3.1. Vegetation and Site Data
We sampled a range of wetland habitats in the SRD and identified 34 taxa or habitat descriptors across the 72 sites sampled, including habitats dominated by open water and floating leaved plants (e.g., Nymphoides, Nuphar, Nymphaea), floating plants (Lemna, Spirodela, Utricularia), rooted water plants (e.g., Potamogeton, Myriophyllum, Ceratophyllum), emergent plants (e.g., Polygonum, Hippuris), and several ruderal plants from grazed areas (e.g., Trifolium, Galeopsis, and Agrostis). The most commonly identified genera and/or wetland habitats included open water, Nymphoides, thatch (i.e., the previous seasonʼs often unidentifiable senescent or dead vegetation), Calamagrostis, Carex, and Equisetum. These occurred at >10% cover (i.e., the lower limit at which a vegetation class or habitat was recorded for this study) at >5% of the plots.
3.2. Multi-Scale Hierarchical Habitat Classification
Although we aggregated classes from the finest level to the most general, the results are more logically described from the coarsest to the finest resolution. We anticipate that end-users would select the hierarchical level at which their needs would be met. At the coarsest scale, or Level 1, we described the entirety of the inland freshwater deltaic wetland (data not shown). This top-level class corresponds to the “System” level in the Cowardin wetland classification scheme and the sub-type “permanent inland deltas (L)” of the general wetland type “inland wetlands” in the RCS [
49,
50]. The second level consisted of two classes (
Figure A1): (1) open water and aquatic bed and (2) emergent herbaceous and woody wetland. The Level 3 classification resulted in five basic aquatic and wetland types: (1) stream, river and lake bed; (2) unconsolidated bottom; (3) aquatic bed; (4) emergent herbaceous wetland; and (5) scrub-shrub wetland (
Figure 6). Level 3 generally corresponds with the “Class” level in the Cowardin classification scheme. Level 4, with 13 classes, approximates the “Subclass” level in the Cowardin scheme [
49,
51] (
Figure 7). In Level 5, which is the finest scale and which served as the initial classification from which all other groupings were aggregated, we identified 22 classes of aquatic substrates and wetland vegetation cover at genus and community levels (
Figure 8).
Figure 6.
Level 3 classification at the Class level of the Cowardin scheme.
Figure 6.
Level 3 classification at the Class level of the Cowardin scheme.
3.3. Accuracy Assessment
We conducted an accuracy assessment of the habitat classification at the finest scale (Level 5), as subsequent and coarser classifications originated from the collapsing of Level 5 classes to more general groups (
Table 1). Our accuracy assessment was quantitatively evaluated based on a stratified random sample of 16,544 validation pixels from ground-truth site polygons independent of training pixels. As we were limited to the number of sampling points we could visit across the study area, our use of site polygons may over-estimate the accuracy of our study. However, we found overall classification accuracy, computed by taking the total number of correctly classified pixels (
i.e., diagonal cells of confusion matrix) and dividing by the total number of samples, to be 86.5%. We calculated the Kappa coefficient, which is a measure of the likelihood that the observed classification is due to chance (Kappa = 0) or true agreement (Kappa = 1.0) [
52], to be 0.85. Individual class accuracies were evaluated by producer’s accuracy (PA) and user’s accuracy (UA) measures (see
Table 1). The PA for 22 classes ranged from 60.7% to 98.3%, and four classes (Classes 2, 7, 9, and 19) had PA values <70% (
i.e., omission error larger than 30%). The UA for 22 classes ranged from 52.3% to 99.5%, and two classes (Classes 9 and 18) had UA values of <70% (
i.e., commission error larger than 30%).
Figure 7.
Level 4 classification at the Subclass level of the Cowardin scheme.
Figure 7.
Level 4 classification at the Subclass level of the Cowardin scheme.
Figure 8.
Genus and community aquatic and wetland habitat classification at Level 5.
Figure 8.
Genus and community aquatic and wetland habitat classification at Level 5.
Table 1.
Percent classification error for 22 classes of aquatic and wetland habitats at Level 5. Class numbers correspond with those in the legend of
Figure 8.
Table 1.
Percent classification error for 22 classes of aquatic and wetland habitats at Level 5. Class numbers correspond with those in the legend of Figure 8.
Class | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | PA | UA |
---|
1 | 92.5 | - | 0.1 | - | - | - | 26.3 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 92.5 | 92.7 |
2 | - | 60.7 | - | - | - | - | 0.5 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 60.7 | 99.2 |
3 | 7.4 | 36.8 | 91.6 | 4.7 | - | 0.2 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 91.6 | 79.7 |
4 | - | - | 8.3 | 94.6 | 2.3 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 94.6 | 82.3 |
5 | - | - | - | 0.4 | 97.6 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 97.6 | 99.5 |
6 | - | - | - | - | - | 93.5 | 5.3 | 2.1 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 93.5 | 96.6 |
7 | 0.1 | 2.5 | - | - | - | 4.0 | 67.8 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 67.8 | 86.8 |
8 | - | - | 0.1 | - | - | 0.7 | - | 70.6 | 16.9 | - | - | - | - | - | - | - | - | - | - | - | - | - | 70.6 | 82.4 |
9 | - | - | - | - | - | 1.2 | - | 9.6 | 69.9 | 8.5 | 2.0 | 0.6 | - | - | - | - | - | - | - | - | - | 0.2 | 69.9 | 52.3 |
10 | - | 0.2 | - | - | - | 0.4 | - | 17.8 | 1.2 | 86.5 | 4.5 | 0.2 | - | - | - | - | - | - | - | - | - | - | 86.5 | 71.4 |
11 | - | - | - | - | - | - | - | - | 2.4 | 5.0 | 86.8 | 4.9 | - | - | - | - | - | - | - | - | - | - | 86.8 | 88.0 |
12 | - | - | - | - | - | - | - | - | - | - | 6.8 | 91.2 | 1.5 | - | - | - | - | - | - | - | - | - | 91.2 | 95.2 |
13 | - | - | - | - | - | - | - | - | - | - | - | 2.7 | 98.3 | - | - | - | - | - | - | - | - | - | 98.3 | 96.2 |
14 | - | - | - | 0.4 | 0.2 | - | - | - | 8.4 | - | - | - | - | 97.2 | 3.7 | - | - | - | 0.7 | - | - | 0.1 | 97.2 | 90.2 |
15 | - | - | - | - | - | - | - | - | - | - | - | - | - | 0.8 | 79.3 | 4.8 | - | - | - | - | - | 0.1 | 79.3 | 84.3 |
16 | - | - | - | - | - | - | - | - | - | - | - | - | - | 1.2 | 16.6 | 94.2 | - | 0.4 | 2.2 | - | - | 0.8 | 94.2 | 88.2 |
17 | - | - | - | - | - | - | - | - | 1.2 | - | - | - | 0.2 | 0.4 | 0.5 | - | 83.9 | 19.8 | 7.9 | - | - | 0.1 | 83.9 | 78.9 |
18 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 0.7 | 12.2 | 74.4 | 27.7 | 1.3 | - | 1.0 | 74.4 | 53.4 |
19 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 2.5 | 3.0 | 60.9 | - | - | - | 60.9 | 93.0 |
20 | - | - | - | - | - | - | - | - | - | - | - | 0.2 | - | - | - | - | 1.0 | 2.2 | 0.1 | 94.4 | 8.1 | 14.2 | 94.6 | 74.5 |
21 | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | - | 89.1 | 2.1 | 89.1 | 94.7 |
22 | - | - | - | - | - | - | - | - | - | - | - | 0.3 | - | 0.4 | - | 0.3 | 0.4 | 0.2 | 0.4 | 4.3 | 2.8 | 81.4 | 81.4 | 95.0 |
3.4. ISA/Vegetation Analyses
We identified 11 indicator taxa/habitats using ISA for the 22 wetland classes at Level 5 (
Table 2 and
Figure 8), and in no cases were multiple plants/habitats identified with sufficient fidelity and specificity to be indicators of the same wetland class. This suggests that at 22 classes, our analyses identified plants and habitats that tended to dominate plots (
Nymphoides,
Potamogeton,
Phragmites, Equisetum, thatch (often composed of last season’s
Equisetum), bare ground,
etc.) in the multiple sites in which they were found. These taxa and habitats tended to be common throughout the delta. We found >100 measureable occurrences of these 11 indicators.
Table 2.
Indicator and dominant species of Level 5 of the classification. Higher mean indicator values suggest greater fidelity and specificity to habitat classes, and the p-value is based on the Monte Carlo class randomization with 9999 runs.
Table 2.
Indicator and dominant species of Level 5 of the classification. Higher mean indicator values suggest greater fidelity and specificity to habitat classes, and the p-value is based on the Monte Carlo class randomization with 9999 runs.
Class # | Class Name (Indicator and/or Hab. Descriptor) | Indicator Taxa/Habitat | Indicator Value Mean | Indicator p-Value | Dominant Species/Substrate |
---|
1 | Deep Water with Sand Bottom | Not detected | | | Open water |
2 | Shallow Water with Sediment | Not detected | | | Open water |
3 | Shallow Water with Mud Bottom | Not detected | | | Open water |
4 | Very Shallow Water with Sand Bottom | Not detected | | | Open water |
5 | Shallow Water with Sand Bottom | Not detected | | | Open water |
6 | Submerged Aquatic Vascular (Potamogeton) | Potamogeton | 21.1 | 0.0464 | Potamogeton |
7 | Submerged Aquatic Vascular (Sparganium) | Sparganium | 20.9 | 0.0158 | Sparganium |
8 | Submerged Aquatic Vascular (Utricularia) | Utricularia | 31.1 | 0.0661 | Utricularia |
9 | Submerged and Floating Vascular (Agrostis/Eleocharis) | Not detected | | | Agrostis and Eleocharis |
10 | Very Sparse Floating Vascular (Nymphoides) | Not detected | | | Nymphoides |
11 | Sparse Floating Vascular (Nymphoides) | Not detected | | | Nymphoides |
12 | Dense Floating Vascular (Nymphoides) | Not detected | | | Nymphoides |
13 | Very Dense Floating Vascular (Nymphoides) | Nymphoides | 14.3 | 0.0001 | Nymphoides |
14 | Persistent Emergent (Phragmites) | Phragmites | 25.6 | 0.0044 | Phragmites |
15 | Persistent Emergent (Bare Ground and Carex) | Bare Ground | 22.3 | 0.0276 | Bare Ground and Carex |
16 | Persistent Emergent (Equisetum) | Equisetum | 16.3 | 0.0262 | Equisetum |
17 | Persistent Emergent (Thatch) | Thatch | 15.7 | 0.0006 | Thatch |
18 | Persistent Emergent (Carex) | Carex | 17.0 | 0.0028 | Carex |
19 | Persistent Emergent (Calamagrostis) | Calamagrostis | 16.6 | 0.0365 | Calamagrostis |
20 | Persistent Emergent (Scolochloa) | Scolochloa | 22.3 | 0.0097 | Scolochloa |
21 | Persistent Emergent (Amoria/Galeopsis/Trifolium) | Not detected | | | Amoria, Galeopsis, and Trifolium |
22 | Scrub-shrub (Salix with Calamagrostis) | Not detected | | | Salix and Calamagrostis |
The ISA was also applied to each of 13 broad classes at Level 4 of the classification hierarchy (
Table 3), resulting in 11 indicator taxa, including open water, bare ground, and thatch habitats. Only one class (Class 8) had more than one indicator with both
Equisetum and bare ground.
Equisetum forms extensive monocultures that, upon senescence, can be easily removed through flooding or ice scour. It is likely this mechanism results in stands of
Equisetum that often are mixed with extensive areas of bare soil. As found with our Level 5 classification, Level 4 identified the same taxa and habitats that were monoculture forming (e.g.,
Phragmites,
Nymphoides,
Equisetum) or commonly found in unique hydrogeomorphic settings (e.g.,
Calamagrostis was frequently interspersed with
Salix on distributary stream banks). These taxa and habitats dominate the SRD. Open water classes at Level 4 appeared to be bifurcated to more resolved Level 5 classes, including
Potamogeton,
Sparganium, and
Utricularia. Interestingly,
Myriophyllum, a cosmopolitan submerged aquatic genus, and
Nuphar, a floating-leaved vascular plant, were indicators at Level 4, but not at Level 5. These taxa are sufficiently common throughout the SRD to be found at many plots, but were sparse enough in most plots that, with more finely resolved data, they may have been further split into groups based on habitat descriptors (e.g., based on substratum).
Table 3.
Indicator and dominant species of 13 classes of aquatic and wetland habitats for Level 4 of the classification.
Table 3.
Indicator and dominant species of 13 classes of aquatic and wetland habitats for Level 4 of the classification.
Class # | Class Name (Indicator and/or hab. Descriptor) | Indicator Taxa/Habitat | Indicator Value Mean | Indicator p-Value | Dominant Species/Substrate |
---|
1 | Deep Water with Sand Bottom | Open water | 15.4 | 0.0127 | Open Water |
2 | Shallow Water with Mud Bottom | Not detected | | | Open Water |
3 | Shallow Water with Sand bottom | Not detected | | | Open Water |
4 | Submerged Aquatic Vascular (Myriophyllum) | Myriophyllum | 17.9 | 0.0119 | Myriophyllum |
5 | Sparse Floating Vascular (Nuphar) | Nuphar | 19.7 | 0.0411 | Nuphar |
6 | Dense Floating Vascular (Nymphoides) | Nymphoides | 15.9 | 0.0001 | Nymphoides |
7 | Persistent Emergent (Phragmites) | Phragmites | 17.8 | 0.0017 | Phragmites |
8 | Persistent Emergent (Equisetum/Bare Ground) | Equisetum Bare Ground | 16.7, 18.3 | 0.0508, 0.0217 | Equisetum and Bare Ground |
9 | Persistent Emergent (Thatch) | Thatch | 16.4 | 0.0014 | Thatch |
10 | Persistent Emergent (Carex) | Carex | 17.6 | 0.0274 | Carex |
11 | Persistent Emergent (Scolochloa) | Scolochloa | 17.8 | 0.0069 | Scolochloa |
12 | Persistent Emergent (Amoria) | Not detected | | | Amoria |
13 | Scrub-Shrub (Salix with Calamagrostis) | Calamagrostis | 16.9 | 0.0873 | Salix and Calamagrostis |
We did not apply ISA to the classes at Level 3 or above, since the wetland and aquatic habitat classes at these levels are too broad and ecologically heterogeneous to identify indicator or dominant species. Instead, the broad classes at Levels 3 and 2 are to be interpreted and labeled in terms of their topographical positions, substrate composition, and taxonomical relations with the fine-scale wetland habitat classes at Level 4.
3.5. Improved Classification with Additional Spectral Bands and Metrics
In this study, we evaluated the utility of four new spectral bands, NDVI, and a textural measure to differentiate among wetland habitats. The contribution of new bands to wetland habitat discrimination was evaluated by observing the changes in overall classification accuracy, PA and UA for each class, when one or more new bands was included in the classification in addition to four traditional bands (
Table 4). Using the finest resolved hierarchical classification of 22 classes, the overall accuracy with the four traditional spectral bands was only 79.0% and the Kappa coefficient was 0.77. When the coastal band was included in the classification along with the four traditional bands, the UA for Class 1 (Deep Water with Sand Bottom) increased from 86.2% to 97.8% and the PA for Class 7 (Submerged Aquatic Vascular—
Sparganium) increased from 86.1% to 96.4%, despite only a small increase in the overall accuracy from 79.0% to 80.4%. The new yellow, red-edge, and NIR2 bands were useful for discriminating many different types of vegetated habitats, including scrub-shrub, emergent herbaceous, and submerged aquatic vascular. When the yellow, red-edge, and NIR2 bands were used along with the four traditional bands, the overall classification accuracy increased from 79.0% to 82.0%. In particular, the PA for Class 22 (Scrub-shrub) increased from 59.6% to 73.0% and the UA increased from 82.0% to 92.2%. Both the UA and PA for Class 21 (Persistent Emergent—
Amoria/
Galeopsis/
Trifolium) and Class 20 (Persistent Emergent—
Scolochloa) also significantly increased. The UA for Class 16 (Persistent Emergent—
Equisetum) and the PA for Class 7 (Submerged Aquatic Vascular—
Sparganium) improved considerably as well (see
Table A1).
Table 4.
Variation of classification accuracy with different combinations of WV2 spectral bands, NDVI, and texture for 22 classes of wetland and aquatic habitats.
Table 4.
Variation of classification accuracy with different combinations of WV2 spectral bands, NDVI, and texture for 22 classes of wetland and aquatic habitats.
Input | Overall Accuracy |
---|
4 traditional bands only (red, blue, green, NIR) | 79.0% |
4 traditional bands plus coastal band | 80.4% |
4 traditional bands plus yellow, red-edge, NIR2 bands | 82.0% |
8 bands (4 traditional bands plus 4 new bands) | 82.9% |
8 bands plus NDVI | 83.9% |
8 bands plus Texture | 84.8% |
8 bands plus NDVI and Texture | 86.5% |
While some particular classes improved substantially, the combined benefit of the four new bands increased the overall classification accuracy by only about 4%. Understanding the ecology and relative abundance of different habitats within a study system may, then, dictate the utility of the additional bands and processing steps. The classification scale may also help to determine whether the time and resources to acquire and process the additional bands are necessary. When dealing with large systems in remote areas with substantial vegetative heterogeneity, it is prudent to err on the side of caution when conducting vegetative assessments and system classification by acquiring the additional bands to calculate the NDVI and textural measures.
Table 5.
Mean and standard deviation (Stdv) of NDVI and textural homogeneity using the Level 5 classification. NDVI values < 0.0 represent little to no detectable photosynthetic activity, while higher values suggest increased green biomass and synthetic activity. Lower homogeneity values (i.e., 0.0–1.0) represent areas of greater textural heterogeneity.
Table 5.
Mean and standard deviation (Stdv) of NDVI and textural homogeneity using the Level 5 classification. NDVI values < 0.0 represent little to no detectable photosynthetic activity, while higher values suggest increased green biomass and synthetic activity. Lower homogeneity values (i.e., 0.0–1.0) represent areas of greater textural heterogeneity.
Class # | Class Name | NDVI | Homogeneity |
---|
Mean | Stdv | Mean | Stdv |
---|
1 | Deep Water with Sand Bottom | −0.33 | 0.04 | 0.72 | 0.09 |
2 | Shallow Water with Sediment | −0.19 | 0.05 | 0.86 | 0.16 |
3 | Shallow Water with Mud Bottom | −0.27 | 0.11 | 0.87 | 0.16 |
4 | Very Shallow Water with Sand Bottom | −0.18 | 0.09 | 0.87 | 0.13 |
5 | Shallow Water with Sand Bottom | 0.00 | 0.04 | 0.66 | 0.16 |
6 | Submerged Aquatic Vascular (Potamogeton) | −0.05 | 0.08 | 0.90 | 0.16 |
7 | Submerged Aquatic Vascular (Sparganium) | −0.23 | 0.04 | 0.97 | 0.05 |
8 | Submerged Aquatic Vascular (Utricularia) | 0.19 | 0.06 | 0.80 | 0.15 |
9 | Submerged and Floating Vascular (Agrostis/Eleocharis) | 0.36 | 0.08 | 0.45 | 0.19 |
10 | Very Sparse Floating Vascular (Nymphoides) | 0.25 | 0.08 | 0.61 | 0.15 |
11 | Sparse Floating Vascular (Nymphoides) | 0.32 | 0.06 | 0.64 | 0.14 |
12 | Dense Floating Vascular (Nymphoides) | 0.41 | 0.05 | 0.60 | 0.14 |
13 | Very Dense Floating Vascular (Nymphoides) | 0.57 | 0.04 | 0.52 | 0.16 |
14 | Persistent Emergent (Phragmites) | 0.32 | 0.07 | 0.64 | 0.12 |
15 | Persistent Emergent (Bare Ground and Carex) | 0.41 | 0.06 | 0.72 | 0.12 |
16 | Persistent Emergent (Equisetum) | 0.54 | 0.04 | 0.71 | 0.14 |
17 | Persistent Emergent (Thatch) | 0.37 | 0.06 | 0.67 | 0.16 |
18 | Persistent Emergent (Carex) | 0.51 | 0.04 | 0.59 | 0.11 |
19 | Persistent Emergent (Calamagrostis) | 0.39 | 0.04 | 0.75 | 0.11 |
20 | Persistent Emergent (Scolochloa) | 0.67 | 0.05 | 0.50 | 0.17 |
21 | Persistent Emergent (Amoria/Galeopsis/Trifolium) | 0.78 | 0.03 | 0.63 | 0.15 |
22 | Scrub-shrub (Salix with Calamagrostis) | 0.66 | 0.06 | 0.24 | 0.11 |
The capabilities of the NDVI and image texture in differentiating wetland habitats were evaluated similarly (
Table 5). In general, water bodies (Classes 1 to 5) and aquatic bed with submerged vascular vegetation (Classes 6 to 8) demonstrated a negative or approximately zero NDVI value. Aquatic beds with floating vascular vegetation (Classes 9 to 11) and some emergent wetlands (Classes 14, 17, and 19) had NDVI values <0.4. Aquatic beds with dense floating vascular vegetation (Classes 12 and 13) and other emergent wetlands (Classes 15, 16, and 18) had NDVI values ranging between 0.4 and 0.6, while persistent emergent and scrub-shrub wetlands (Classes 20 to 22) had NDVI values >0.65. The NDVI layer contributed to the improvement of the classification mainly through better distinguishing of aquatic habitats with floating vegetation (
i.e., those with moderate NDVI values) in contrast to aquatic beds with submerged vegetation or open water bodies (
i.e., those with low NDVI values; see
Figure 3). When the NDVI layer was included along with the eight spectral bands in the classification, the overall classification accuracy increased from 82.9% to 83.9% (see
Table 4). In addition, inclusion of the NDVI improved both the UA and PA for Class 10 (13.8% and 5.1%, respectively) and Class 11 (8.7% and 10.2%, respectively). The UA for Class 2 improved by 9.6% and PA for Class 19 improved by 10.9%.
Water surfaces of lakes, streams (Classes 1 to 4), and aquatic beds with submerged vegetation (
i.e., Classes 6 to 8) had smooth textures, with homogeneity values >0.7 (see
Table 5). Some types of emergent herbaceous stands (Classes 14 to 17, 19, and 21) also had quite fine and smooth textures, and their homogeneity values were >0.6. Scrub-shrub covered areas (Class 22) had coarse and rough textures with the lowest homogeneity value (0.24), strongly contrasting with the surrounding emergent herbaceous wetlands (see
Figure 4). Class 9 (Submerged and Floating Vascular—
Agrostis/
Eleocharis) and Class 20 (Persistent Emergent—
Scolochloa) also had relatively coarse and rough textures resulting in low homogeneity values. The inclusion of image texture with the eight spectral bands increased the overall classification accuracy by 2% (see
Table 4), but most greatly affected the PA for Class 22 (Scrub-Shrub), which improved by 12% (from 75.7% to 87.7%). Both the PA and UA for Class 21 (Persistent Emergent—
Amoria/
Galeopsis/
Trifolium) and Class 19 (Persistent Emergent—
Calamagrostis) were also considerably improved, suggesting that image texture plays a critical role in discriminating scrub-shrub habitats from emergent herbaceous vegetation and open waters.