Spatial and temporal patterns of aboveground net primary productivity (ANPP) along two freshwater-estuarine transects in the Florida Coastal Everglades

We present here a 4-year dataset (2001–2004) on the spatial and temporal patterns of aboveground net primary production (ANPP) by dominant primary producers (sawgrass, periphyton, mangroves, and seagrasses) along two transects in the oligotrophic Florida Everglades coastal landscape. The 17 sites of the Florida Coastal Everglades Long Term Ecological Research (FCE LTER) program are located along fresh-estuarine gradients in Shark River Slough (SRS) and Taylor River/C-111/Florida Bay (TS/Ph) basins that drain the western and southern Everglades, respectively. Within the SRS basin, sawgrass and periphyton ANPP did not differ significantly among sites but mangrove ANPP was highest at the site nearest the Gulf of Mexico. In the southern Everglades transect, there was a productivity peak in sawgrass and periphyton at the upper estuarine ecotone within Taylor River but no trends were observed in the C-111 Basin for either primary producer. Over the 4 years, average sawgrass ANPP in both basins ranged from 255 to 606 g m−2 year−1. Average periphyton productivity at SRS and TS/Ph was 17–68 g C m−2 year−1 and 342–10371 g C m−2 year−1, respectively. Mangrove productivity ranged from 340 g m−2 year−1 at Taylor River to 2208 g m−2 year−1 at the lower estuarine Shark River site. Average Thalassia testudinum productivity ranged from 91 to 396 g m−2 year−1 and was 4-fold greater at the site nearest the Gulf of Mexico than in eastern Florida Bay. There were no differences in periphyton productivity at Florida Bay. Interannual comparisons revealed no significant differences within each primary producer at either SRS or TS/Ph with the exception of sawgrass at SRS and the C−111 Basin. Future research will address difficulties in assessing and comparing ANPP of different primary producers along gradients as well as the significance of belowground production to the total productivity of this ecosystem.

Soil carbon dynamics in primary and secondary tropical forests in Colombia

Soils play a central role in the dynamics of biospheric carbon and in climate change. They contain the largest carbon stock of terrestrial ecosystems and return to the atmosphere a significant proportion of carbon fixed by photosynthesis. Soils of tropical forests are tremendously important in the carbon cycle because they receive the largest organic matter inputs, they have the largest respiration rates, and they are among the largest carbon reservoirs among world soils. This research assesses the main components of the soil carbon dynamics in primary (PF) and secondary (SF) tropical forests in Colombia. I evaluated the production, stocks, and decomposition rates of aboveground detritus as well as the stocks, growth, mortality, and decomposition of fine roots in these two forest types. Soil carbon outputs were evaluated as total soil, heterotrophic, and root respiration. The stocks of soil organic carbon down to 4 m deep in these two cover types and in degraded pastures (PAS) were also evaluated. ^ Soil inputs of organic carbon from above and belowground sources were lower in SF than in PF. Litterfall in SF was 58% and production of fine root detritus was 60% of that in PF. When production of woody detritus and palm fronds was considered, the difference between these forest types was even larger. However, outputs of mineral carbon through heterotrophic soil respiration were similar; in SF they equaled 97% of those in PF. As a result, soil carbon balance was positive in PF and negative in SF. Despite that soil carbon balances suggest that soils of SF are losing carbon, soil carbon stocks of SF were higher than of degraded pastures, suggesting that they have already started to recover soil carbon stocks lost under degraded pastures. This discrepancy can be partially explained by the effect of drier conditions on heterotrophic soil respiration as a consequence of a moderate El Niño event during the period of soil respiration measurements. The positive carbon balance in soils of PF despite the El Niño event, suggests that soils of PF accumulated about 664 Kg C ha−1 yr−1. Therefore, soil carbon dynamics mainly depended on successional status of vegetation and on climatic conditions. ^

Biogeochemical factors affecting the fate and transport of mercury in the terrestrial environment at Oak Ridge, Tennessee

Mercury (Hg) contamination problem in the United Sates has been an important issue due to its potential threat to human and ecological health. This thesis presents a study of two Hg-contaminated sites along the East Fork Poplar Creek (EFPC) at Oak Ridge. The top soils from the terrestrial areas, along with the soils from three vertical soil horizons at the EFPC bank were sampled and analyzed for total-Hg (THg), methyl-Hg, total-organic-carbon (TOC), and pH. The stream bank soils were also analyzed for the stable-Hg-isotopes (198Hg, 199Hg, 200Hg, 201Hg, and 202Hg). Furthermore, some of the soil samples (n=7) from the same study sites were investigated for phytoavailability of mercury as measured by degree of Hg translocation in aboveground biomass of Impatiens walleriana plants grown in the soils.^ The results showed a significant difference (p<0.001) in THg concentrations for the forest soils (42.40±4.98 mg/kg) and the grassland soils (8.71±2.30 mg/kg). The higher THg and methyl-Hg concentrations were commensurate with the higher TOC content in the soils (p<0.001). Also, the THg concentrations for the upstream site was higher (129.08±34.14 mg/kg) than the downstream site (24.31±3.47 mg/kg). The two sites also differed in their stable Hg isotope compositions (p<0.001 for δ199Hg). The stable isotope analysis indicated the increased level of mass dependent isotopic fractionation with increasing depths along the EFPC bank. The difference between the two study sites was also prominent in case of the Hg uptake by the plants, with higher Hg uptake from the upstream soils compared to that from the downstream soils. A significant correlation, r=0.93 p<0.01, was observed between the Hg uptake and the soil-THg concentrations. THg was higher in the leaves (1161.87±310.01 μg/kg) than in the flowers (206.13±55.23 μg/kg) or the stems (634.54±403.35μg/kg). ^ The level of Hg contamination increased with decreasing distance from the point source and was highly influenced by plants/microbes, soil-organic-content, and Hg-speciation. The isotopic study indicated the existence of an additional Hg source in the EFPC watershed, possibly atmospheric Hg-deposition. These findings are worth taking into account while planning any Hg remediation effort and developing Hg loading criteria as per the National Pollutant Discharge Elimination System (NPDES) Program.^

Long-Term Effects of Adding Nutrients to an Oligotrophic Coastal Environment

Management of ecological disturbances requires an understanding of the time scale and dynamics of community responses to disturbance events. To characterize long-term seagrass bed responses to nutrient enrichment, we established six study sites in Florida Bay, USA. In 24 plots (0.25 m2) at each site, we regularly added nitrogen (N) and phosphorus (P) in a factorial design for 7 years. Five of the six sites exhibited strong P limitation. Over the first 2 years, P enrichment increased Thalassia testudinum cover in the three most P-limited sites. After 3 years, Halodule wrightii began to colonize many of the P-addition plots, but the degree of colonization was variable among sites, possibly due to differences in the supply of viable propagules. Thalassia increased its allocation to aboveground tissue in response to P enrichment; Halodule increased in total biomass but did not appear to change its aboveground: belowground tissue allocation. Nutrient enrichment did not cause macroalgal or epiphytic overgrowth of the seagrass. Nitrogen retention in the study plots was variable but relatively low, whereas phosphorus retention was very high, often exceeding 100% of the P added as fertilizer over the course of our experiments. Phosphorus retentions exceeding 100% may have been facilitated by increases in Thalassia aboveground biomass, which promoted the settlement of suspended particulate matter containing phosphorus. Our study demonstrated that lowintensity press disturbance events such as phosphorus enrichment can initiate a slow, ramped successional process that may alter community structure over many years.

Responses of sawgrass and spikerush to variation in hydrologic drivers and salinity in Southern Everglades marshes

Aboveground net primary production (ANPP) by the dominant macrophyte and plant community composition are related to the changing hydrologic environment and to salinity in the southern Everglades, FL, USA. We present a new non-destructive ANPP technique that is applicable to any continuously growing herbaceous system. Data from 16 sites, collected from 1998 to 2004, were used to investigate how hydrology and salinity controlled sawgrass (Cladium jamaicense Crantz.) ANPP. Sawgrass live biomass showed little seasonal variation and annual means ranged from 89 to 639 gdw m)2. Mortality rates were 20–35% of live biomass per 2 month sampling interval, for biomass turnover rates of 1.3–2.5 per year. Production by C. jamaicense was manifest primarily as biomass turnover, not as biomass accumulation. Rates typically ranged from 300 to 750 gdw m)2 year)1, but exceeded 1000 gdw m)2 year)1 at one site and were as high as 750 gdw m)2 year)1 at estuarine ecotone sites. Production was negatively related to mean annual water depth, hydroperiod, and to a variable combining the two (depth-days). As water depths and hydroperiods increased in our southern Everglades study area, sawgrass ANPP declined. Because a primary restoration goal is to increase water depths and hydroperiods for some regions of the Everglades, we investigated how the plant community responded to this decline in sawgrass ANPP. Spikerush (Eleocharis sp.) was the next most prominent component of this community at our sites, and 39% of the variability in sawgrass ANPP was explained by a negative relationship with mean annual water depth, hydroperiod, and Eleocharis sp. density the following year. Sawgrass ANPP at estuarine ecotone sites responded negatively to salinity, and rates of production were slow to recover after high salinity years. Our results suggest that ecologists, managers, and the public should not necessarily interpret a decline in sawgrass that may result from hydrologic restoration as a negative phenomenon.

Ecological effects of low-level phosphorus additions on two plant communities in a neotropical freshwater wetland ecosystem

We conducted a low-level phosphorus (P) enrichment study in two oligotrophic freshwater wetland communities (wet prairies [WP] and sawgrass marsh [SAW]) of the neotropical Florida Everglades. The experiment included three P addition levels (0, 3.33, and 33.3 mg P m−2 month−1), added over 2 years, and used in situ mesocosms located in northeastern Everglades National Park, Fla., USA. The calcareous periphyton mat in both communities degraded quickly and was replaced by green algae. In the WP community, we observed significant increases in net aboveground primary production (NAPP) and belowground biomass. Aboveground live standing crop (ALSC) did not show a treatment effect, though, because stem turnover rates of Eleocharis spp., the dominant emergent macrophyte in this community, increased significantly. Eleocharis spp. leaf tissue P content decreased with P additions, causing higher C:P and N:P ratios in enriched versus unenriched plots. In the SAW community, NAPP, ALSC, and belowground biomass all increased significantly in response to P additions. Cladium jamaicense leaf turnover rates and tissue nutrient content did not show treatment effects. The two oligotrophic communities responded differentially to P enrichment. Periphyton which was more abundant in the WP community, appeared to act as a P buffer that delayed the response of other ecosystem components until after the periphyton mat had disappeared. Periphyton played a smaller role in controlling ecosystem dynamics and community structure in the SAW community. Our data suggested a reduced reliance on internal stores of P by emergent macrophytes in the WP that were exposed to P enrichment. Eleocharis spp. rapidly recycled P through more rapid aboveground turnover. In contrast, C. jamaicense stored added P by initially investing in belowground biomass, then shifting growth allocation to aboveground tissue without increasing leaf turnover rates. Our results suggest that calcareous wetland systems throughout the Caribbean, and oligotrophic ecosystems in general, respond rapidly to low-level additions of their limiting nutrient.

Responses of seagrass communities to fertilization along a gradient of relative availability of nitrogen and phosphorus in a carbonate environment

Patterns of relative nutrient availability in south Florida suggest spatial differences regarding the importance of nitrogen (N) and phosphorus (P) to benthic primary producers. We did a 14-month in situ fertilization experiment to test predictions of N and P limitation in the subtropical nearshore marine waters of the upper Florida Keys. Six sites were divided into two groups (nearshore, offshore) representing the endpoints of an N: P stoichiometric gradient. Twenty-four plots were established at each site with six replicates of each treatment (1N, 1P, 1N1P, control), for a total of 144 experimental plots. The responses of benthic communities to N and P enrichment varied appreciably between nearshore and offshore habitats. Offshore seagrass beds were strongly limited by nitrogen, and nearshore beds were affected by nitrogen and phosphorus. Nutrient addition at offshore sites increased the length and aboveground standing crop of the two seagrasses, Thalassia testudinum and Syringodium filiforme, and growth rates of T. testudinum. Nutrient addition at nearshore sites increased the relative abundance of macroalgae, epiphytes, and sediment microalgae. N limitation of seagrass in this carbonate system was clearly demonstrated. However, added phosphorus was retained in the system more effectively than N, suggesting that phosphorus might have important long-term effects on these benthic communities. The observed species-specific responses to nutrient enrichment underscores the need to monitor all primary producers when addressing questions of nutrient limitation and eutrophication in seagrass communities.

Phosphorus Availability and Salinity Control Productivity and Demography of the Seagrass Thalassia testudinum in Florida Bay

Biomass, net primary productivity (NPP), foliar elemental content, and demography of Thalassia testudinum were monitored in populations from five sites across Florida Bay beginning in January 2001. Sites were selected to take advantage of the spatial variability in phosphorus (P) availability and salinity climates across the bay. Aboveground biomass and NPP of T. testudinum were determined five to six times annually. Short-shoot demography, belowground biomass, and belowground NPP were assessed from a single destructive harvest at each site and short-shoot cohorts were estimated from leaf scar counts multiplied by site-specific leaf production rates. Biomass, relative growth rate (RGR), and overall NPP were positively correlated with P availability. Additionally, a positive correlation between P availability and the ratio of photosynthetic to non-photosynthetic biomass suggests that T. testudinum increases allocation to aboveground biomass as P availability increases. Population turnover increased with P availability, evident in positive correlations of recruitment and mortality rates with P availability. Departures from seasonally modeled estimates of RGR were found to be influenced by salinity, which depressed RGR when below 20 psu or above 40 psu. Freshwater management in the headwaters of Florida Bay will alter salinity and nutrient climates. It is becoming clear that such changes will affect T. testudinum, with likely feedbacks on ecosystem structure, function, and habitat quality.

Phosphorus budgets in Everglades wetland ecosystems: the effects of hydrology and nutrient enrichment

The Florida Everglades is a naturally oligotrophic hydroscape that has experienced large changes in ecosystem structure and function as the result of increased anthropogenic phosphorus (P) loading and hydrologic changes. We present whole-ecosystem models of P cycling for Everglades wetlands with differing hydrology and P enrichment with the goal of synthesizing existing information into ecosystem P budgets. Budgets were developed for deeper water oligotrophic wet prairie/slough (‘Slough’), shallower water oligotrophic Cladium jamaicense (‘Cladium’), partially enriched C. jamaicense/Typha spp. mixture (‘Cladium/Typha’), and enriched Typha spp. (‘Typha’) marshes. The majority of ecosystem P was stored in the soil in all four ecosystem types, with the flocculent detrital organic matter (floc) layer at the bottom of the water column storing the next largest proportion of ecosystem P pools. However, most P cycling involved ecosystem components in the water column (periphyton, floc, and consumers) in deeper water, oligotrophic Slough marsh. Fluxes of P associated with macrophytes were more important in the shallower water, oligotrophic Cladium marsh. The two oligotrophic ecosystem types had similar total ecosystem P stocks and cycling rates, and low rates of P cycling associated with soils. Phosphorus flux rates cannot be estimated for ecosystem components residing in the water column in Cladium/Typha or Typha marshes due to insufficient data. Enrichment caused a large increase in the importance of macrophytes to P cycling in Everglades wetlands. The flux of P from soil to the water column, via roots to live aboveground tissues to macrophyte detritus, increased from 0.03 and 0.2 g P m−2 yr−1 in oligotrophic Slough and Cladium marsh, respectively, to 1.1 g P m−2 yr−1 in partially enriched Cladium/Typha, and 1.6 g P m−2 yr−1 in enriched Typha marsh. This macrophyte translocation P flux represents a large source of internal eutrophication to surface waters in P-enriched areas of the Everglades.

Using soil profiles of seeds and molecular markers as proxies for sawgrass and wet prairie slough vegetation in Shark Slough, Everglades National Park

We measured the abundance of Cladium jamaicense (Crantz) seeds and three biomarkers in freshwater marsh soils in Shark River Slough (SRS), Everglades National Park (ENP) to determine the degree to which these paleoecological proxies reflect spatial and temporal variation in vegetation. We found that C. jamaicense seeds and the biomarkers Paq, total lignin phenols (TLP) and kaurenes analyzed from surface soils were all significantly correlated with extant aboveground C. jamaicense biomass quantified along a vegetation gradient from a C. jamaicense to a wet prairie/slough (WPS) community. Our results also suggest that these individual proxies may reflect vegetation over different spatial scales: Paq and kaurenes correlated most strongly (R 2 = 0.88 and 0.99, respectively) with vegetation within 1 m of a soil sample, while seeds and TLP reflected vegetation 0–20 m upstream of soil samples. These differences in the spatial scale depicted by the different proxies may be complementary in understanding aspects of historic landscape patterning. Soil profiles of short (25 cm) cores showed that downcore variation in C. jamaicense seeds was highly correlated with two of the three biomarkers (Paq, R 2 = 0.84, p<0.005; TLP, R 2 = 0.97, p<0.0001), and all four of the proxies indicated a recent increase in C. jamaicense biomass at the site. Using a preliminary depth-to-age relationship based on matching charcoal peaks with available ENP fire records (1980-present) specific to our coring site, we found that peak-depths in C. jamaicense seed concentration appeared to correspond to recent minimum water levels (e.g., 1989 and 2001), and low seed abundance corresponded to high water levels (e.g., 1995), consistent with the known autecology of C. jamaicense. In summary, the combination of C. jamaicense seeds and biomarkers may be useful for paleoecological reconstruction of vegetation change and ultimately in guaging the success of ongoing efforts to restore historic hydrologic conditions in the South Florida Everglades.

Possible climate change impacts on the hydrological and vegetative character of Everglades National Park, Florida

The increasing threat of global climate change is predicted to have immense influences on ecosystems worldwide, but could be particularly severe to vulnerable wetland environments such as the Everglades. This work investigates the impact global climate change could have on the hydrologic and vegetative makeup of Everglades National Park (ENP) under forecasted emissions scenarios. Using a simple stochastic model of aboveground water levels driven by a fluctuating rainfall input, we link across ENP a location's mean depth and percent time of inundation to the predicted changes in precipitation from climate change. Changes in the hydrologic makeup of ENP are then related to changes in vegetation community composition through the use of relationships developed between two publically available datasets. Results show that under increasing emissions scenarios mean annual precipitation was forecasted to decrease across ENP leading to a marked hydrologic change across the region. Namely, areas were predicted to be shallower in average depth of standing water and inundated less of the time. These hydrologic changes in turn lead to a shift in ENP's vegetative makeup, with xeric vegetative communities becoming more numerous and hydric vegetative communities becoming scarcer. Noticeably, the most widespread of vegetative communities, sawgrass, decreases in abundance under increasing emissions scenarios. These results are an important indicator of the effects climate change may have on the Everglades region and raise important management implications for those seeking to restore this area to its historical hydrologic and vegetative condition.

Allocation of Biomass and Net Primary Productivity of Mangrove Forests along Environmental Gradients in the Florida Coastal Everglades, USA

Vegetation patterns of mangroves in the Florida Coastal Everglades (FCE) result from the interaction of environmental gradients and natural disturbances (i.e., hurricanes), creating an array of distinct riverine and scrub mangroves across the landscape. We investigated how landscape patterns of biomass and total net primary productivity (NPPT), including allocation in above- and below-ground mangrove components, vary inter-annually (2001–2004) across gradients in soil properties and hydroperiod in two distinct FCE basins: Shark River Estuary and Taylor River Slough. We propose that the allocation of belowground biomass and productivity (NPPB) relative to aboveground allocation is greater in regions with P limitation and permanent flooding. Porewater sulfide was significantly higher in Taylor River (1.2 ± 0.3 mM) compared to Shark River (0.1 ± 0.03 mM) indicating the lack of a tidal signature and more permanent flooding in this basin. There was a decrease in soil P density and corresponding increase in soil N:P from the mouth (28) to upstream locations (46–105) in Shark River that was consistent with previous results in this region. Taylor River sites showed the highest P limitation (soil N:P > 60). Average NPPT was double in higher P environments (17.0 ± 1.1 Mg ha−1 yr−1) compared to lower P regions (8.3 ± 0.3 Mg ha−1 yr−1). Root biomass to aboveground wood biomass (BGB:AWB) ratio was 17 times higher in P-limited environments demonstrating the allocation strategies of mangroves under resource limitation. Riverine mangroves allocated most of the NPPT to aboveground (69%) while scrub mangroves showed the highest allocation to belowground (58%). The total production to biomass (P:B) ratios were lower in Shark River sites (0.11 yr−1); whereas in Taylor River sites P:B ratios were higher and more variable (0.13–0.24 yr−1). Our results suggest that the interaction of lower P availability in Taylor River relative to Shark River basin, along with higher sulfide and permanent flooding account for higher allocation of belowground biomass and production, at expenses of aboveground growth and wood biomass. These distinct patterns of carbon partitioning between riverine and scrub mangroves in response to environmental stress support our hypothesis that belowground allocation is a significant contribution to soil carbon storage in forested wetlands across FCE, particularly in P-limited scrub mangroves. Elucidating these biomass strategies will improve analysis of carbon budgets (storage and production) in neotropical mangroves and understanding what conditions lead to net carbon sinks in the tropical coastal zone.

Drivers of Decadal-Scale Change in Southern Everglades Wetland Macrophyte Communities of the Coastal Ecotone

Understanding the relationships between hydrology and salinity and plant community structure and production is critical to allow predictions of wetland responses to altered water management, changing precipitation patterns and rising sea-level. We addressed how salinity, water depth, hydroperiod, canal inflows, and local precipitation control marsh macrophyte aboveground net primary production (ANPP) and structure in the coastal ecotone of the southern Everglades. We contrasted responses in two watersheds - Taylor Slough (TS) and C-111 - systems that have and will continue to experience changes in water management. Based on long-term trajectories in plant responses, we found continued evidence of increasing water levels and length of inundation in the C-111 watershed south of the C-111 canal. We also found strong differentiation among sites in upper TS that was dependent on hydrology. Finally, salinity, local precipitation and freshwater discharge from upstream explained over 80 % of the variance in Cladium ANPP at a brackish water site in TS. Moreover, our study showed that, while highly managed, the TS and C-111 watersheds maintain legacies in spatial pattern that would facilitate hydrologic restoration. Based on the trajectories in Cladium and Eleocharis, shifts in plant community structure could occur within 5–10 years of sustained water management change.

Effect of Hydrologic Restoration on the Habitat of The Cape Sable Seaside Sparrow, Annual Report of 2004-2005

The major activities in Year 3 on ‘Effect of hydrologic restoration on the habitat of the Cape Sable seaside sparrow (CSSS)’ included presentations, field work, data analysis, and report preparation. During this period, we made 4 presentations, two at the CSSS – fire planning workshops at Everglades National Park (ENP), one at the Society of Wetland Scientists’ meeting in Charleston, SC, and a fourth at the Marl Prairie/CSSS performance measure workshop at ENP. We started field work in the third week of January and continued till June 3, 2005. Early in the field season, we completed vegetation surveys along two transects, B and C (~15.1 km). During April and May, vegetation sampling was completed at 199 census sites, bringing to 608 the total number of CSSS census sites with quantitative vegetation data. We updated data sets from all three years, 2003-05, and analyzed them using cluster analysis and ordination as in previous two years. However, instead of weighted averaging, we used weighted-averaging partial least square regression (WA-PLS) model, as this method is considered an improvement over WA for inferring values of environmental variables from biological species composition. We also validated the predictive power of the WA-PLS regression model by applying it to a sub-set of 100 census sites for which hydroperiods were “known” from two sources, i.e., from elevations calculated from concurrent water depth measurements onsite and at nearby water level recorders, and from USGS digital elevation data. Additionally, we collected biomass samples at 88 census sites, and determined live and dead aboveground plant biomass. Using vegetation structure and biomass data from those sites, we developed a regression model that we used to predict aboveground biomass at all transects and census sites. Finally, biomass data was analyzed in relation to hydroperiod and fire frequency.

Buckwheat as a Cover Crop in Florida: Mycorrhizal Status, Soil Analysis, and Economic Assessment

This thesis analyses buckwheat as a cover crop in Florida. The study was designed to demonstrate: soil enrichment with nutrients, mycorrhizal arbuscular fungi interactions, growth in different soil types, temperature limitations in Florida, and economic benefits for farmers. Buckwheat was planted at the FIU organic garden (Miami, FL) in early November and harvested in middle December. After incorporation of buckwheat residues, soil analyses indicated the ability of buckwheat to enrich soil with major nutrients, in particular, phosphorus. Symbiosis with arbuscular mycorrhizal fungi increased inorganic phosphorus uptake and plant growth. Regression analysis on aboveground buckwheat biomass weight and soil characteristics showed that high soil pH was the major limiting factor that affected buckwheat growth. Spatial analysis illustrated that buckwheat could be planted in South Florida throughout the year but might not be planted in North and Central Florida in winter. An economic assessment proved buckwheat to be a profitable cover crop.