Decomposition rates of native versus non-native litter in a northwest temperate forest Jane Q. Sample Hypothetical University, Planet Earth Introduction: Soils of the earth harbor much more carbon than is contained in the earth’s atmosphere (Davidson and Janssens 2006). Fallen litter provides pulses of nutrients for forest plants, soil invertebrates, and microbes and creates stocks of organic carbon in the soil. Current literature suggests that an increase in temperature of 1°C in areas with an annual mean temperature of 5°C will result in a loss of over 10% soil organic carbon. The amount of loss at an area with a mean annual temperature of 30°C will only be 3% (Kirschbaum 1995). Tropical forests, dominated by fleshy deciduous leaves, experience extreme spikes in available nutrients in the fall when its leaves senesce. These leaves decompose rather quickly, resulting in long periods of low nutrients during the spring and summer months. Conversely, our forests in the temperate Northwest are dominated by coniferous trees. As a result, our forests accumulate deep layers of duff through the shedding of needles, but don’t experience the nutrient spikes of the tropics. Climate change is expected to create a positive feedback in the global carbon cycle through decreases in soil organic carbon levels (Kirschbaum 1995). In this study our aim was to measure decomposition rates of native versus introduced leaf litter. Biological invasion theories assert that success rates of invasions are determined in part by the invader being released from natural predators. In our case, we consider these predators to include shredding insects and soil microbes that decompose leaf material in order to obtain the carbohydrates and sugars contained within. Because decomposition rates depend on the triangle of soil fauna/soil moisture/soil temperature, we also consider climate to be an important abiotic “predator”. We therefore predict that native leaf litter, remaining under pressure from their native predators, will decompose at a faster rate than introduced litter, released from the predators of its native range. Materials and Methods: Litter bags were constructed as recommended in the protocol provided by the Ecoplexity Project (http://ecoplexity.org/node/617). We randomly selected bags to receive treatments of Alder, live Douglas fir needles, and dead Douglas fir needles from the H. J. Andrews Research Forest in Oreong; Cecropea from the Luquillo Tropical Forest in Puerto Rico; grass from the Shortgrass Steppe in Colorado, mesquite from the Chihuahuan desert of New Mexico, and paper from an urban site in Pheonix, Arizona. All litter was dried to a constant dry weight at 10⁰C prior to placement in the bags. At the H. J. Andrews Research Forest in Blue River, Oregon, we constructed two strings of bags per litter type, five bags per string, with the bags six inches apart. We placed one string of each litter type above ground (Fig. 1), and one string of each just below the surface of the forest floor soil. Fourteen strings were constructed in this manner for a total of 70 bags. We allowed the litter bags to remain undisturbed from 2/1/2008 through 6/24/2008. At the termination of the experiment, we collected the bags and again weighed to a dry constant weight. Figure 1. Example of “above ground” litter bag filled with leaf litter. (Image courtesy of S. Kirkpatrick) Results: While the bags of paper placed above and below ground experienced equal weight loss, all others placed below the soil surface lost a greater proportion of their starting weights (Fig. 2). The bags containing paper lost the greatest proportion of weight and the Cecropea samples lost the least amount (Table 1). All non-native litter samples, with the exception of Cecropea, experienced the greatest amount of mass lost during the experiment (Fig. 3). Mean Weight Loss Percentage 0.60 0.50 0.40 0.30 0.20 0.10 0.00 Alder Live Doug Dead Cecropea fir Doug fir Grass Mesquite Paper Figure 2. Percentage of beginning weight lost during experiment. Values are the collective mean of the five bags above ground and five bags below ground for each litter type. Species Alder Live Doug fir Dead Doug fir Cecropea Grass Mesquite Paper Above Donor Ground Region Loss (%) HJA 12 HJA 32 HJA 12 LUQ 0 COL 15 NM 41 AZ 52 Below Ground Loss (%) 23 34 15 5 25 47 52 Table 1. Mean percentage of weight lost in litter bags above and below ground level. Donor region codes are as follows: HJA = H. J. Andrews Research Forest, Temperate Rain Forest; LUQ = Luquillo Research Forest, Puerto Rico, Tropical Rain Forest; COL = Colorado, Shortgrass Steppe; NM = New Mexico, Desert; and AZ = Arizona, Urban. Percent Weight Loss‐ Plants Native to H. J. Andrews 0.60 0.50 0.40 0.30 above ground 0.20 below ground 0.10 0.00 Alder Live Doug fir Dead Doug fir Percent Weight Loss‐ Plants Not Native to H. J. Andrews 0.60 0.50 0.40 0.30 above ground 0.20 below ground 0.10 0.00 Cecropea Grass Mesquite Paper Figure 3. Weight loss relative to litter type and nativity. Because initial weights varied slightly from species to species, proportion of weight loss is reported here. Discussion: Litter decomposition rates are determined by the soil fauna present, soil moisture content, and soil temperatures. It was our expectation that due to having its native predators present, native leaf litter would decompose at a faster rate than non-native litter. Contrary to our expectation, however, all but one non-native litter type lost a greater proportion of their starting weights than did native litters. There are a number of possible explanations for this finding, only a couple of which we will discuss here. First, Douglas fir needles have thick, waxy cuticles that prevent the plant from desiccating during the hot and dry summers that are typical at the H. J. Andrews Forest. By design, this cuticle is slow to break down. Cecropea, grass, Mesquite, and paper do not have waxy surfaces and would therefore decompose at faster rates. This does not explain the unexpected results of the alder sample, as alder is a deciduous tree without a thick leaf cuticle. It is important to note, however, that alder is a nitrogen-fixing species. While this may at first appear irrelevant, we believe this to be a very real explanation of why the alder did not lose as much mass as was expected. Alders indirectly benefit Douglas firs in that the alder fixes nitrogen which feeds the native mycorrhizal fungus that dramatically increases the Douglas firs’ root surface area, allowing increased nutrient and water uptake. It might very well be the case that the alder litter itself decomposed more than appears and the hyphae of the mycorrhizal fungus is masking that fact by contributing to the final weight. As discussed above, Micorrhizae obtain sugars they need in part by developing mutualistic relationships with deciduous trees in our temperate forests, chiefly alder trees. A second possible explanation for the unexpected results involves the fact that the introduced types of litter, with the exception of the paper, are deciduous. Leaves of deciduous plants are typically fleshier and contain higher levels of sugar than do coniferous leaves. The Micorrhizae may show a preference for the fleshy introduced litter, extracting the sugars and carbohydrates and dramatically decreasing their masses. Future studies that test native deciduous against native coniferous litter are needed to investigate this hypothesis further. The final litter mass may not be fully representative of processes in the field due to the construction of the litter bags themselves. As is typical in leaf litter decomposition studies, the underside of the bags consisted of fine mesh so as to not lose litter material out of the bottom of the bag. While not as fine as the underside, the top of the bags were also made of fairly fine mesh, most likely to the exclusion of shredding insects. During the decomposition process, it is these shredding insects that remove the greatest proportion of leaf material, either using it for themselves or breaking the larger litter into more usable units for other detritivores. We recommend similar experiments be conducted that assign bag mesh size as a treatment level in order to elucidate the effects on variability. Due to climate change we are seeing plants growing at higher elevations and in habitats once considered novel. It is not unreasonable to assume that over time, our temperate coniferous forests will contain a greater number of deciduous trees introduced into the system as a result. Studies such as this are important in understanding the effects this will have on decomposition rates, and therefore the organic carbon supply, of our forests. This study should be viewed as only a starting point upon which to build. References: Davidson, Eric A. and Ivan A. Janssens. 2006. Temperature sensitivity of soil carbon decomposition and feedbacks to climate change. Nature 440: 165-173. Miko U. F. Kirschbaum. 1995. The temperature dependence of soil organic matter decomposition, and the effect of global warming on soil organic C storage. Soil Biology and Biochemistry 27(6): 753-760.
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