Headwater Basins Research Articles

Modeling increases in snowmelt yield and desynchronization resulting from forest gap‐thinning treatments in a northern mountain headwater basin

Key Points Forest thinning treatments in mountain regions may substantially alter snowmelt Changes in snowmelt timing from forest thinning depend on slope and aspect. Changes in snowmelt timing are caused primarily by shifts in radiation to snow.

Watershed-scale controls on snow accumulation in a small montane watershed, southwestern Alberta, Canada

Snow accumulation in mountain headwater basins is a major water source, particularly in semi-arid environments such as southern Alberta where water resources are stressed and snowmelt supplies more than 80% of downstream runoff. Relationships between landscape predictor variables and snow water equivalent (SWE) were quantified by combining field and LiDar measurements with classification and regression tree analysis over two winter seasons (2010 and 2011) in a small, montane watershed. 2010 was a below average snow accumulation year, while 2011 was well above normal. In both the field and regression tree data, elevation was the dominant control on snow distribution in both years, although snow distribution was driven by melt processes in 2010 and accumulation processes in 2011. The importance of solar radiation and wind exposure was represented in the regression trees in both years. The regression trees also noted the lower importance of canopy closure, slope, and aspect, which was not observed in the field data. This technique could provide an additional method of forecasting annual water supply from melting snow. However, further research is required to address the lack of data collected above treeline, to provide a full-basin estimate of SWE. Copyright © 2012 John Wiley & Sons, Ltd.

Hydrologic sensitivity of the Upper San Joaquin River Watershed in California to climate change scenarios

To examine the hydrological system sensitivity of the southern Sierra Nevada Mountains of California to climate change scenarios (CCS), five headwater basins in the snow-dominated Upper San Joaquin River Watershed (USJRW) were selected for hydrologic simulations using the Hydrological Simulation Program-Fortran (HSPF) model. A pre-specified set of CCS as projected by the Intergovernmental Panel on Climate Change (IPCC) were adopted as inputs for the hydrologic analysis. These scenarios include temperature increases between 1.5 and 4.5 °C and precipitation variation between 80 and 120% of the baseline conditions. The HSPF model was calibrated and validated with measured historical data. It was then used to simulate the hydrologic responses of the watershed to the projected CCS. Results indicate that the streamflow of USJRW is sensitive to the projected climate change. The total volume of annual streamflow would vary between −41 and +16% compared to the baseline years (1970–1990). Even if the precipitation remains unchanged, the total annual flow would still decrease by 8–23% due to temperature increases. A larger portion of the streamflow would occur earlier in the water year by 15–46 days due to the temperature increases, causing higher seasonal variability of streamflow.

Vegetation activity monitoring as an indicator of eco-hydrological impacts of extreme events in the southeastern USA

A vegetation disturbance persistence metric based on Moderate-Resolution Imaging Spectroradiometer North American Carbon Program vegetation indices (8-day composite at 500 m spatial resolution) was developed with the objective of assessing the eco-hydrological impact of extreme events generally, and hurricanes and tropical cyclones in particular, in the southeastern USA. The metric is based on the relationship between vegetation stress and the persistence of standardized enhanced vegetation index anomalies. The data show that in the first water year (October–September) after hurricane landfall, pronounced decreases in chlorophyll activity are found predominantly at very low elevations – including coastal marshes, wetlands, and the floodplains aligned with drainage networks of major river systems along the storm track. Inland impacts are mainly located in agricultural areas and woody wetlands in the floodplains, and in forested headwater basins in the mountains. Overall, the data analysis indicates that the major impacts of these storms are on agricultural and natural land-margin ecosystems predominantly via flooding associated with the storm surge along the coast, and flood propagation through the river networks inland. The timescale of impacts in coastal areas is up to two years, consistent with persistent impacts of salt intrusion in shallow coastal aquifers, erosion of root zone soils, and depletion of nutrients due to flooding. By contrast, ecological drought disturbance patterns exhibit larger spatial persistence but significantly shorter timescales (3–4 months).

Spatial variability in the flood vulnerability of urban areas in the headwater basins of Slovakia

AbstractIn the new paradigm of flood defence, the control of flooding by technical means is supported by an emphasis on the reduction of vulnerability to flooding by economic, social and environmental means. The aim of the present work was to assess the vulnerability of basic urban units (BUUs) located in the headwater basins of Slovakia. Vulnerability was expressed in terms of proxy variables. Using a combination of these variables, specific indices were established. These indices represented the potential for susceptibility to damage, resistance to flooding and resilience after flooding of the social and economic systems in these BUUs. The assessment of vulnerability was based on an application of the multi‐atribute decision‐making approach. The overall vulnerability of a BUU was calculated via a summation of the specific vulnerability indices. Using visualisation in GIS, the BUUs were grouped into three classes, of low, medium and high vulnerability. Second, evaluation of BUUs from the point of view of expected negative consequences of flooding was done.

Development of a stream–aquifer numerical flow model to assess river water management under water scarcity in a Mediterranean basin

Stream flow, as a part of a basin hydrological cycle, will be sensible to water scarcity as a result of climate change. Stream vulnerability should then be evaluated as a key component of the basin water budget. Numerical flow modeling has been applied to an alluvial formation in a small mountain basin to evaluate the stream–aquifer relationship under these future scenarios. The Arbúcies River basin (116km2) is located in the Catalan Inner Basins (NE Spain) and its lower reach, which is related to an alluvial aquifer, usually becomes dry during the summer period. This study seeks to determine the origin of such discharge losses whether from natural stream leakage and/or induced capture due to groundwater withdrawal. Our goal is also investigating how discharge variations from the basin headwaters, representing potential effects of climate change, may affect stream flow, aquifer recharge, and finally environmental preservation and human supply.A numerical flow model of the alluvial aquifer, based on MODFLOW and especially in the STREAM routine, reproduced the flow system after the usual calibration. Results indicate that, in the average, stream flow provides more than 50% of the water inputs to the alluvial aquifer, being responsible for the amount of stored water resources and for satisfying groundwater exploitation for human needs. Detailed simulations using daily time-steps permit setting threshold values for the stream flow entering at the beginning of the studied area so surface discharge is maintained along the whole watercourse and ecological flow requirements are satisfied as well. The effects of predicted rainfall and temperature variations on the Arbúcies River alluvial aquifer water balance are also discussed from the outcomes of the simulations. Finally, model results indicate the relevance of headwater discharge management under future climate scenarios to preserve downstream hydrological processes. They also point out that small mountain basins could be self-sufficient units so long as the response of the main hydrological components to external forces that produce water scarcity, as climate change or human pressures, is appropriately considered in water resource planning.

Aquatic insect assemblages associated with subalpine stream segment types in relict glaciated headwaters

Abstract Aquatic habitats and biotic assemblages in subalpine headwaters are sensitive to climate and human impacts. Understanding biotic responses to such perturbations and the contribution of high‐elevation headwaters to riverine biodiversity requires the assessment of assemblage composition among habitat types. We compared aquatic insect assemblages among headwater stream segment types in relict glaciated subalpine basins in Mt. Rainier National Park, Washington, USA. Aquatic insects were collected during summer and autumn in three headwater basins. In each basin, three different stream segment types were sampled: colluvial groundwater sources, alluvial lake inlets, and cascade‐bedrock lake outlets. Ward's hierarchical cluster analysis revealed high β diversity in aquatic insect assemblages, and non‐metric multidimensional scaling indicated that spatial and temporal patterns in assemblage composition differed among headwater stream segment types. Aquatic insect assemblages showed more fidelity to stream segment types than to individual basins, and the principal environmental variables associated with assemblage structure were temperature and substrate. Indicator species analyses identified specific aquatic insects associated with each stream segment type. Several rare and potentially endemic aquatic insect taxa were present, including the recently described species, Lednia borealis (Baumann and Kondratieff). Our results indicate that aquatic insect assemblages in relict glaciated subalpine headwaters were strongly differentiated among stream segment types. These results illustrate the contribution of headwaters to riverine biodiversity and emphasise the importance of these habitats for monitoring biotic responses to climate change. Monitoring biotic assemblages in high‐elevation headwaters is needed to prevent the potential loss of unique and sensitive biota.

Evaluation of SNODAS snow depth and snow water equivalent estimates for the Colorado Rocky Mountains, USA

AbstractThe National Weather Service's Snow Data Assimilation (SNODAS) program provides daily, gridded estimates of snow depth, snow water equivalent (SWE), and related snow parameters at a 1‐km2 resolution for the conterminous USA. In this study, SNODAS snow depth and SWE estimates were compared with independent, ground‐based snow survey data in the Colorado Rocky Mountains to assess SNODAS accuracy at the 1‐km2 scale. Accuracy also was evaluated at the basin scale by comparing SNODAS model output to snowmelt runoff in 31 headwater basins with US Geological Survey stream gauges. Results from the snow surveys indicated that SNODAS performed well in forested areas, explaining 72% of the variance in snow depths and 77% of the variance in SWE. However, SNODAS showed poor agreement with measurements in alpine areas, explaining 16% of the variance in snow depth and 30% of the variance in SWE. At the basin scale, snowmelt runoff was moderately correlated (R2 = 0.52) with SNODAS model estimates. A simple method for adjusting SNODAS SWE estimates in alpine areas was developed that uses relations between prevailing wind direction, terrain, and vegetation to account for wind redistribution of snow in alpine terrain. The adjustments substantially improved agreement between measurements and SNODAS estimates, with the R2 of measured SWE values against SNODAS SWE estimates increasing from 0.42 to 0.63 and the root mean square error decreasing from 12 to 6 cm. Results from this study indicate that SNODAS can provide reliable data for input to moderate‐scale to large‐scale hydrologic models, which are essential for creating accurate runoff forecasts. Refinement of SNODAS SWE estimates for alpine areas to account for wind redistribution of snow could further improve model performance. Published 2012. This article is a US Government work and is in the public domain in the USA.

Runoff responses to forest thinning at plot and catchment scales in a headwater catchment draining Japanese cypress forest

We examined the effect of forest thinning on runoff generation at plot and catchment scales in headwater basins draining a Japanese cypress (Chamaecyparis obtusa) forest. We removed 58.3% of the stems (corresponding to 43.2% of the basal area) in the treated headwater basin (catchment M5), and left the control catchment (M4) untreated. In both catchments, we monitored overland flow from hillslope plots and stream runoff from catchments at basin outlets over a 2-year pre-thinning period and a 2-year post-thinning period. Paired catchment analysis revealed that annual catchment runoff increased 240.7 mm after thinning. Delayed runoff increased significantly, while quick runoff followed similar patterns in the pre- and post-thinning periods. Flow duration in the ephemeral channel in catchment M5 increased from 56.9% in the pre-thinning period to 73.3% in the post-thinning period. Despite the changes in hydrological responses at the catchment scale, increases in overland flow were not significant. The increased availability of water in the soil matrix, caused by decreased interception loss and evapotranspiration, increased base flow after thinning. Based on the summarized data of previous studies together with this study, the effects of forest thinning on increases in runoff were less than partial harvesting in which the managed areas were concentrated within a watershed. We demonstrated that the effect of forest thinning was strongly scale dependent, an important finding for optimizing water and forest management in forested watersheds.

Surface texture analysis of a high-resolution DTM: Interpreting an alpine basin

This study focuses on a geomorphometric analysis of a small headwater basin of the Dolomites (Eastern Italian Alps). The analysis of the fine-scale surface morphology was performed on a high-resolution digital elevation model (cell size 2m) derived from an airborne LiDAR survey. The topography was synthesized by a set of spatial-statistical indexes representing a signature of the local surface morphology, and the relationships between the geomorphometric indexes and morphological features were evaluated. The studied basin was then segmented by means of a fuzzy k-means classification algorithm into different spatial subdomains characterized by specific local geomorphic signatures. This method allows us to objectively choose the optimal combination of the number of classes and degree of fuzziness characterizing the studied basin. Accordingly, the analysis can be adopted to the characteristics of the studied area, improving the interpretation of surface morphology and offering a valuable tool for comparative analysis between different geomorphological contexts. The results of the classification were compared with the actual surface morphology based on high-resolution orthophotos, aerial photos, and field surveys.

Ecosystem Processes and Human Influences Regulate Streamflow Response to Climate Change at Long-Term Ecological Research Sites

Analyses of long-term records at 35 headwater basins in the United States and Canada indicate that climate change effects on streamflow are not as clear as might be expected, perhaps because of ecosystem processes and human influences. Evapotranspiration was higher than was predicted by temperature in water-surplus ecosystems and lower than was predicted in water-deficit ecosystems. Streamflow was correlated with climate variability indices (e.g., the El Nino—Southern Oscillation, the Pacific Decadal Oscillation, the North Atlantic Oscillation), especially in seasons when vegetation influences are limited. Air temperature increased significantly at 17 of the 19 sites with 20- to 60-year records, but streamflow trends were directly related to climate trends (through changes in ice and snow) at only 7 sites. Past and present human and natural disturbance, vegetation succession, and human water use can mimic, exacerbate, counteract, or mask the effects of climate change on streamflow, even in reference basin...

Uncertainties in Hydrologic and Climate Change Impact Analyses in Headwater Basins of British Columbia

Abstract Three headwater basins located across British Columbia (BC) were analyzed using a hydrologic model driven by five global climate models (GCMs) and three scenarios from the Special Report on Emissions Scenarios (SRES) to project future changes in seasonal water budgets and assess the uncertainty in the projections arising from GCMs, emissions scenarios, and hydrologic model parameterizations under two future time periods. Future projected changes in temperature are for annual increases of approximately +2°C by the 2050s and +3°C by the 2080s. The 2050s and 2080s precipitation projections are for increased winter precipitation in all basins and decreases in summertime precipitation for two of the three basins—with increases projected in the northeastern BC subwatershed. The study found that the hydrologic parameter uncertainty ranged up to 55%, (average 31%) for winter runoff anomalies, which was less than the uncertainty associated with GCMs and emissions scenarios that ranged up to 135% and 78% (average 84% and 58%, respectively). The uncertainty results were variable across the three hydroclimate regimes. Coastal headwater systems in British Columbia experience more uncertainty associated with changes during winter and the summer, whereas interior systems experience the greatest uncertainties during the winter and spring. Changes projected for the 2050s at the coastal site fell outside of the range of natural variability, a robust shift that may result in a very different regime for this basin within the short planning horizon of 50 years. A small, semiarid watershed located on the Chilcotin Plateau exhibited changes to the hydrologic regime that were projected to be small in absolute terms and fell within the range of natural variability.

First impact assessment of hydrological cycle in the Tana River Basin, Kenya, under a changing climate in the late 21st Century

This study projects the hydrological cycle in the Tana River Basin, Kenya, under a changing climate due to anthropogenic greenhouse gas emissions in the late 21st century, using a 20-km mesh atmospheric global climate model (AGCM) and a 0.5°–mesh global river-routing model. In addition, 60-km mesh AGCM ensemble experiments forced with four different projected sea-surface temperatures were performed to quantify the uncertainty in the climate projections. All four climatological annual mean hydroclimate variables: precipitation, evaporation, total runoff, and soil water storage: are projected to increase in the future climate. With the exception of total runoff and river discharge in the headwater basin, these climatological annual mean increases are generally robust. The four climatological monthly mean hydroclimate variables at Garissa are projected to exhibit different seasonal changes. The long rains season is projected to have almost the same peak precipitation amount in April with an earlier onset, while the short rains season is projected to have significant precipitation increases. River discharges are projected to increase significantly in November to March with high consistent change in sign and in June. These results should guide the awareness programs of the changing hydrological cycles in the Tana River Basin and assist in formulating mitigation and adaptation strategies to meet these challenges.

Climate Change Impacts on the Seasonal Distribution of Runoff in a Snowy Headwater Basin, Niigata

Climate Change Impacts on the Seasonal Distribution of Runoff in a Snowy Headwater Basin, Niigata

Upper Colorado River and Great Basin streamflow and snowpack forecasting using Pacific oceanic–atmospheric variability

Summary Water managers in western U.S., including areas such as the State of Utah, are challenged with managing scarce resources and thus, rely heavily on forecasts to allocate and meet various water demands. The need for improved streamflow and snowpack forecast models in the Upper Colorado River and Great Basin is of the utmost importance. In this research, the use of oceanic and climatic variables as predictors to improve the long lead-time (three to nine months) forecast of streamflow and snowpack was investigated. Singular Value Decomposition (SVD) analysis was used to identify a region of Pacific Ocean SSTs and a region of 500 mbar geopotential height (Z 500 ) that were teleconnected with streamflow (and snowpack) in Upper Colorado River and Great Basin headwaters. The resulting Pacific Ocean SSTs and Z 500 regions were used to create indices that were then used as predictors in a non-parametric forecasting model. The majority of forecasts resulted in positive statistical skill, which indicated an improvement of the forecast over the climatology or no-skill forecast. The results indicated that derived indices from Pacific Ocean SSTs were better suited for long lead-time (six to nine month) forecasts of streamflow (and snowpack) while the derived indices from Z 500 improved short-lead time (3 month) forecasts. In all, the results of the forecast model indicated that incorporating Pacific oceanic-atmospheric climatic variability in forecast models can lead to improved forecasts for both streamflow and snowpack.

Post-glacial landscape response to climate variability in the southeastern San Juan Mountains of Colorado, USA

Geomorphic mapping in the upper Conejos River Valley of the San Juan Mountains has shown that three distinct periods of aggradation have occurred since the end of the last glacial maximum (LGM). The first occurred during the Pleistocene–Holocene transition (~ 12.5–9.5 ka) and is interpreted as paraglacial landscape response to deglaciation after the LGM. Evidence of the second period of aggradation is limited but indicates a small pulse of sedimentation at ~ 5.5 ka. A third, more broadly identifiable period of sedimentation occurred in the late Holocene (~ 2.2–1 ka). The latest two periods of aggradation are concurrent with increases in the frequency of climate change in the region suggesting that Holocene alpine and sub-alpine landscapes respond more to rapid changes in climate than to large singular climatic swings. Soil development and radiocarbon dating indicate that hillslopes were stable during the Holocene even while aggradation was occurring in valley bottoms. Thus, we can conclude that erosion does not occur equally throughout the landscape but is focused upslope of headwater streams, along tributary channels, or on ridge tops. This is in contrast to some models which assume equal erosion in headwater basins.

Land use and climate change impacts on the hydrology of the upper Mara River Basin, Kenya: results of a modeling study to support better resource management

Abstract. Some of the most valued natural and cultural landscapes on Earth lie in river basins that are poorly gauged and have incomplete historical climate and runoff records. The Mara River Basin of East Africa is such a basin. It hosts the internationally renowned Mara-Serengeti landscape as well as a rich mixture of indigenous cultures. The Mara River is the sole source of surface water to the landscape during the dry season and periods of drought. During recent years, the flow of the Mara River has become increasingly erratic, especially in the upper reaches, and resource managers are hampered by a lack of understanding of the relative influence of different sources of flow alteration. Uncertainties about the impacts of future climate change compound the challenges. We applied the Soil Water Assessment Tool (SWAT) to investigate the response of the headwater hydrology of the Mara River to scenarios of continued land use change and projected climate change. Under the data-scarce conditions of the basin, model performance was improved using satellite-based estimated rainfall data, which may also improve the usefulness of runoff models in other parts of East Africa. The results of the analysis indicate that any further conversion of forests to agriculture and grassland in the basin headwaters is likely to reduce dry season flows and increase peak flows, leading to greater water scarcity at critical times of the year and exacerbating erosion on hillslopes. Most climate change projections for the region call for modest and seasonally variable increases in precipitation (5–10 %) accompanied by increases in temperature (2.5–3.5 °C). Simulated runoff responses to climate change scenarios were non-linear and suggest the basin is highly vulnerable under low (−3 %) and high (+25 %) extremes of projected precipitation changes, but under median projections (+7 %) there is little impact on annual water yields or mean discharge. Modest increases in precipitation are partitioned largely to increased evapotranspiration. Overall, model results support the existing efforts of Mara water resource managers to protect headwater forests and indicate that additional emphasis should be placed on improving land management practices that enhance infiltration and aquifer recharge as part of a wider program of climate change adaptation.

Development of streamflow projections under changing climate conditions over Colorado River basin headwaters

Abstract. The current drought over the Colorado River Basin has raised concerns that the US Department of the Interior, Bureau of Reclamation (Reclamation) may impose water shortages over the lower portion of the basin for the first time in history. The guidelines that determine levels of shortage are affected by relatively short-term (3 to 7 month) forecasts determined by the Colorado Basin River Forecast Center (CBRFC) using the National Weather Service (NWS) River Forecasting System (RFS) hydrologic model. While these forecasts by the CBRFC are useful, water managers within the basin are interested in long-term projections of streamflow, particularly under changing climate conditions. In this study, a bias-corrected, statistically downscaled dataset of projected climate is used to force the NWS RFS utilized by the CBRFC to derive projections of streamflow over the Green, Gunnison, and San Juan River headwater basins located within the Colorado River Basin. This study evaluates the impact of changing climate to evapotranspiration rates and contributes to a better understanding of how hydrologic processes change under varying climate conditions. The impact to evapotranspiration rates is taken into consideration and incorporated into the development of streamflow projections over Colorado River headwater basins in this study. Additionally, the NWS RFS is modified to account for impacts to evapotranspiration due to changing temperature over the basin. Adjusting evapotranspiration demands resulted in a 6 % to 13 % average decrease in runoff over the Gunnison River Basin when compared to static evapotranspiration rates. Streamflow projections derived using projections of future climate and the NWS RFS provided by the CBRFC resulted in decreased runoff in 2 of the 3 basins considered. Over the Gunnison and San Juan River basins, a 10 % to 15 % average decrease in basin runoff is projected through the year 2099. However, over the Green River basin, a 5 % to 8 % increase in basin runoff is projected through 2099. Evidence of nonstationary behavior is apparent over the Gunnison and San Juan River basins.

Effect of forest thinning on overland flow generation on hillslopes covered by Japanese cypress

AbstractWe examined the effect of forest thinning on overland flow generation on hillslopes covered by a Japanese cypress plantation in central Japan. We established hillslope plots in two small catchments for monitoring overland flow generation. Plot 1 was a treatment plot in which 58·3% of stems were removed, whereas plot 2 remained untreated as a control. Overland flow generation was examined using (1) time‐trend analysis that compared overland flow to precipitation 2 years pre‐ and 2 years post‐thinning in plot 1 and (2) a paired‐plot approach that compared the treated hillslope plot to the control hillslope plot. The latter approach was more reliable for identifying treatment effects than time‐trend analysis because it minimizes effects of climate variables such as precipitation. Unlike our initial hypothesis for decreasing overland flow after thinning, paired‐plot analysis indicated that after thinning, mean overland flow increased 7·4% (corresponding to 2·1 mm) in response to individual storms and 4·8% (corresponding to 7·9 mm) based on monthly data. Although the increased overland flow is likely associated with reduced canopy interception, the reduction of canopy interception after the thinning was estimated to be greater than the increases in overland flow. Therefore, some of the increased throughfall was infiltrated into the soil matrix. Such changes in hillslope‐scale runoff in response to thinning alter the distribution of water in the surface and subsurface layers of forested headwater basins. Copyright © 2010 John Wiley & Sons, Ltd.

Aquatic Biodiversity in the Amazon: Habitat Specialization and Geographic Isolation Promote Species Richness.

Simple SummaryThe immense rainforest ecosystems of tropical America represent some of the greatest concentrations of biodiversity on the planet. Prominent among these are evolutionary radiations of freshwater fishes, including electric eels, piranhas, stingrays, and a myriad of small-bodied and colorful tetras, cichlids, and armored catfishes. In all, the many thousands of these forms account for nearly 10% of all the vertebrate species on Earth. This article explores the complimentary roles that ecological and geographic filters play in limiting dispersal in aquatic species, and how these factors contribute to the accumulation of species richness over broad geographic and evolutionary time scales.The Neotropical freshwater ichthyofauna has among the highest species richness and density of any vertebrate fauna on Earth, with more than 5,600 species compressed into less than 12% of the world's land surface area, and less than 0.002% of the world's total liquid water supply. How have so many species come to co-exist in such a small amount of total habitat space? Here we report results of an aquatic faunal survey of the Fitzcarrald region in southeastern Peru, an area of low-elevation upland (200–500 m above sea level) rainforest in the Western Amazon, that straddles the headwaters of four large Amazonian tributaries; the Juruá (Yurúa), Ucayali, Purús, and Madre de Dios rivers. All measures of fish species diversity in this region are high; there is high alpha diversity with many species coexisting in the same locality, high beta diversity with high turnover between habitats, and high gamma diversity with high turnover between adjacent tributary basins. Current data show little species endemism, and no known examples of sympatric sister species, within the Fitzcarrald region, suggesting a lack of localized or recent adaptive divergences. These results support the hypothesis that the fish species of the Fitzcarrald region are relatively ancient, predating the Late Miocene-Pliocene (c. 4 Ma) uplift that isolated its several headwater basins. The results also suggest that habitat specialization (phylogenetic niche conservatism) and geographic isolation (dispersal limitation) have contributed to the maintenance of high species richness in this region of the Amazon Basin.