1. Introduction
Physical hydraulic and geomorphic characteristics of river systems serve as important controls on the habitat for aquatic species. Characterization of these physical habitat variables is important for describing and comparing different river systems, examining changes in habitat resulting from natural or human-caused disturbances, and developing and implementing conservation and restoration goals. To aid in these objectives, numerous channel habitat unit classification systems have been developed. Many of these systems are based on hydraulic and geomorphic variables. For example, Frissell et al. [
1] developed a hierarchical classification for stream habitats, with channel habitat units classified on the basis of reach class, bed topography, water surface slope, morphogenetic structure or process, substrate, and bank configuration. Hawkins et al. [
2] developed a channel habitat unit classification system based on a division between fast water (which is further subdivided into rough and smooth) and slow water (which is further subdivided into scour pools and dammed pools). Montgomery and Buffington [
3] classified mountain channels based on typical bed material, bedform pattern, dominant roughness elements, dominant sediment sources, sediment storage elements, typical confinement, and typical pool spacing. Thomson et al. [
4] used the River Styles framework to classify channel habitat units according to planform, channel geometry, and textural controls. A common feature of hydraulic and geomorphic classifications is that they are based primarily on physical variables such as slope, substrate, and width-to-depth ratio.
Other channel habitat unit classification approaches relate species assemblages to physical habitat characteristics. For example, Buffagni et al. [
5] used macroinvertebrate species assemblages to identify channel habitat units, resulting in classes of margin with macrophytes, margin without macrophytes, backwater, run-riffle, and macrophytes in the flow, based on flow velocity, depth, Froude number, granulometric composition and substratum roughness, and cover. Newson and Newson [
6] proposed using a “habitat hydraulics” approach to channel habitat unit classification, based on flow types and physical biotopes analogous to mesohabitats and functional habitats in ecology. In these ecological approaches, the species assemblages are classified first, and the physical variables that control their spatial patterns are determined afterwards.
Regardless of whether channel habitat units are classified using geomorphic or ecological approaches, the role of classification in meeting conservation goals is similar. Many aquatic species have specific physical habitat requirements, which may differ for different stages of their life cycles. A diversity of channel habitat units is therefore typically recognized as being conducive to healthy aquatic ecosystems. For example, Crispin et al. [
7] found that restoration of large woody debris resulted in a fivefold increase in summer habitat for coho salmon and a sixfold increase in winter habitat through the creation of additional pool-and-riffle channel habitat units. Quinn and Peterson [
8] also found that wild juvenile coho salmon survival rates were strongly correlated with complexity of channel habitat units. Angermeier and Karr [
9] found that all fish feeding guilds other than aquatic insectivores were most concentrated in deep pools in streams in Panama. Willis et al. [
10] analyzed fish assemblages in a Venezuelan river and found that species density was negatively associated with flow velocity and positively associated with habitat complexity. Montaña and Winemiller [
11] found that cichlid species partitioned habitat at the local scale, based on features such as leaf litter, sand banks, rocky shoals, and woody debris. Delineation of channel habitat units is an important preliminary step in assessing the baseline condition of a river system and in setting goals for conservation and restoration.
Despite the importance of being able to delineate channel habitat units, the process is not always straightforward. Channel habitat units are often delineated by visual assessment during a field visit, which is problematic for two reasons. First, visual assessments can be subjective, even for well-defined channel habitat units such as pools, riffles, and runs. Roper and Scarnecchia [
12] determined that there was significant variability in how trained observers classified channel habitat units, especially when there was a large number of habitat unit types to choose from, although the variability decreased with a higher level of training. To alleviate the problem of observer subjectivity, some researchers have proposed ecohydraulic channel habitat unit classifications, such as those based on velocity/depth ratio, Froude number, and slope [
13]. Such ecohydraulic approaches make for more objective and robust classifications.
A more fundamental limitation of the surveying-based approach to channel habitat unit delineation is that it requires costly and time-consuming fieldwork. This limitation can be addressed through the use of remote sensing and hydraulic modeling. With some field data available for validation, it is possible to use remotely-sensed imagery to estimate some hydraulic variables needed for ecohydraulic characterization of channel habitat units (i.e., depth). Remotely-sensed data can in turn be used to initialize hydraulic models that can simulate additional variables needed for classification (i.e., velocity). A combination of minimal field measurements, remote sensing, and hydraulic modeling can be used to characterize channel habitat units for longer segments of river than would be feasible to survey manually, or to delineate channel habitat units for remote regions that are difficult to access.
Tropical rivers are especially rich targets for remotely-sensed channel habitat unit classifications, because they are often relatively difficult to access and lack extensive observed data. In part because of the difficulty of fieldwork on tropical rivers, these systems are generally under-studied in comparison with midlatitude rivers, for which the vast majority of channel habitat unit classification systems were developed. Although both midlatitude and tropical rivers are highly variable, there are some common characteristics of tropical rivers that make them different enough that ideas regarding channel habitat units on midlatitude rivers may not necessarily apply [
14]. For example, tropical climates are the wettest in the world, so tropical rivers have high rates of discharge per unit area. Despite overall high rates of precipitation and discharge, many tropical climates are also highly variable in their hydroclimatic regime, with distinct wet and dry seasons, and often large interannual variability resulting from oceanic-atmospheric phenomena such as the El Niño-Southern Oscillation. These pronounced differences in precipitation between wet and dry periods result in large variations in discharge. During high flows, sediment transport capacity is large, resulting in significant bedload sediment transport. Sediment supply is likely to be large, because the warm temperatures and abundant moisture mean that the tropics have the highest weathering rates in the world. Decomposition rates are also high, so large woody debris does not persist in channels for as long as in midlatitude rivers, although the supply of new large woody debris is large because of the dense vegetation. Except at the highest elevations, tropical river basins were never glaciated and can therefore generally be expected to be better-graded than many higher-latitude rivers that are still adjusting to the retreat of Pleistocene ice sheets. In general—because of the high rates of discharge and sediment transport, lack of stabilized large woody debris, and history of no glaciation—tropical rivers could reasonably be expected to adjust their channels more rapidly to changes in external drivers, compared to many midlatitude rivers.
In this paper, we develop a method for delineating channel habitat units from remotely-sensed imagery and hydraulic modeling, and apply the method to a tropical montane river. We also examine the spatial patterns of delineated channel habitat units on the study river and compare them to those that would be expected based on studies of midlatitude rivers. The method described here is novel, and this study is also among the first that focuses on channel habitat units of tropical rivers.
3. Results
Figure 2b shows the HAB-2 water depths for the Bladen River, which reach a maximum depth of 4 m and are significantly correlated with observed depths in Reach 1 and Reach 2 (
Figure 5 and
Table 3).
Figure 2c shows the water-surface elevation, which was estimated by extending a constant water-surface slope upstream from the measured water-surface elevation of Reach 1. The HAB-2 depths (
Figure 2b) were subtracted from the estimated water-surface elevation (
Figure 2c) to create the bed elevation used as input to the FaSTMECH model (
Figure 2d).
Figure 6 shows the FaSTMECH modeling results. The modeled water depths (
Figure 6a) are substantially higher than the observed values in Reach 1 and Reach 2 (
Figure 5b), and the correlation between the two is lower than between the HAB-2 estimated depths and the measured values, although the two are significantly correlated (
Table 3). The modeled flow velocities (
Figure 6b) are also substantially higher than the observed values. The distribution of velocities is clustered around relatively small values (<2 m s
−1), but there are several instances of high velocity (>3 m s
−1) that show up in both the observations and the model output (
Figure 5c). These high-velocity points are associated with rapids exiting a deep pool near the upstream end of Reach 1. Overall, the correlation between observed and modeled flow velocity is significant, and the R
2 is higher for FaSTMECH-simulated velocity than for FaSTMECH-simulated depth, but is not as high as for HAB-2 depths. This finding indicates that the hydraulic modeling component introduces more uncertainty into the overall channel habitat unit delineation process than does the remote sensing. Both depths estimated from HAB-2 and depths and velocities modeled by FaSTMECH are positively biased, meaning depth and velocity are both overestimated by the modeling procedures (
Figure 5). The most likely explanation for this systematic bias is that discharge was higher during the period of aerial imagery acquisition than during the field data collection period, resulting in higher estimates of both depth and velocity.
We used the modeled depth and velocity to classify the entire study segment into three types of channel habitat unit: pools, riffles, and runs. Although there are more types of channel habitat unit, we selected these three because (1) they are the most commonly identified channel habitat units that are known to provide a range of different habitat conditions of importance to aquatic species [
1,
2,
3,
4]; and (2) these channel habitat units can be classified on the basis of depth and velocity alone, without the need for additional data, such as substrate conditions, that are not available for the entire study segment.
Figure 7a shows the delineated channel habitat units, based on applying on the Jowett [
13] velocity-depth thresholds to the FaSTMECH modeling results, for the entire Bladen River study segment, while
Figure 7c shows a detailed close-up of a particular reach.
Figure 7b shows average velocity-depth ratios for 100-m bins of the channel from upstream to downstream, along with the thresholds for the channel habitat units. In general, the most frequent shifts between channel habitat units are concentrated in the middle segment, which is where the transition from the confined montane to alluvial floodplain channel occurs.
To validate the channel habitat unit classification, we also applied the Jowett [
13] velocity-depth ratios to 186 points in Reach 1 and Reach 2 for which we had field measurements of velocity and depth, and compared these points to the channel habitat units delineated for those locations using the HAB-2/FaSTMECH procedure. Overall, 83% of points were classified accurately, with accuracies ranging from 81% for runs to 88% for riffles (
Table 4). This finding indicates that, despite the previously noted systematic biases in the HAB-2 depths and FaSTMECH-simulated depth and velocities, the classification procedure results in reasonably accurate channel habitat units.
According to the channel habitat unit classification, the unit that covered the greatest length in the study segment was runs, which made up nearly half (47%) of the total segment length (
Table 5). The dominance of runs may be explained by the intermediate values of velocity-depth ratio used to define a run, which makes it essentially a catchall channel habitat unit for any portion of the channel that is not a pool or riffle. The second most prevalent habitat unit was pools, which made up 32% of the total segment length. As expected, pools were most commonly found on the outside of bends, and also in some areas along one side of the channel with a riffle on the other side (
Figure 7c. The least prevalent channel habitat unit was riffles, which made up just 21% of the total segment length. The low prevalence of riffles may be explained in part by the igneous lithology of the study segment, in which quartz-rich parent material weathers predominantly to sand. The sand is deposited relatively evenly in the channel to create runs, rather than the concentrated deposition needed to create riffles, which would be expected for coarser bed material that requires a higher competence, such as gravels [
19].
Table 4 also shows the average spacing between each delineated channel habitat unit. The spacing ranges from 284 m for pools to 550 m for runs. This indicates that, although runs covered more of the segment length than any other channel habitat unit, they were predominantly found in several very long expanses that were far apart from one another (
Figure 7a). Pools and riffles, in contrast, while covering less of the total segment length, tended to alternate more frequently with other channel habitat units than did runs. The commonly reported value for pool and riffle spacing is five to seven bankfull channel widths [
20]. Based on our field-measured average bankfull width value for Reach 1 of ~35 m, the expected pool and riffle spacing for the Bladen River study segment would be only 175 to 245 m. Our delineated channel habitat units all had greater average spacing than this expected value, which suggests the potential for greater overall amplitude of channel habitat units in the study segment.
4. Discussion
This study involved the development of an innovative approach to delineating channel habitat units using a combination of fieldwork, remote sensing, and hydraulic modeling. Channel habitat units have often been delineated from aerial imagery, but usually this is done using supervised classification. For example, Marcus et al. [
21] classified in-stream habitat using supervised classification of high-resolution hyperspectral imagery. An advantage of the approach presented here is that it is based on hydraulic variables (depth and velocity), rather than purely on the spectral signatures generated by different channel habitat units. This incorporation of hydraulic variables can only be accomplished if hydraulic modeling is combined with remote sensing. For example, Legleiter and Goodchild [
22] incorporated hydraulic modeling into the remote sensing process with a combination of supervised classification and fuzzy clustering to delineate channel habitat units. Such combined approaches, however, have rarely been applied, especially on tropical rivers.
A limitation of optical remote sensing to delineate channel habitat units is that the algorithms do not work in turbid water, or where canopy cover obstructs the channel. The Bladen River presented an ideal case study for optical remote sensing of channel habitat units, because its water is very clear during the dry season and it is wide enough that the channel is never completely obstructed by vegetation. Such ideal conditions, however, are not common in tropical rivers, which often experience high turbidity and have dense riparian vegetation. In such cases, alternative sensors such as blue-green LiDAR may be preferable for mapping channel habitat units. For example, McKean et al. [
23] used a narrow-beam aquatic-terrestrial LiDAR to map and calculate the volume of pools. Because of the great expense of LiDAR sensors and the need to have the imagery flown, however, approaches based on optical imagery still fulfill a necessary function.
In this study, both the HAB-2 estimates of depth and the FaSTMECH-simulated depths and velocities exhibited a positive bias. The likely cause is the fact that the dates of aerial image acquisition and field data collection were not the same. If the discharge was higher when the aerial imagery was flown compared to when we collected the field data, this would result in overestimates of depth in the HAB-2 model. This positive bias in depth would then propagate to the depths and velocities simulated by FaSTMECH. Future research on initializing hydraulic models with remotely-sensed depth estimates should endeavor to collect field data concurrently with image acquisition, to avoid such cases of systematic bias. The bias problem could also be alleviated with observed discharge data from gaging stations, which would allow for more precise calibration of the β diffuse attenuation coefficient in the HAB-2 model. These corrective steps were not possible on the Bladen River because it lacks gaging stations, so the systematic bias in the depth and velocity measurements remains a fundamental limitation of the study. It should be noted, however, that the ratio between velocity and depth is more important than their absolute values in delineating channel habitat units, and the comparison of the observed and modeled ratios indicates that the classification procedure was largely successful in accurately distinguishing channel habitat units in Reach 1 and Reach 2.
The results of this study indicate that the greatest source of uncertainty in the approach taken here is the hydraulic modeling. The depth estimates from the HAB-2 model match reasonably well to observations, but the FaSTMECH-modeled depth and velocity did not match observations as well. A number of limitations of this study contribute to FaSTMECH’s modeling performance. In particular, the estimation of water-surface elevation in the upstream segment and the estimation of the upstream discharge boundary condition using downstream discharge are likely to be the main sources of error in the hydraulic modeling. These issues could be remedied by a minimal amount of additional field data collection at the upstream end of the study segment, including measurement of water-surface elevation and discharge to serve as upstream boundary conditions. In particular, both issues would be addressed if a stream gage were available to record stage and discharge at the upstream end of the study segment. Availability of gage data would also allow the low-flow conditions during the dry season simulated here to be extended to high flows during the rainy season, to allow for analysis of change over time in channel habitat units. The Bladen River is currently ungaged, which is common in remote tropical rivers. Future research should consider using aerial photography specifically acquired for processing through Structure-from-Motion, so that water level could be mapped directly from the imagery [
24].
This study found some evidence that the spacing of channel habitat units on this segment of the Bladen River is larger than the typical values reported for midlatitude rivers. This difference could potentially suggest an overall amplified wavelength of channel habitat units, which would be consistent with the high erosion rates resulting from high discharge per unit area and high weathering rates in tropical rivers. Many other empirical relationships developed to predict characteristics of rivers, primarily based on midlatitude rivers, have been found inadequate when applied to tropical rivers. For example, Baker [
25] found that the sediment-size sinuosity relationships developed for temperate rivers do not apply to tropical rivers. As noted by Latrubesse et al. [
26], the empirical equations used to relate channel morphology and sediment load to predict channel patterns often fail to predict tropical river planforms. Because this study examined only a short segment of a single river system, there is not enough information to conclude that tropical rivers generally have larger spacing of channel habitat units than is typical for midlatitude rivers, but future research should investigate this possibility by examining longer river segments in a diversity of tropical environments.
Moreover, many tropical rivers experience significant flow variability between the dry and wet seasons, which could result in channel habitat units being more dynamic than in many midlatitude rivers. For example, Rayner et al. [
27] found that small tropical rivers that lack extensive floodplain storage experience a reduction in habitat heterogeneity during the wet season, when bed sediments are mobilized and aquatic vegetation is scoured. To address the question of whether channel habitat units in tropical rivers are more dynamic seasonally or change over time more rapidly than those in midlatitude rivers, additional research is needed to delineate changes in channel habitat units over time.
In addition to what it reveals about fundamental fluvial processes, the delineation of channel habitat units has important implications for river management. Physical habitat assessments are widely used for targeting priority areas for conservation, designing stream restoration projects, and setting environmental instream flow standards [
28]. Managing rivers for conservation objectives is especially critical in tropical rivers, given their high levels of aquatic biodiversity relative to higher-latitude rivers [
29]. Native aquatic species in the Bladen River, including several cichlid species, have been found to be sensitive to physical habitat variables [
30,
31]. Given the high aquatic biodiversity of tropical rivers—and threats to those rivers from climate change, land-use change, dam construction, and invasive species—it is imperative to build more comprehensive understanding of the controls on and spatial distribution of channel habitat units in tropical rivers. In particular, species richness is expected to be higher in heterogeneous habitat types, such as sequences of pools and riffles, rather than in relatively uniform stretches of runs.