Simulating Forest Shade to Study the Developmental Ecology of Tropical Plants: Juvenile Growth in Three Vines in India

Both light quantity and quality affect the development and autoecology of plants under shade conditions, as in the understorey of tropical forests. However, little research has been directed towards the relative contributions of lowered photosynthetic photon flux density (PPFD) versus altered spectral distributions (as indicated by quantum ratios of 660 to 730 nm, or R:FR) of radiation underneath vegetation canopies. A method for constructing shade enclosures to study the contribution of these two variables is described. Three tropical leguminous vine species (Abrus precatorius L., Caesalpinia bondicela Fleming and Mucuna pruriens (L.) DC.) were grown in two shade enclosures with 3-4% of solar PPFD with either the R:FR of sunlight (1.10) or foliage shade (0.33), and compared to plants grown in sunlight. Most species treated with low R:FR differed from those treated with high R:FR in (1) percent allocation to dry leaf weight, (2) internode length, (3) dry stem weight/length, (4) specific leaf weight, (5) leaf size, and (6) chlorophyll a/b ratios. However, these plants did not differ in chlorophyll content per leaf dry weight or area. In most cases the effects of low R:FR and PPFD were additional to those of high R:FR and low PPFD. Growth patterns varied among the three species, but both low PPFD and diminished R:FR were important cues in their developmental responses to light environments. This shadehouse system should be useful in studying the effects of light on the developmental ecology of other tropical forest plants.

Are mangroves in the tropical Atlantic ripe for invasion? Exotic mangrove trees in the forests of South Florida

Abstract Two species of mangrove trees of Indo- Pacific origin have naturalized in tropical Atlantic mangrove forests in South Florida after they were planted and nurtured in botanic gardens. Two Bruguiera gymnorrhiza trees that were planted in the intertidal zone in 1940 have given rise to a population of at least 86 trees growing interspersed with native mangrove species Rhizophora mangle, Avicennia germinans and Laguncularia racemosa along 100 m of shoreline; the population is expanding at a rate of 5.6% year-1. Molecular genetic analyses confirm very low genetic diversity, as expected from a population founded by two individuals. The maximumnumber of alleles at any locus was three, and we measured reduced heterozygosity compared to native-range populations. Lumnitzera racemosa was introduced multiple times during the 1960s and 1970s, it has spread rapidly into a forest composed of native R. mangle, A. germinans, Laguncularia racemosa and Conocarpus erectus and now occupies 60,500 m2 of mangrove forest with stem densities of 24,735 ha-1. We estimate the population growth rate of Lumnitzera racemosa to be between 17 and 23% year-1. Populations of both species of naturalized mangroves are dominated by young individuals. Given the long life and water-dispersed nature of propagules of the two exotic species, it is likely that they have spread beyond our survey area. We argue that the species-depauperate nature of tropical Atlantic mangrove forests and close taxonomic relatives in the more species-rich Indo-Pacific region result in the susceptibility of tropical Atlantic mangrove forests to invasion by Indo-Pacific mangrove species.

Epidermal cells functioning as lenses in leaves of tropical rain-forest shade plants

A ray tracing model has been developed to investigate the possible focusing effects of the convexly curved epidermal cell walls which characterize a number of shade-adapted plants. The model indicates that such focusing occurs, resulting in higher photosynthetic photon flux densities at certain locations within the leaf. It is postulated that there will be a corresponding increase in the rate of photosynthesis. In addition, leaf reflectance measurements indicate that this is generally less for the shade plants compared with sun species and would be advantageous in increasing the efficiency of energy capture. Either effect is important for plants which must survive at extremely low light levels.

Are mangroves in the tropical Atlantic ripe for invasion? Exotic mangrove trees in the forests of South Florida

Two species of mangrove trees of Indo- Pacific origin have naturalized in tropical Atlantic mangrove forests in South Florida after they were planted and nurtured in botanic gardens. Two Bruguiera gymnorrhiza trees that were planted in the intertidal zone in 1940 have given rise to a population of at least 86 trees growing interspersed with native mangrove species Rhizophora mangle, Avicennia germinans and Laguncularia racemosa along 100 m of shoreline; the population is expanding at a rate of 5.6% year-1. Molecular genetic analyses confirm very low genetic diversity, as expected from a population founded by two individuals. The maximumnumber of alleles at any locus was three, and we measured reduced heterozygosity compared to native-range populations. Lumnitzera racemosa was introduced multiple times during the 1960s and 1970s, it has spread rapidly into a forest composed of native R. mangle, A. germinans, Laguncularia racemosa and Conocarpus erectus and now occupies 60,500 m2 of mangrove forest with stem densities of 24,735 ha-1. We estimate the population growth rate of Lumnitzera racemosa to be between 17 and 23% year-1. Populations of both species of naturalized mangroves are dominated by young individuals. Given the long life and water-dispersed nature of propagules of the two exotic species, it is likely that they have spread beyond our survey area. We argue that the species-depauperate nature of tropical Atlantic mangrove forests and close taxonomic relatives in the more species-rich Indo-Pacific region result in the susceptibility of tropical Atlantic mangrove forests to invasion by Indo-Pacific mangrove species.

Factors affecting the nodulation and growth of tropical woody legume seedlings

This study surveys the occurrence of nodulation in woody legume species in Panamá and Costa Rica, describes nodule and root characteristics, and researches host-bacteria specificity, nodulation potential of soils, and the effects of light, added nitrogen, and rhizobia and VA mycorrhizal fungi inoculation on seedling growth. I examined 83 species in 37 genera and found 80% to be nodulated. Percent nodulated species in the Caesalpinioideae, Mimosoideae, and Papilionoideae was 17, 95, and 86, respectively, with no correlation between nodule morphology and tribal classification. Nodules formed mainly at root branch points which supports epidermal breaks as an important rhizobia infection route. More non-nodulated than nodulated species had root hairs. Several species emitted volatile sulfur-containing compounds, including the toxic compound ethylmercaptan, from roots, germinating seeds, and other tissues. These emissions may have an allelopathic action against pathogens, predators, or other plants. In contrast to the general non-specificity of most legumes for rhizobia, Mimosa pigra L. was highly specific and only nodulated in flooded soils. This species' specificity, combined with a limited occurrence of its root nodule bacteria may limit its natural distribution, but its spread as an invasive weed is facilitated when fill material from rivers is deposited in other areas. ^ An experimental light level of 1.5% of full sun completely inhibited seedling nodulation, as do similar naturally low levels in forest understory. In the forest, trees and seedlings were not nodulated. in some soils with suspected high N content. For six experimental species, added N progressively increased seedling growth while decreasing nodule biomass; at the highest level of added N nodulation was completely suppressed. Species and individuals showed variation in nodule biomass at high N applications which may indicate an opportunity for genetic selection for optimal N acquisition. Rhizobia inoculation had a small positive effect on seedling shoot growth, but VA mycorrhiza inoculation overwhelmingly increased seedling size, biomass, and leaf mineral concentration. In lowland tropical forest, VA mycorrhizal colonization appears indispensable for legume nodulation because of the fungus' ability to supply P in deficient soils. This requirement makes the legume-rhizobia-mycorrhiza association obligately tripartite. ^

The effects of climate on the growth and physiology of tropical rainforest trees

Tropical rainforests account for more than a third of global net primary production and contain more than half of the global forest carbon. Though these forests are a disproportionately important component of the global carbon cycle, the relationship between rainforest productivity and climate remains poorly understood. Understanding the link between current climate and rainforest tree stem diameter increment, a major constituent of forest productivity, will be crucial to efforts at modeling future climate and rainforest response to climate change. This work reports the physiological and stem growth responses to micrometeorological and phenological states of ten species of canopy trees in a Costa Rican wet tropical forest at sub-annual time intervals. I measured tree growth using band dendrometers and estimated leaf and reproductive phenological states monthly. Electronic data loggers recorded xylem sap flow (an indicator of photosynthetic rate) and weather at half-hour intervals. An analysis of xylem sap flow showed that physiological responses were independent of species, which allowed me to construct a general model of weather driven sap flow rates. This model predicted more than eighty percent of climate driven sap flow variation. Leaf phenology influenced growth in three of the ten species, with two of these species showing a link between leaf phenology and weather. A combination of rainfall, air temperature, and irradiance likely provided the cues that triggered leaf drop in Dipteryx panamensis and Lecythis ampla. Combining the results of the sap flow model, growth, and the climate measures showed tree growth was correlated to climate, though the majority of growth variation remained unexplained. Low variance in the environmental variables and growth rates likely contributed to the large amount of unexplained variation. A simple model that included previous growth increment and three meteorological variables explained from four to nearly fifty percent of the growth variation. Significant growth carryover existed in six of the ten species, and rainfall was positively correlated to growth in eight of the ten species. Minimum nighttime temperature was also correlated to higher growth rates in five of the species and irradiance in two species. These results indicate that tropical rainforest tree trunks could act as carbon sinks if future climate becomes wetter and slightly warmer. ^

Plant biogeography and conservation on a tropical island: Isla del Coco, Costa Rica

Isla del Coco (Cocos Island) is a small volcanic island located in the Pacific 500 km west of Costa Rica. Three collecting trips to Isla del Coco, in addition to herbarium research, were completed in order to assess the floristic diversity of the island. The current flora of Isla del Coco contains 262 plant species of which 37 (19.4%) are endemic. This study reports 58 species as new to the island. Seventy-one species (27.1%) were identified as introduced by humans. In addition, five potentially invasive plant species are identified. Seven vegetation types are identified on the island: bayshore, coastal cliff, riparian, low elevation humid forest, high elevation cloud forest, landslide and islet. ^ The biogeographic affinities of the native and endemic species are with Central America/northern South America and to a lesser extent, the Caribbean. Endemic species in the genus Epidendrum were investigated to determine whether an insular radiation event had produced two species found on Isla del Coco. Phylogenetic analysis of the internal transcribed spacer (ITS) of nuclear ribosomal DNA was not able to disprove that the endemic species in this genus are not sister species. Molecular biogeographic analyses of ITS sequence data determined that the Isla del Coco endemic species in the genera Epidendrum, Pilea and Psychotria are most closely related to Central American/northern South American taxa. No biogeographical links were found between the floras of Isla del Coco and the Galápagos Islands. ^ The native and endemic plant diversity of Isla del Coco is threatened with habitat degradation by introduced pigs and deer, and to a lesser extent, by exotic plant species. The IUCN Red List and RAREplants criteria were used to assess the extinction threat for the 37 endemic plant taxa found on the island. All of the endemic species are considered threatened with extinction at the Critically Endangered (CR) by the IUCN criteria or either CR or Endangered (EN) using RAREplants methodology. ^

The potential role of plant oxygen and sulphide dynamics in die-off events of the tropical seagrass, Thalassia testudinum

1 Oxygen and sulphide dynamics were examined, using microelectrode techniques, in meristems and rhizomes of the seagrass Thalassia testudinum at three different sites in Florida Bay, and in the laboratory, to evaluate the potential role of internal oxygen variability and sulphide invasion in episodes of sudden die-off. The sites differed with respect to shoot density and sediment composition, with an active die-off occurring at only one of the sites. 2 Meristematic oxygen content followed similar diel patterns at all sites with high oxygen content during the day and hyposaturation relative to the water column during the night. Minimum meristematic oxygen content was recorded around sunrise and varied among sites, with values close to zero at the die-off site. 3 Gaseous sulphide was detected within the sediment at all sites but at different concentrations among sites and within the die-off site. Spontaneous invasion of sulphide into Thalassia rhizomes was recorded at low internal oxygen partial pressure during darkness at the die-off site. 4 A laboratory experiment showed that the internal oxygen dynamics depended on light availability, and hence plant photosynthesis, and on the oxygen content of the water column controlling passive oxygen diffusion from water column to leaves and belowground tissues in the dark. 5 Sulphide invasion only occurred at low internal oxygen content, and the rate of invasion was highly dependent on the oxygen supply to roots and rhizomes. Sulphide was slowly depleted from the tissues when high oxygen partial pressures were re-established through leaf photosynthesis. Coexistence of sulphide and oxygen in the tissues and the slow rate of sulphide depletion suggest that sulphide reoxidation is not biologically mediated within the tissues of Thalassia. 6 Our results support the hypothesis that internal oxygen stress, caused by low water column oxygen content or poor plant performance governed by other environmental factors, allows invasion of sulphide and that the internal plant oxygen and sulphide dynamics potentially are key factors in the episodes of sudden die-off in beds of Thalassia testudinum . Root anoxia followed by sulphide invasion may be a more general mechanism determining the growth and survival of other rooted plants in sulphate-rich aquatic environments.

Forest Resource Islands in a Sub-tropical Marsh: Soil–Site Relationships in Everglades Hardwood Hammocks

Spatial heterogeneity in soils is often characterized by the presence of resource-enriched patches ranging in size from a single shrub to wooded thickets. If the patches persist long enough, the primary constraint on production may transition from one limiting environmental factor to another. Tree islands that are scattered throughout the Florida Everglades basin comprise nutrient-enriched patches, or resource islands, in P-limited oligotrophic marshes. We used principal component analysis and multiple regressions to characterize the belowground environment (soil, hydrology) of one type of tree island, hardwood hammocks, and examined its relationship with the three structural variables (basal area, biomass, and canopy height) indicative of site productivity. Hardwood hammocks in the southern Everglades grow on two distinct soil types. The first, consisting of shallow, organic, relatively low-P soils, is common in the seasonally flooded Marl Prairie landscape. In contrast, hammocks on islands embedded in long hydroperiod marsh have deeper, alkaline, mineral soils with extremely high P concentrations. However, this edaphic variation does not translate simply into differences in forest structure and production. Relative water depth was unrelated to all measures of forest structure and so was soil P, but the non-carbonate component of the mineral soil fraction exhibited a strong positive relationship with canopy height. The development of P-enriched forest resource islands in the Everglades marsh is accompanied by the buildup of a mineral soil; however, limitations on growth in mature islands appear to differ substantively from those that dominate incipient stages in the transformation from marsh to forest. Key words: resource island; tree

Benthic diatom assemblages as indicators of water quality in the Everglades and three tropical karstic wetlands

Limestone-based (karstic) freshwater wetlands of the Everglades, Belize, Mexico, and Jamaica are distinctive in having a high biomass of CaCO3-rich periphyton mats. Diatoms are common components of these mats and show predictable responses to environmental variation, making them good candidates for assessing nutrient enrichment in these naturally ultraoligotrophic wetlands. However, aside from in the Everglades of southern Florida, very little research has been done to document the diatoms and their environmental preferences in karstic Caribbean wetlands, which are increasingly threatened by eutrophication. We identified diatoms in periphyton mats collected during wet and dry periods from the Everglades and similar freshwater karstic wetlands in Belize, Mexico, and Jamaica. We compared diatom assemblage composition and diversity among locations and periods, and the effect of the limiting nutrient, P, on species composition among locations. We used periphyton-mat total P (TP) as a metric of availability. A total of 176 diatom species in 45 genera were recorded from the 4 locations. Twenty-three of these species, including 9 that are considered indicative of Everglades diatom flora, were found in all 4 locations. In Everglades and Caribbean sites, we identified assemblages and indicator species associated with low and high periphyton-mat TP and calculated TP optima and tolerances for each indicator species. TP optima and tolerances of indicator species differed between the Everglades and the Caribbean, but weighted averaging models predicted periphyton-mat TP concentrations from diatom assemblages at Everglades (R2  =  0.56) and Caribbean (R2  =  0.85) locations. These results show that diatoms can be effective indicators of water quality in karstic wetlands of the Caribbean, but application of regionally generated transfer functions to distant sites provides less reliable estimates than locally developed functions.

Mechanisms governing the eyewall replacement cycle in numerical simulations of tropical cyclones

Eyewall replacement cycle (ERC) is frequently observed during the evolution of intensifying Tropical Cyclones (TCs). Although intensely studied in recent years, the underlying mechanisms of ERC are still poorly understood, and the forecast of ERC remains a great challenge. To advance our understanding of ERC and provide insights in improvement of numerical forecast of ERC, a series of numerical simulations is performed to investigate ERCs in TC-like vortices on a f-plane. The simulated ERCs possess key features similar to those observed in real TCs including the formation of a secondary tangential wind maximum associated with the outer eyewall. The Sawyer-Eliassen equation and tangential momentum budget analyses are performed to diagnose the mechanisms underlying the secondary eyewall formation (SEF) and ERC. Our diagnoses reveal crucial roles of outer rainband heating in governing the formation and development of the secondary tangential wind maximum and demonstrate that the outer rainband convection must reach a critical strength relative to the eyewall before SEF and the subsequent ERC can occur. A positive feedback among low-level convection, acceleration of tangential winds in the boundary layer, and surface evaporation that leads to the development of ERC and a mechanism for the demise of inner eyewall that involves interaction between the transverse circulations induced by eyewall and outer rainband convection are proposed. The tangential momentum budget indicates that the net tendency of tangential wind is a small residual resultant from a large cancellation between tendencies induced by the resolved and sub-grid scale (SGS) processes. The large SGS contribution to the tangential wind budget explains different characteristics of ERC shown in previous numerical studies and poses a great challenge for a timely correct forecast of ERC. The sensitivity experiments show that ERCs are strongly subjected to model physics, vortex radial structure and background wind. The impact of model physics on ERC can be well understood with the interaction among eyewall/outer rainband heating, radilal inflow in the boundary layer, surface layer turbulent processes, and shallow convection in the moat. However, further investigations are needed to fully understand the exhibited sensitivities of ERC to vortex radial structure and background wind.

Evidence of climate variability and tropical cyclone activity from diatom assemblage dynamics in coastal southwest Florida

Estuaries are dynamic on many spatial and temporal scales. Distinguishing effects of unpredictable events from cyclical patterns can be challenging but important to predict the influence of press and pulse drivers in the face of climate change. Diatom assemblages respond rapidly to changing environmental conditions and characterize change on multiple time scales. The goals of this research were to 1) characterize diatom assemblages in the Charlotte Harbor watershed, their relationships with water quality parameters, and how they change in response to climate; and 2) use assemblages in sediment cores to interpret past climate changes and tropical cyclone activity. ^ Diatom assemblages had strong relationships with salinity and nutrient concentrations, and a quantitative tool was developed to reconstruct past values of these parameters. Assemblages were stable between the wet and dry seasons, and were more similar to each other than to assemblages found following a tropical cyclone. Diatom assemblages following the storm showed a decrease in dispersion among sites, a pattern that was consistent on different spatial scales but may depend on hydrological management regimes. ^ Analysis of sediment cores from two southwest Florida estuaries showed that locally-developed diatom inference models can be applied with caution on regional scales. Large-scale climate changes were suggested by environmental reconstructions in both estuaries, but with slightly different temporal pacing. Estimates of salinity and nutrient concentrations suggested that major hydrological patterns changed at approximately 5.5 and 3 kyrs BP. A highly temporally-resolved sediment core from Charlotte Harbor provided evidence for past changes that correspond with known climate records. Diatom assemblages had significant relationships with the three-year average index values of the Atlantic Multidecadal Oscillation and the El Niño Southern Oscillation. Assemblages that predicted low salinity and high total phosphorus also had the lowest dispersion and corresponded with some major storms in the known record, which together may provide a proxy for evidence of severe storms in the paleoecological record. ^

Mechanisms Governing the Eyewall Replacement Cycle in Numerical Simulations of Tropical Cyclones

Eyewall replacement cycle (ERC) is frequently observed during the evolution of intensifying Tropical Cyclones (TCs). Although intensely studied in recent years, the underlying mechanisms of ERC are still poorly understood, and the forecast of ERC remains a great challenge. To advance our understanding of ERC and provide insights in improvement of numerical forecast of ERC, a series of numerical simulations is performed to investigate ERCs in TC-like vortices on a f-plane. The simulated ERCs possess key features similar to those observed in real TCs including the formation of a secondary tangential wind maximum associated with the outer eyewall. The Sawyer-Eliassen equation and tangential momentum budget analyses are performed to diagnose the mechanisms underlying the secondary eyewall formation (SEF) and ERC. Our diagnoses reveal crucial roles of outer rainband heating in governing the formation and development of the secondary tangential wind maximum and demonstrate that the outer rainband convection must reach a critical strength relative to the eyewall before SEF and the subsequent ERC can occur. A positive feedback among low-level convection, acceleration of tangential winds in the boundary layer, and surface evaporation that leads to the development of ERC and a mechanism for the demise of inner eyewall that involves interaction between the transverse circulations induced by eyewall and outer rainband convection are proposed. The tangential momentum budget indicates that the net tendency of tangential wind is a small residual resultant from a large cancellation between tendencies induced by the resolved and sub-grid scale (SGS) processes. The large SGS contribution to the tangential wind budget explains different characteristics of ERC shown in previous numerical studies and poses a great challenge for a timely correct forecast of ERC. The sensitivity experiments show that ERCs are strongly subjected to model physics, vortex radial structure and background wind. The impact of model physics on ERC can be well understood with the interaction among eyewall/outer rainband heating, radilal inflow in the boundary layer, surface layer turbulent processes, and shallow convection in the moat. However, further investigations are needed to fully understand the exhibited sensitivities of ERC to vortex radial structure and background wind.