Thesis (M.S.)--Oregon State University, 2007. / Printout. Includes bibliographical references (leaves 42-54). Also available on the World Wide Web.

Fire ecology encompasses the study of the factors, biotic and abiotic, that influence the occurrence of fire in an area, as well as the effects fire has on the flora and fauna native and non-native to the region (Whelan 1995). Fire has had a major influence on shaping biomes as we see them today. Fire has had an effect on vegetation much before the evolution of Homo on Earth (Keeley and Rundel 2005, Pausas and Keeley 2009, Midgley and Bond 2011). With the evolution and expansion of Homo across Earth, fire has been tamed, and then generated and used over time to yield landscapes that were suitable for their existence (Pyne 1991, Bowman et al. 2009, Archibald et al. 2012). Thus, fire, vegetation and humans were, and still are, inextricably linked in certain biomes on Earth. The best examples are observed in tropical savannas and grasslands, biomes that experience distinct seasonality in climate and are thus prone to frequent fires caused either by lightning or by humans (Keeley and Rundel 2005, Archibald et al. 2012). At the other end of the spectrum of tropical vegetation types are rainforests where the occurrence of fires is constrained by a perpetually moist environment (Meyn et al. 2007, van der Werf et al. 2008), in the absence of manipulation of the forest landscape by humans. Frequent fires have been documented to alter structure and cause a decline in forest diversity in rainforests (Cochrane and Schulze 1999, Cochrane 2003), whereas fire exclusion in mesic savannas leads to increases in biomass and transition to forest ecosystems (Bond et al. 2003, Bond et al. 2005 and references therein). A tropical biome that lies between these two extremes of vegetation types is the Seasonally Dry Tropical Forest (SDTF) where the occurrence of fire is common, but for which there are contrasting views on the effect of fire on this system (Saha and Howe 2003, Otterstrom et al. 2006 as examples). Current forest management policies in SDTF areas, especially in India, actively aim to exclude fire from these forests mostly because of the perception held by forest managers and the general public that fire has negative effects on forests. However, very few scientific studies have explored the ecology of fire in SDTFs. In order to formulate fire management policies, it is necessary to have a more comprehensive understanding of the ecology of fire in this tropical forest type. This thesis addresses two components of fire ecology as applied to SDTFs. The first is how fire is influenced by the environment, and the second, how fires influence the biotic community particular to SDTFs. The study was carried out in an SDTF in southern India where fire is a common occurrence -the forests of Mudumalai – a protected area that exhibits a range of SDTF vegetation types, from moist deciduous to dry thorn forest, corresponding to a rainfall gradient. Fire influenced by the environment: For this section, the influence of fuel load, fuel moisture and ambient weather on area burnt, fire occurrence and fire temperatures were studied in the SDTF vegetation types of Mudumalai. The extent of fire (area burnt) in an ecosystem differs according to the relative contribution of fuel load and fuel moisture available (Meyn et al. 2007). At a global scale, these factors vary along a spatial gradient of climatic conditions and are thus “varying constraints” (Krawchuk and Moritz 2011) on fire activity in natural ecosystems (Meyn et al. 2007, Krawchuk and Moritz 2011). Moist ecosystems such as tropical rainforests are at one end of the spectrum where fire activity is constrained by fuel moisture. At the other end are arid ecosystems, such as deserts, where fire activity is limited by the presence of fuels. The potential for the globally widespread seasonally dry tropical forests (SDTFs) to be placed as a single entity in this framework was examined by analyzing the interacting effects of fuel load and fuel moisture on the extent of fire in Mudumalai. Logistic regression was used to model proportion area burnt in a given year with factors that would influence fuel load and fuel moisture – these were proportion area burnt the previous year, wet season rainfall the previous year and early dry season rainfall. Modelling was conducted at two levels – the overall landscape and within four defined moisture regimes (between 700 and 1700 mm yr-1) – using a dataset of area burnt and seasonal rainfall from 1990 to 2010. The landscape scale model showed that the extent of fire in a given year within this SDTF is dependent on the combined interaction of seasonal rainfall and extent burnt the previous year. However, within individual moisture regimes the relative contribution of these factors to the annual extent burnt varied – early dry season rainfall (i.e. a moderator of fuel moisture) was the predominant factor in the wettest regime, while the previous year’s wet season rainfall (i.e. a proxy for fuel load) had a large influence on fire extent in the driest regime. Thus, the diverse structural vegetation types associated with SDTFs across a wide range of rainfall regimes would have to be examined at finer regional or local scales to understand the specific environmental drivers of fire. While the extent burnt in SDTFs is largely dependent on climatic influences, the probability of ignition has not been characterized for SDTFs. Anthropogenic fires are a regular occurrence during the dry season in SDTFs (Stott et al. 1990). We investigated if the occurrences of anthropogenic fire in Mudumalai were associated with any particular weather conditions during the dry season. Logistic regression between probability of a fire day and weather variables -seasonal rainfall, ambient relative humidity and temperature -was examined during the dry seasons of 20042010 in Mudumalai. Fire incidence data was obtained from the Fire Information for Resource Management System (FIRMS; NASA 2002) and weather data from two automatic weather stations within Mudumalai. The analysis showed that days with high probabilities of fire occurrence were associated with low levels of early dry season rainfall, low daily average relative humidity, and high daily average temperatures. These weather conditions are known to influence moisture levels of fine fuels (Viney 1991, Archibald et al. 2009). In Mudumalai as well as other SDTFs the primary fuels for fires are fine fuels such as litter and dried grass that accumulate on the forest floor during the dry season. Our results suggest that the occurrence of fire is moderated by environmental conditions that reduce or enhance the flammability of fine fuels in the dry tropics. A quantitative framework for assessing risk of a fire day has been proposed as an outcome of this analysis to assist forest managers in anticipating fire occurrences in this SDTF, and possibly for those across south Asia. Of the various components of a fire regime, fire intensity is an important aspect. High fire temperatures (one measure of fire intensity, Keeley 2009) and resulting soil temperatures would have an effect on soil properties as well as plant species demography and community structure (Moreno and Oechel 1991, Neary et al. 1999, Morrison 2002). Fires that occur frequently in a region could vary in their intensity and severity depending upon the amount of fuel available and ambient weather conditions (Stinson and Wright 1969, Stott 1986, Stronach and McNaughton 1989, Ansley et al. 1998, Wotton et al. 2012). However, this relationship has not been examined in a multiple regression framework for SDTFs. Fire temperature was recorded and its relationship with ambient weather and fuel load was studied in two SDTFs of southern India -Mudumalai and Biligiri Rangaswamy Temple (BRT) Wildlife Sanctuary in Karnataka. During “controlled burns” conducted by the forest department staff in these reserves in February and March 2010, temperature indicating lacquers on mica sheets were used to measure fire temperature at several points at ground level and one cm below the ground. Biomass was harvested close to the temperature measurement points to estimate fuel load and fuel moisture. Ambient weather conditions were recorded during the controlled burn when the flame passed over the indicators. Temperatures recorded at ground level ranged from <79oC to 760oC, with the most frequently recorded temperatures between 343-399 oC and 510566 oC. Temperatures measured one cm below the ground ranged from <79oC to 302oC, with a majority of the indicators recording temperatures in the <79oC category. Ground-level temperatures increased with increasing biomass. A linear regression of ground-level temperatures with fuel load and ambient weather conditions of relative humidity and temperature was found to explain most of the variation in the data. Ground-level fire temperatures increased with increasing fuel load, but were also found to be lower at higher relative humidities at a given temperature. In order to reduce the intensity of forest fires that occur accidentally during the dry season, we recommend that fuel loads be reduced in the forest by prescribed burning early in the dry season. This applies especially to areas where there is accumulation of biomass over years, such as that of the tall grass Themeda cymbaria found predominantly in dry deciduous forest types. If prescribed burning is incorporated in fire management policies for these forests, then the season of burning will be important to consider. It is known from ecosystems where prescribed burning is regularly applied that early dry season fires are less intense than late dry season fires (Williams et al. 1998). However, this has not been systematically investigated for SDTFs. Through a burning experiment carried out in private land with vegetation type similar to tropical dry thorn forest, we investigated differences in area burnt, ground-level fire temperatures and soil temperatures one cm below the ground in the early dry season in January, late dry season in April and and early wet season in June. We also examined differences in fuel load, fuel moisture, soil moisture and weather conditions of ambient relative humidity (RH), temperature and wind speed in these phases; these factors could be responsible for observed differences in fire and soil temperatures or area burnt. Although area burnt was not significantly different between the early and late phases of the dry season, fire and soil temperatures were significantly lower in the former. The late dry season was characterized by distinctly higher fuel loads, lower fuel moisture, lower relative humidity, higher ambient temperatures and higher wind speeds compared that measured in the early dry season. Differences in soil temperature between these months may be attributed to the increase in fuel load since there were no significant differences in soil moisture. Fire spread was limited in the experimental plots in the early wet season in June, probably due to significantly higher levels of fuel moisture in this month; the resultant fire and soil temperatures recorded were low. Forest management should, therefore, consider early dry season burns in the month of January for prescribed burns in the sanctuary, although this would have to be tested in other SDTF vegetation types with more variable fuel load, fuel moisture and weather conditions. Fire’s influence on the biotic community: Concerns regarding the regeneration capacity of woody species in SDTFs have been voiced with respect to increasing frequencies of fire (Saha and Howe 2006, Kodandapani et al. 2008). Fire is known to cause high mortality of individuals of small size (Swaine et al. 1990, Suresh et al. 2010). However, mortality has been examined for large size classes, and not for seedlings. It is essential to understand the dynamics of seedlings and their contribution to the regeneration potential of SDTFs. Woody species in SDTFs are known to have traits that help them recover from recurring disturbances, such as sprouting from underground root stocks (Vieira and Scariot 2006). Another trait may relate to growth rates of seedlings. Growth rates of seedlings (defined in this study as established individuals between 10 and 100cm height) after dry season (February-March) fires were compared between adjacent pairs of burnt and unburnt transects established at eight sites in Mudumalai across vegetation types of moist deciduous, dry deciduous and dry thorn forest. The growth of grasses, a possible competitor for resources, was also monitored at each site. Seedling and grass heights were monitored at 3-month intervals between August 2009 and August 2010. A second fire in March 2010 affected transects at two sites in Mudumalai. Seedling and grass heights were monitored for two enumerations till August 2010 subsequent to the second fire at these two sites. A total of 1032 individuals across 58 woody species were enumerated. High seedling survivorship (>95%) was observed in both burnt and unburnt areas. Although seedling heights were significantly different between burnt and unburnt areas at the start of the enumeration in August 2009, heights were comparable within a year and a half of the fire. Comparable seedling heights in such a short time span were because of distinctly higher growth rates of seedlings in burnt areas compared to unburnt areas after the fire event, particularly during the pre-monsoon season. Grass biomass (volume), on the other hand, was significantly different between burnt and unburnt areas at both the first and last enumerations. Grass growth (change in volume) did not differ between burnt and unburnt areas. Rapid growth by seedlings after a fire implies adaptation through the use of stored resources for growth, possibly aided by lower competition from grasses, in order to attain a certain size before the subsequent return of unfavourable factors such as a recurrent fire event. Conclusions: The results from the study point to climatically driven fire regimes in an SDTF in southern India, with daily influences of weather conditions during the dry season on fire occurrences. Fire intensities increase with increasing fuel loads in these forests, moderated by weather conditions such as RH and temperature. Since fires are an anthropogenic phenomenon in these forests, active management with the use of prescribed fires in the early phase of the dry season is a possible option to control late dry season fires that would be higher in intensity. The current woody tree species assemblage in this southern Indian SDTF is resilient to fires at the seedling stage, with established individuals exhibiting high survivorship and rapid growth after a fire. However, the effects of fires of varying intensities on the regenerative capacity of the seedlings are not known. The effect of fire on habitat utilisation by large herbivores, or the impact of fire on the faunal community in general has not been studied for vegetation types that comprise SDTFs. The effect of fire exclusion on the ecology of SDTFs will provide useful information that can feed into management policies for this ecosystem type. These are potential areas of research for the future. Fire, if managed wisely, can be an effective tool for the conservation of SDTFs across south and southeast Asia.

Increases in temperature, altered precipitation patterns, and the occurrence and severity of extreme climatic events have been important characteristics of the climate change observed to date. This has had many and diverse impacts upon the living world, with one recent observation being a global reduction in the net primary production of all terrestrial vegetation. Increases in temperature and the frequency of extreme events are predicted to continue throughout the 21st century, and can be expected to have far reaching effects on global terrestrial ecosystems. Increases in temperature and drought occurrence could fundamentally impact upon the growth rates, species composition and biogeography of forests in many regions of the world, with many studies indicating that this process is already underway. European beech, Fagus sylvatica, is one of Europe’s most widespread and significant broadleaved tree species, forming an important and frequently dominant component of around 17 million hectares of forest. However, the species is also considered to be drought sensitive. Thus, much research interest has focused on eliciting the details of its physiological response to increased water stress, whilst dendroecological studies have attempted to identify sites and regions where reductions in growth might be found. A significant knowledge gap exists regarding a multi-regional, range-wide view of growth trends, growth variability, climate sensitivity, and drought response for the species. Predicting the potential effects of climate change on competition and species composition in mixed species forests remains an important challenge. In order to address this knowledge gap, a multi-regional tree-ring network was constructed comprising of 46 sites in a latitudinal transect across the species’ Western European range. This consisted of 2719 tree cores taken from 1398 individual trees, which were used to construct tree-ring chronologies for each site in the network. As a first step in a multi-regional assessment for F. sylvatica, a combination of the tree-ring chronologies and environmental data derived from a large scale gridded climate dataset were used in a multivariate analysis. Sites in the latitudinal transect were partitioned into geographically meaningful regions for further analysis. The resulting regions were then studied using climate-growth analysis, pointer year analysis of drought years, analysis of growth trends and growth variability, in order to examine regional variation in the response of the species to climate. Furthermore, a combination of long-term monitoring data from one specific site was combined with tree-ring sampling of multiple cohorts of F. sylvatica and one co-dominant competitor, Quercus petraea, to study the effects of an extreme drought event in 1976 on mortality and subsequent recovery. Key results of the multi-regional analysis are that large scale growth reductions are not evident in even the most southerly and driest portions of the species’ range. Radial growth is increasing, both in the north and in the core of the species’ range, with southern range edge forests maintaining stable growth. However, the variability of growth from year to year is increasing for all regions, indicative of growing stress. Crucially, the southern range edge, which previous studies had identified as an ‘at risk’ region, was shown to be more robust than expected. Climate sensitivity and drought impacts were low for this region. Instead, forests in the core of the species range, both in continental Europe and in the south of the UK, were identified as having the highest climate sensitivity, highest drought impacts, and experiencing periodic reductions in growth as a result. Northern range edge forests showed little sign of being affected by drought, instead having low climate sensitivity and strongly increasing growth trends. Extreme drought was found to affect species differently: the dominant species (F. sylvatica) failed to recover pre-drought levels of growth, whilst a transient effect of competitive release occurred for the co-dominant species (Q. petraea). There was also a long term effect on the relative abundance of the two species within the woodland, due to differences in the levels of drought induced mortality experienced by the species. This shows that in the case of extreme climatic events where thresholds in the ability of species to tolerate water stress are breached, the effects of drought can be rapid and long lasting. Drought impacts can cascade beyond that experienced by the most drought sensitive species, due to changes in competitive interactions between species in mixed species forests. The implications of this work suggest opportunities, risks and strengths for F. sylvatica. In the northern portion of the species’ range, predicted increases in productivity are confirmed by recent growth trends, indicating a good outlook for the species. At the southern range edge, F. sylvatica forests exist either in locations where precipitation is high or locations where local environmental conditions buffer them from an inhospitable regional climate. These factors result in southern range edge forests which are highly resilient to the effects of increasing climate stress. It is instead in the core of the species’ range where the most sensitive forests are found. The effects of extreme drought on a range core forest demonstrated here provide a cautionary note: where drought stress tolerance thresholds are breached, rapid and long lasting effects on growth and mortality can occur, even in regions where drought has not previously been considered to pose a strong risk to the species.

This thesis examines factors limiting understory herb presence and flowering in young second-growth Douglas-fir (Pseudotsuga menziesii) forests on the west side of the Cascade Mountains, Oregon, USA. I studied the belowground effects of canopy trees on understory herbs and shrubs in old-growth forests using trenched plots from which tree roots were excluded. Effects of tree density and stand age were tested by comparing the understory community composition of old-growth stands and pre-commercially thinned and unthinned young second-growth stands. I also examined the effect of conifer basal area on understory herb presence and flowering within one young second-growth watershed. In young stands, I focused on three groups of understory herb species: disturbance-responsive (release), forest generalist and old-growth associated. The effects of root trenching on vegetation and soil moisture were tested in closed-canopy and gap locations in two old-growth Douglas-fir (Pseudotsuga menziesii) forests. Ten years after installation, trenched plots averaged 92% total understory cover while untrenched plots averaged 47% cover. Trenched plots under closed canopies were moister than control plots throughout the growing season; the trenching effect on soil moisture became apparent in the generally wetter gaps only at the end of the growing season. Vegetation responses to trenching were concomitantly larger under closed canopies than in gaps. Stands that had been pre-commercially thinned 20 years earlier exhibited understory composition more similar to old growth than did unthinned stands. Thinned stands exhibited higher frequencies, abundances and density of flowering of old-growth associated herbs than did unthinned stands, but lower than did old-growth stands. Forest generalist and release species showed mixed responses to thinning. I used both general linear models and classification and regression tree models to explore the association of herb species presence and flowering with conifer basal area and abiotic variables. Both modeling approaches yielded similar biological insights. Flowering was more sensitive than presence to current stand basal area. Flowering of old-growth associated and release species was negatively correlated with conifer basal area. Linear models allowed clearer hypothesis tests, while tree-based models had greater explanatory power and provided information about interactions between variables. / Graduation date: 2004

One of the human activities impacting biodiversity is the cutting of old-growth forests. In response to the controversy surrounding the cutting of old-growth in the coastal rainforest of BC, the Ministries of the Environment and Forests have produced biodiversity guidelines that are to be applied when manipulating stands in the provincial forest. This study augments these guidelines by investigating the diversity differences between second-growth and old-growth forests in relation to site quality. We demonstrate how standlevel plant diversity differs between 40-year-old and old-growth stands in the Very Wet Coastal Western Hemlock subzone (CWHvm) on Vancouver Island. This information is intended to provide foresters with an understanding of the effects of age, disturbance and site quality on stand-level plant diversity, thereby allowing for informed professional management decisions.

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Thesis (Ph. D.)--Ohio State University, 2007. / Title from first page of PDF file. Includes bibliographical references (p. 160-168).

Expanding anthropogenic development within the tropical forest biome is driving the loss of an irreplaceable global resource. Mega-diverse tropical forests are vital for regulating the global carbon cycle, and are essential for climate change mitigation. Today, over half of the world’s remaining tropical forest is degraded or regenerating secondary forest. Tropical forests are becoming increasingly fragmented through the expansion of agriculture and roads. Landscape-scale flooding of terrestrial habitats caused by dam construction is an emerging driver of habitat loss and fragmentation. Much attention has been paid to the long-term impacts of tropical forest fragmentation for biodiversity, ecosystem functioning, and carbon emissions. Most of our understanding of the impacts associated with habitat fragmentation originates from systems in which the habitat matrix surrounding remnant forest patches is another, albeit low quality, terrestrial habitat. However, dam-induced habitat fragmentation results in remnant terrestrial biological communities becoming isolated on islands within a water matrix. A water matrix presents the worst-case scenario for remnant habitat fragments. In Chapter 2 I synthesise the results of numerous studies reporting the responses of taxonomic groups to isolation on reservoir land-bridge islands, and uncover a globally-applicable pattern of extinction debt acting upon remnant biological communities on reservoir islands. All islands, regardless of taxonomic group, habitat type, or island area lose species as island isolation time increases. Moreover, I show that contrary to existing ecological theory, once terrestrial habitat becomes isolated within a water matrix, it is effectively too isolated for species losses to be buffered by metapopulation dynamics. Dam development is rapidly expanding in the largest remaining tract of intact tropical forest, the Amazon Basin. In Chapters 3 and 4 I study the Balbina mega-dam system in the central Brazilian Amazon. Here, I use detailed field inventories of trees and lianas on islands and in continuous mainland habitat to determine the impact of landscape-scale habitat fragmentation caused by reservoir creation on these taxonomic groups. I find that islands maintain tree communities at significantly lower densities, richness and diversity compared to continuous forest. Furthermore, tree communities on islands exhibit compositional divergence from those found in mainland continuous forest. Island tree assemblages are dominated by low-wood density species, and may be on a trajectory towards communities characteristic of early successional forests with reduced carbon storage capacity. In contrast, liana assemblages remain compositionally intact and are becoming increasingly dominant relative to trees. Thus, lianas appear robust to many of the negative impacts associated with landscape-scale habitat fragmentation. As insular tree communities continue to degrade through area- and edge-effects, lianas may become a key feature of this archipelagic landscape due to their competitive advantage over trees in disturbed forest habitats. Lianas significantly inhibit tree recruitment and carbon storage. Thus, findings from Chapters 3 and 4 provide strong evidence for additional, and currently unaccounted-for biodiversity and carbon impacts associated with tropical dams. As development of tropical forest regions increases, there is an urgent need to reconcile the need for resources with the need for ecosystem service provision, such as carbon storage, particularly as we attempt to mitigate the impacts of rising atmospheric carbon. Recent studies have shown that secondary tropical forests have the potential to rapidly uptake atmospheric carbon, and act as a powerful tool in climate change mitigation policy. Broad-scale estimates of secondary forest carbon uptake are currently based on above-ground biomass alone. In Chapter 5 I present carbon stock estimates of additional tropical forest carbon pools - soil and dead woody biomass - in secondary forests ranging from 40-120 years. I find that soil fertility (nitrogen concentration) is key in determining carbon storage in secondary forests, and that the stability of carbon stocks held in dead woody biomass increases with secondary forest stand age. I highlight the need to integrate detailed site-specific information into broad-scale predictive models of secondary tropical forest carbon sequestration. This thesis links ecological theory and landscape-scale field inventories, to provide new understanding of the long-term costs of tropical forest fragmentation for biodiversity conservation and carbon storage, and provides further evidence of the important role secondary tropical forests may play in carbon sequestration and climate change mitigation.

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No description available.