Leaf morphology scales multi-annual trends in nutrient cycling and leaf, flower, and fruiting phenology among species in the sub-tropical hardwood forests of the northern Florida Keys

The subtropical hardwood forests of southern Florida are formed by 120 frost-sensitive, broadleaved angiosperm species that range throughout the Caribbean. Previous work on a series of small sized forest component patches of a 20 km2, forest preserve in northern Key Largo indicate that a shift in species composition was associated with a 100 year forest developmental sequence, and this shift was associated with an increasingly evergreen canopy. This document investigates the underlying differences of the biology of trees that live in this habitat, and is specifically focused on the impact of leaf morphology on changing nutrient cycling patterns. Measurements of the area, thickness, dry mass, nutrient content and longevity of several leaves from 3-4 individuals of ten species were conducted in combination with a two-year leaf litter collection and nutrient analysis to determine that species with thicker, denser leaves cycled scarce nutrients up to 2-3 times more efficiently than thin leaved tree species, and the leaf thickness/density index predicts role in forest development in a parallel direction as the index predicts nutrient cycling efficiency. A three year set of observations on the relative abundance of new leaves, flowers and fruits of the same tree species provides an opportunity to evaluate the consequences the leaf morphology/nutrient cycling/forest development relationship to forest habitat quality. Results of the three documents support a mechanistic link between forest development and nutrient cycling, and suggests that older forests are likely to be better habitats based on the availability of valuable forest products like new leaves, flowers, and fruits throughout the year.

Importance of water source in controlling leaf leaching losses in a dwarf red mangrove (Rhizophora mangle L.) wetland

The southern Everglades mangrove ecotone is characterized by extensive dwarf Rhizophora mangle L. shrub forests with a seasonally variable water source (Everglades – NE Florida Bay) and residence times ranging from short to long. We conducted a leaf leaching experiment to understand the influence that water source and its corresponding water quality have on (1) the early decay of R. mangle leaves and (2) the early exchange of total organic carbon (TOC) and total phosphorus (TP) between leaves and the water column. Newly senesced leaves collected from lower Taylor River (FL) were incubated in bottles containing water from one of three sources (Everglades, ambient mangrove, and Florida Bay) that spanned a range of salinity from 0 to 32‰, [TOC] from 710 to 1400 μM, and [TP] from 0.17 to 0.33 μM. We poisoned half the bottles in order to quantify abiotic processes (i.e., leaching) and assumed that non-poisoned bottles represented both biotic (i.e., microbial) and abiotic processes. We sacrificed bottles after 1,2, 5, 10, and 21 days of incubation and quantified changes in leaf mass and changes in water column [TOC] and [TP]. We saw 10–20% loss of leaf mass after 24 h—independent of water treatment—that leveled off by Day 21. After 3 weeks, non-poisoned leaves lost more mass than poisoned leaves, and there was only an effect of salinity on mass loss in poisoned incubations—with greatest leaching-associated losses in Everglades freshwater. Normalized concentrations of TOC in the water column increased by more than two orders of magnitude after 21 days with no effect of salinity and no difference between poisoned and non-poisoned treatments. However, normalized [TP] was lower in non-poisoned incubations as a result of immobilization by epiphytic microbes. This immobilization was greatest in Everglades freshwater and reflects the high P demand in this ecosystem. Immobilization of leached P in mangrove water and Florida Bay water was delayed by several days and may indicate an initial microbial limitation by labile C during the dry season.

Temporally dependent C, N, and P dynamics associated with the decay of Rhizophora mangle L. leaf litter in oligotrophic mangrove wetlands of the Southern Everglades

We performed two litter decomposition experiments using nearly-senesced red mangrove (Rhizophora mangle L.) leaves collected from an Everglades dwarf mangrove wetland to understand the short-term (3 weeks) and long-term (1 year) changes in mass, as well as C-, N-, and P-content of decomposing leaf litter. We expected that leaves decomposing in this oligotrophic environment would be short-term sources of C, N, and P, but potential long-term sinks for N and P. In May 1998, we conducted a 3-week leaching experiment, incubating fresh, individual leaves in seawater for up to 21 days. From May 1997 to May 1998, leaf litter in mesh bags decomposed on the forest floor at two dwarf mangrove sites. Leaching accounted for about 33% loss of dry mass from R. mangle leaves after 3 weeks. Leaching losses were rapid, peaking by day 2, and large, with leachate concentrations of total organic carbon (TOC) and total phosphorus (TP) increasing by more than an order of magnitude after 3 weeks. Mean leaf C:N increased from 105 to 115 and N:P increased from a mean of 74 to 95 after 21 days, reflecting the relatively large leaching losses of N and P. Loss of mass in the litterbags leveled off after 4 months, with roughly 60%dry mass remaining (DMR) after nearly 1 year of decomposition. The mass of carbon in each litterbag declined significantly after 361 days, but the mass of nitrogen and phosphorus doubled, indicating long-term accumulation of these constituents into the detritus. Subsequently, the leaf C:N ratio dropped significantly from 90 to 34 after 361 days. Following an initial 44-day increase, leaf N:P decreased from 222 to 144, reflecting high accumulation of P relative to N. A review of several estuarine macrophyte decomposition studies reveals a trend in nitrogen accumulation through time regardless of site, but suggests no clear pattern for C and P. We believe that the increase in litter P observed in this study was indicative of the P-limited status of the greater Everglades ecosystem and that decomposing mangrove litter may represent a substantial phosphorus pool in the system.

Leaf gas exchange characteristics of three neotropical mangrove species in response to varying hydroperiod

We determined how different hydroperiods affected leaf gas exchange characteristics of greenhouse-grown seedlings (2002) and saplings (2003) of the mangrove species Avicennia germinans (L.) Stearn., Laguncularia racemosa (L.) Gaertn. f., and Rhizophora mangle L. Hydroperiod treatments included no flooding (unflooded), intermittent flooding (intermittent), and permanent flooding (flooded). Plants in the intermittent treatment were measured under both flooded and drained states and compared separately. In the greenhouse study, plants of all species maintained different leaf areas in the contrasting hydroperiods during both years. Assimilation-light response curves indicated that the different hydroperiods had little effect on leaf gas exchange characteristics in either seedlings or saplings. However, short-term intermittent flooding for between 6 and 22 days caused a 20% reduction in maximum leaf-level carbon assimilation rate, a 51% lower light requirement to attain 50% of maximum assimilation, and a 38% higher demand from dark respiration. Although interspecific differences were evident for nearly all measured parameters in both years, there was little consistency in ranking of the interspecific responses. Species by hydroperiod interactions were significant only for sapling leaf area. In a field study, R. mangle saplings along the Shark River in the Everglades National Park either demonstrated no significant effect or slight enhancement of carbon assimilation and water-use efficiency while flooded. We obtained little evidence that contrasting hydroperiods affect leaf gas exchange characteristics of mangrove seedlings or saplings over long time intervals; however, intermittent flooding may cause short-term depressions in leaf gas exchange. The resilience of mangrove systems to flooding, as demonstrated in the permanently flooded treatments, will likely promote photosynthetic and morphological adjustment to slight hydroperiod shifts in many settings.

Effect of leaf-cutting ant nests on plant growth in an oligotrophic Amazon rain forest

This study examined whether high nutrient concentrations associated with leaf-cutting ant nests influence plant growth and plant water relations in Amazon rain forests. Three nests of Atta cephalotes were selected along with 31 Amaioua guianensis and Protium sp. trees that were grouped into trees near and distant (>10 m) from nests. A 15N leaf-labelling experiment confirmed that trees located near nests accessed nutrients from nests. Trees near nests exhibited higher relative growth rates (based on stem diameter increases) on average compared with trees further away; however this was significant for A. guianensis (near nest 0.224 y−1 and far from nest 0.036 y−1) but not so for Protium sp. (0.146 y−1 and 0.114 y−1 respectively). Water relations were similarly species-specific; for A. guianensis, near-nest individuals showed significantly higher sap flow rates (16 vs. 5 cm h−1), higher predawn/midday water potentials (−0.66 vs. −0.98 MPa) and lower foliar δ13C than trees further away indicating greater water uptake in proximity to the nests while the Protium sp. showed no significant difference except for carbon isotopes. This study thus shows that plant response to high nutrient concentrations in an oligotrophic ecosystem varies with species. Lower seedling abundance and species richness on nests as compared with further away suggests that while adult plants access subterranean nutrient pools, the nest surfaces themselves do not encourage plant establishment and growth.

Leaf Gas Exchange and Nutrient Use Efficiency Help Explain the Distribution of Two Neotropical Mangroves under Contrasting Flooding and Salinity

Rhizophora mangle and Laguncularia racemosa cooccur along many intertidal floodplains in the Neotropics. Their patterns of dominance shift along various gradients, coincident with salinity, soil fertility, and tidal flooding. We used leaf gas exchange metrics to investigate the strategies of these two species in mixed culture to simulate competition under different salinity concentrations and hydroperiods. Semidiurnal tidal and permanent flooding hydroperiods at two constant salinity regimes (10 g L−1 and 40 g L−1) were simulated over 10 months. Assimilation ( ), stomatal conductance ( ), intercellular CO2 concentration ( ), instantaneous photosynthetic water use efficiency (PWUE), and photosynthetic nitrogen use efficiency (PNUE) were determined at the leaf level for both species over two time periods. Rhizophora mangle had significantly higher PWUE than did L. racemosa seedlings at low salinities; however, L. racemosa had higher PNUE and and, accordingly, had greater intercellular CO2 (calculated) during measurements. Both species maintained similar capacities for A at 10 and 40 g L−1 salinity and during both permanent and tidal hydroperiod treatments. Hydroperiod alone had no detectable effect on leaf gas exchange. However, PWUE increased and PNUE decreased for both species at 40 g L−1 salinity compared to 10 g L−1. At 40 g L−1 salinity, PNUE was higher for L. racemosa than R. mangle with tidal flooding. These treatments indicated that salinity influences gas exchange efficiency, might affect how gases are apportioned intercellularly, and accentuates different strategies for distributing leaf nitrogen to photosynthesis for these two species while growing competitively.

Leaf Functional Traits and Forest Structure of Tropical Dry Forest Species Along a Rainfall Gradient in Florida and Puerto Rico

This study examined how different rainfall regimes affect a set of leaf functional traits related to plant stress and forest structure in tropical dry forest (TDF) species on limestone substrate. One hundred fifty eight individuals of four tree species were sampled in six ecological sites in south Florida and Puerto Rico, ranging in mean annual rainfall from 858 to 1933 mm yr-1. Leaf nitrogen content, specific leaf area (SLA), and N:P ratio of evergreen species, but not deciduous species, responded positively to increasing rainfall. Phosphorus content was unaffected in both groups. Canopy height and basal area reached maxima of 10.3 m and 31.4 m2 ha-1, respectively, at 1168 mm annual rainfall. Leaf traits reflected soil properties only to a small extent. This led us to the conclusion that water is a major limiting factor in TDF and some species that comprise TDF ecosystems are limited by nitrogen in limestone sites with less than ~1012 mm rainfall, but organismal, biological and/or abiotic forces other than rainfall control forest structure in moister sites.

Plant leaf wax (n-alkane) data from sediment core MD96-2048

In this study we investigate Pleistocene vegetation and climate change in southern East Africa by examining plant leaf waxes in a marine sediment core that receives terrestrial runoff from the Limpopo River. The plant leaf wax records are compared to a multi-proxy sea surface temperature (SST) record and pollen assemblage data from the same site. We find that Indian Ocean SST variability, driven by high-latitude obliquity, exerted a strong control on the vegetation of southern East Africa during the past 800,000 yr. Interglacial periods were characterized by relatively wetter and warmer conditions, increased contributions of C3 vegetation, and higher SST, whereas glacial periods were marked by cooler and arid conditions, increased contributions of C4 vegetation, and lower SST. We find that Marine Isotope Stages (MIS) 5e, 11c, 15e and 7a-7c are strongly expressed in the plant leaf wax records but MIS 7e is absent while MIS 9 is rather weak. Our plant leaf wax records also record the climate transition associated with the Mid-Brunhes Event (MBE) suggesting that the pre-MBE interval (430-800 ka) was characterized by higher inputs from grasses in comparison to relatively higher inputs from trees in the post-MBE interval (430 to 0 ka). Differences in vegetation and SST of southern East Africa between the pre- and post-MBE intervals appear to be related to shifts in the location of the Subtropical Front. Comparison with vegetation records from tropical East Africa indicates that the vegetation of southern East Africa, while exhibiting glacial-interglacial variability and notable differences between the pre- and post-MBE portions of the record, likely did not experience such dramatic extremes as occurred to the north at Lake Malawi.

Leaf Traits, Neighbors, and Abiotic Factors: Ways That Context Can Mediate the Impact of Invasive Species on Nitrogen Cycling

Species invasions are more prevalent than ever before. While the addition of a species can dramatically change critical ecosystem processes, factors that mediate the direction and magnitude of those impacts have received less attention. A better understanding of the factors that mediate invasion impacts on ecosystem functioning is needed in order to target which exotic species will be most harmful and which systems are most vulnerable. The role of invasion on nitrogen (N) cycling is particularly important since N cycling controls ecosystem services that provision human health, e.g. nutrient retention and water quality.

We conducted a meta-analysis and in-depth studies focused on the invasive grass species, Microstegium vimineum, to better understand how (i) plant characteristics, (ii) invader abundance and neighbor identity, and (iii) environmental conditions mediate the impacts of invasion on N pools and fluxes. The results of our global meta-analysis support the concept that invasive species and reference community traits such as leaf %N and leaf C:N are useful for understanding invasion impacts on soil N cycling, but that trait dissimilarities between invaded and reference communities are most informative. Regarding the in-depth studies of Microstegium, we did not find evidence to suggest that invasion increases net nitrification as other studies have shown. Instead, we found that an interaction between its abundance and the neighboring plant identify were important for determining soil nitrate concentrations and net nitrification rates in the greenhouse. In field, we found that variability in environmental conditions mediated the impact of Microstegium invasion on soil N pools and fluxes, primarily net ammonification, between sites through direct, indirect, and interactive pathways. Notably, we detected a scenario in which forest openness has a negative direct effect and indirect positive effect on ammonification in sites with high soil moisture and organic matter. Collectively, our findings suggest that dissimilarity in plant community traits, neighbor identity, and environmental conditions can be important drivers of invasion impacts on ecosystem N cycling and should be considered when evaluating the ecosystem impacts of invasive species across heterogeneous landscapes.

Host genotype and age shape the leaf and root microbiomes of a wild perennial plant.

Bacteria living on and in leaves and roots influence many aspects of plant health, so the extent of a plant's genetic control over its microbiota is of great interest to crop breeders and evolutionary biologists. Laboratory-based studies, because they poorly simulate true environmental heterogeneity, may misestimate or totally miss the influence of certain host genes on the microbiome. Here we report a large-scale field experiment to disentangle the effects of genotype, environment, age and year of harvest on bacterial communities associated with leaves and roots of Boechera stricta (Brassicaceae), a perennial wild mustard. Host genetic control of the microbiome is evident in leaves but not roots, and varies substantially among sites. Microbiome composition also shifts as plants age. Furthermore, a large proportion of leaf bacterial groups are shared with roots, suggesting inoculation from soil. Our results demonstrate how genotype-by-environment interactions contribute to the complexity of microbiome assembly in natural environments.

Bridging the Scale Gap from Leaf to Canopy in Biosphere-Atmosphere Gas and Particle Exchanges

Terrestrial ecosystems, occupying more than 25% of the Earth's surface, can serve as

`biological valves' in regulating the anthropogenic emissions of atmospheric aerosol

particles and greenhouse gases (GHGs) as responses to their surrounding environments.

While the signicance of quantifying the exchange rates of GHGs and atmospheric

aerosol particles between the terrestrial biosphere and the atmosphere is

hardly questioned in many scientic elds, the progress in improving model predictability,

data interpretation or the combination of the two remains impeded by

the lack of precise framework elucidating their dynamic transport processes over a

wide range of spatiotemporal scales. The diculty in developing prognostic modeling

tools to quantify the source or sink strength of these atmospheric substances

can be further magnied by the fact that the climate system is also sensitive to the

feedback from terrestrial ecosystems forming the so-called `feedback cycle'. Hence,

the emergent need is to reduce uncertainties when assessing this complex and dynamic

feedback cycle that is necessary to support the decisions of mitigation and

adaptation policies associated with human activities (e.g., anthropogenic emission

controls and land use managements) under current and future climate regimes.

With the goal to improve the predictions for the biosphere-atmosphere exchange

of biologically active gases and atmospheric aerosol particles, the main focus of this

dissertation is on revising and up-scaling the biotic and abiotic transport processes

from leaf to canopy scales. The validity of previous modeling studies in determining

iv

the exchange rate of gases and particles is evaluated with detailed descriptions of their

limitations. Mechanistic-based modeling approaches along with empirical studies

across dierent scales are employed to rene the mathematical descriptions of surface

conductance responsible for gas and particle exchanges as commonly adopted by all

operational models. Specically, how variation in horizontal leaf area density within

the vegetated medium, leaf size and leaf microroughness impact the aerodynamic attributes

and thereby the ultrane particle collection eciency at the leaf/branch scale

is explored using wind tunnel experiments with interpretations by a porous media

model and a scaling analysis. A multi-layered and size-resolved second-order closure

model combined with particle

uxes and concentration measurements within and

above a forest is used to explore the particle transport processes within the canopy

sub-layer and the partitioning of particle deposition onto canopy medium and forest

oor. For gases, a modeling framework accounting for the leaf-level boundary layer

eects on the stomatal pathway for gas exchange is proposed and combined with sap

ux measurements in a wind tunnel to assess how leaf-level transpiration varies with

increasing wind speed. How exogenous environmental conditions and endogenous

soil-root-stem-leaf hydraulic and eco-physiological properties impact the above- and

below-ground water dynamics in the soil-plant system and shape plant responses

to droughts is assessed by a porous media model that accommodates the transient

water

ow within the plant vascular system and is coupled with the aforementioned

leaf-level gas exchange model and soil-root interaction model. It should be noted

that tackling all aspects of potential issues causing uncertainties in forecasting the

feedback cycle between terrestrial ecosystem and the climate is unrealistic in a single

dissertation but further research questions and opportunities based on the foundation

derived from this dissertation are also brie

y discussed.