levels of inorganic fertility, which is the result of inadequate mineralization of organic litter, a consequence of the cool, short alpine growing season (Rehder and Schäfer 1978; Gokceoglu and Rehder 1977; Rehder 1976a, 1976b; Fisk and Schmidt 1995; chapters 11, 12). Slow mineralization rates, in turn, result in a soil that is high in organic humus, and more likely than the soil of other ecosystems, to sequester and bind inorganic nutrients, especially N and P. Accordingly, alpine plants are exposed to a difficult situation in their efforts to obtain the inorganic ions required to support growth and reproduction. In accommodating the relative infertility of alpine soils, plants rely on a number of different traits, some of which are ubiquitous and some of which are more restricted in their distribution. Biomass allocation patterns favor high root:shoot ratios, increasing the potential for nutrient absorption by the roots relative to nutrient utilization by the shoot. Nutrient-use efficiencies (biomass produced per mass of senescent nutrient) tend to be high in alpine plants due to efficient resorption prior to leaf senescence. In several alpine growth forms, strict internal controls over seasonal phenology and growth (e.g., preformed buds and strongly enforced dormancy patterns) bring growth demands for nutrients more into balance with the limited supply provided by the soil. Luxury uptake and long-term storage during pulses of high nutrient availability provide plants with a means of bridging the gap between incongruent periods of high nutrient supply and high nutrient demand. Association of fungi with the roots of some alpine plants has the potential to enhance N and P acquisition. Finally, some alpine species can overcome the limitations imposed by scarce inorganic nutrient supplies through high rates of organic nutrient assimilation. It is the aim of this chapter to further consider each of these traits, with particular emphasis on their relationship to N and P acquisition. Topics concerning soil processes and their role in controlling nutrient availability have been covered elsewhere (chapter 8) and will not be repeated. Rather, this review focuses on nutrient relations from the plant’s perspective.

Primary Production

The production of biomass by plants is of central importance to energy, carbon, and nutrient fluxes in ecosystems. Knowledge of the spatial and temporal variation of production and the underlying biotic and physical controls on this variation are central themes in ecosystem science. The goals of this chapter are to present the estimates of spatial patterns in above- and belowground production associated with the major community types found on Niwot Ridge and other alpine areas of the southern Rocky Mountains and to examine the likely environmental causes and underlying mechanisms responsible for spatial and temporal variation in production as elucidated by experimental and observational studies. Rates of primary production and standing crops of plant biomass are low in alpine tundra relative to other ecosystem types (Lieth and Whittaker 1975; Zak et al. 1994). However, within communities (i.e., at the plot level), there is large variation in rates of production, the degree of biotic control over response to environmental change, and the principal environmental constraints of primary production. As a result, the alpine is one of the most dynamic ecosystems for research. For example, there is a tenfold difference in annual aboveground production between the most and least productive sites with continuous plant cover on Niwot Ridge. In addition, the high plant diversity is a source of potential variation in physiological and developmental control of plant response to the environment. Dominant species include sedges, grasses, shrubs, and forbs, among which are N2-fixing Trifolium species. Nearly all of the dominant species may be mycorrhizal. Soil moisture, a driving force for many biotic processes, may vary by an order of magnitude between wet and dry sites following prolonged periods of drought. Thus the alpine tundra of Niwot Ridge, which might appear superficially homogeneous, in fact has complex physical and biotic gradients. This spatial variation prevents simple generalizations about single limiting resources or climatic driving forces determining spatial and temporal variation in productivity. Billings (1973) defined the mesotopographic gradient as a working unit for describing the alpine landscape, as it encompasses the full range of snow accumulation and associated microclimates and thus biological diversity.

Vertebrates

Vertebrates of alpine tundra are near the limits of their genetic tolerance, and thus the alpine provides a natural laboratory for the study of the ecology of these organisms in a climatically stressful environment. The alpine supports a greater species richness of vertebrate herbivores than does arctic tundra (Halfpenny and Southwick 1982). Hoffmann (1974) provided an extensive review of terrestrial vertebrates of arctic and alpine ecosystems, emphasizing circumpolar patterns. For a variety of reasons, however, vertebrates of alpine tundra are considerably less studied than are those of the Arctic, and much remains to be learned about the physiological and behavioral adaptations of vertebrates that allow this group to exist in this extreme and variable ecosystem. May (1980) offered some generalizations about the state of knowledge of alpine animals. Terrestrial systems are better known than aquatic systems; the magnitude of environmental variability is better known than its predictability and significance to populations of animals; life histories of animals are better known than their roles and functions; dynamics of single species are better known than interactions between and among species; habitat selection by animals is more often defined in terms of the perception of the investigator than in terms of the perception of the organism; the response of animals to patterns of vegetation is better known than the influence animals have in creating and maintaining those patterns; and densities of animals are better known than are patterns of dispersion and their causes. Those generalizations remain broadly accurate. The purpose of this chapter is to develop a perspective on the structure and function of the vertebrate fauna of alpine environments of the Southern Rocky Mountains, with an emphasis on the fauna found on Niwot Ridge. It considers the origin and ongoing development of the fauna and its biogeographic and ecological relationships. A pattern of distributions is described that is dynamic in space and time. A principal focus is the role of vertebrates to the structure and function of the tundra ecosystem, including both the biotic and physical impacts of vertebrate populations. Some attention is paid to vertebrate trophic guilds, but plant-animal interactions are detailed in this volume by Dearing (chapter 14).

The Vegetation: Hierarchical Species-Environment Relationships

The vegetation of Niwot Ridge has a rich history of study, beginning with phytosociological studies directly on the Ridge and in the surrounding mountains and incorporating more experimental and dynamic approaches in later years. This chapter provides an overview of the spatial patterns of Niwot Ridge plants and plant communities relative to the primary controlling environmental gradients at scales from the individual to the landscape. The spatial patterns of vegetation at all scales are dominated by physical forces, particularly the interaction of wind, snow, and topography. The controls of biotic factors on the distribution and abundance of plant species on Niwot Ridge have received considerably less attention than have physical factors, but recent studies have revealed the importance of competition and certain mutualisms in structuring community composition. Community research on Niwot Ridge has been organized around a hierarchy of spatial scales, from the plot to the region. Plot-based studies have focused on physiological and ecological dynamics of specific species and communities, and more spatially extensive studies have provided a hierarchical framework for the plot studies. In this chapter, we first present an overview of the broader patterns in the vegetation, followed by descriptions of the communities, and then the specifics of physical and biotic controls on species and plant growth that drive the community patterns. The landscape-scale patterns in the Niwot vegetation are driven by a complex elevation gradient, which is a combination of temperature and snow regime, with wind modifying and interacting with temperature and snow at all points along the gradient (chapter 2). Certainly the most critical boundary in the system is the upper tree limit, which defines the alpine system and which lies roughly between 3400 and 3600 m elevation on Niwot Ridge. Billings (1988) provided a climatic-floristic-physiographic review of major North American alpine systems that helps to place Niwot Ridge into a larger perspective. Climatically, Niwot is intermediate between the dry Sierras, which have greater precipitation but almost none of it falling during the summer, and the wetter northern Appalachians (Mt. Washington), which have fairly even annual precipitation and no drought.

Soil-Atmosphere Gas Exchange

The alpine, while not extensive in global area, has several advantages for trace gas research, particularly the spatial landscape heterogeneity in soil types and plant communities. This variation can be viewed as a “natural experiment,” allowing field measurements under extremes of moisture and temperature. While the atmospheric carbon dioxide (CO2) record at Niwot Ridge extends back to 1968 (chapter 3), and NOAA has done extensive measurements on atmospheric chemistry at the subalpine climate station (e.g., Conway et al. 1994), work on tundra soil-atmosphere interactions were not initiated until recently. In 1992, studies were begun on Niwot Ridge to gain a comprehensive understanding of trace gas fluxes from alpine soils. Our sampling regime was designed to capture the spatial and temporal patterns of trace gas fluxes in the alpine. In addition, we coupled our studies of trace gas fluxes with ongoing studies of nitrogen cycling on Niwot Ridge (Fisk and Schmidt 1995,1996; Fisk et al. 1998; chapter 12). Methane (CH4), carbon dioxide (CO2), and nitrous oxide (N2O) were studied because of their role in global environmental change and because they could be easily monitored at our remote sites. On a per-molecule basis, CH4 and N2O are much more potent as greenhouse gases than CO2 is (Lashof and Ahuja 1990; Rodhe 1990). In addition, N2O plays a role in ozone depletion in the stratosphere. The global CH4 and N2O budgets are still poorly understood and the relative importance of soils in these budgets is even less clear. For example, estimates of the global soil sink for CH4 range from 9.0 to 55.9 Tg per year (Dörr et al. 1993). This range is large compared with the approximately 30 Tg of excess CH4 that is accumulating in the atmosphere every year. To better assess the role of soil in trace gas budgets, our work focused on investigating landscape patterns of gas fluxes (CH4, N2O, and CO2) and environmental controls on these fluxes.