Differential Species Richness and Ecological Success of Epiphytes and Hemiepiphytes of Neotropical Araceae and Cyclanthaceae.

Numerous plant functional traits of ecophysiology and morphology associated with an epiphytic life history have promoted relatively high rates of evolutionary diversification and ecological success in tropical families such as the Orchidaeae, Polypodiaceae, Bromeliaceae, and Cactaceae. Epiphytic life histories are relatively uncommon in the Araceae and rare in the Cyclanthaceae which lack key functional traits for epiphytism. Only two lineages of Neotropical Araceae, Anthurium and Philodendron, include examples of epiphyte life histories. The evolution of a hemiepiphytic life history represented an important development for tropical Araceae by providing functional traits that have greatly expanded opportunities for adaptive radiation and ecological success as indicated by species richness and frequency of occurrence. The key adaptive trait allowing the diversification of hemiepiphytic Araceae was the development of heteroblastic growth of leaves and stems. Although hemiepiphytic life histories are present in the Cyclanthaceae, the family has undergone only modest speciation and limited ecological success in both its epiphytes and hemiepiphytes. Extensive sampling of more than 4600 trees from primary forest on four soil groups in northeastern Costa Rica have found a modest diversity of 15 species of epiphytic Araceae but only two species of epiphytic Cyclanthaceae. In contrast, 38 species of hemiepiphytic Araceae and 5 species of hemiepiphytic Cyclanthaceae were sampled, indicating relatively limited adaptive radiation of hemiepiphytic Cyclanthaceae and lower ecological success. Using summed values of frequency of occurrence as a measure of ecological success, epiphytic Araceae were 18 to 42 times more frequent than epiphytic Cyclanthaceae in swamp, alluvial, and residual soil forests. Summed frequencies of occurrence of hemiepiphytic Araceae were 7 to 13 times higher than those of hemiepiphytic Cyclanthaceae. The four soil groups were similar in their floristic composition of epiphytic and hemiepiphytic Araceae and Cyclanthaceae, but the frequencies of occurrence of both epiphytes and hemiepiphytes were, with few exceptions, highest on swamp soil plots, with alluvial soil plots slightly less favorable.

Collaborative Use of Sensor Networks and Cyberinfrastructure to Understand Complex Ecosystem Interactions in a Tropical Rainforest: Challenges and Lessons Learned.

Collaborations between ecosystem ecologists and engineers have led to impressive progress in developing complex models of biogeochemical fluxes in response to global climate change. Ecology and engineering iteratively inform and transform each other in these efforts. Nested data streams from local sources, adjacent networks, and remote sensing sources together magnify the capacity of ecosystem ecologists to observe systems in near real-time and address questions at temporal and spatial scales that were previously unobtainable. We describe our research experiences working in a Costa Rican rainforest ecosystem with the challenges presented by constant high humidity, 4300 mm of annual rainfall, flooding, small invertebrates entering the tiniest openings, stinging insects, and venomous snakes. Over the past two decades, we faced multiple challenges and learned from our mistakes to develop a broad program of ecosystem research at multiple levels of integration. This program involved integrated networks of diverse sensors on a series of canopy towers linked to multiple belowground soil sensor arrays that could transport sensor data streams from the forest directly to an off-site location via a fiber optic cable. In our commentary, we highlight three components of our work: (1) the eddy flux measurements using canopy towers; (2) the soil sensor arrays for measuring the spatial and temporal patterns of CO2 and O2 fluxes at the soil-atmosphere interface; and (3) focused investigations of the ecosystem impact of leaf-cutter ants as "ecosystem engineers" on carbon fluxes.

Leaf-cutter ants - mycorrhizal fungi: observations and research questions from an unexpected mutualism.

Leaf-cutter ants (LCAs) are widely distributed and alter the physical and biotic architecture above and below ground. In neotropical rainforests, they create aboveground and belowground disturbance gaps that facilitate oxygen and carbon dioxide exchange. Within the hyperdiverse neotropical rainforests, arbuscular mycorrhizal (AM) fungi occupy nearly all of the forest floor. Nearly every cubic centimeter of soil contains a network of hyphae of Glomeromycotina, fungi that form arbuscular mycorrhizae. Our broad question is as follows: how can alternative mycorrhizae, which are-especially ectomycorrhizae-essential for the survival of some plant species, become established? Specifically, is there an ant-mycorrhizal fungus interaction that facilitates their establishment in these hyperdiverse ecosystems? In one lowland Costa Rican rainforest, nests of the LCA Atta cephalotes cover approximately 1.2% of the land surface that is broadly scattered throughout the forest. On sequencing the DNA from soil organisms, we found the inocula of many AM fungi in their nests, but the nests also contained the inocula of ectomycorrhizal, orchid mycorrhizal, and ericoid mycorrhizal fungi, including Scleroderma sinnamariense, a fungus critical to Gnetum leyboldii, an obligate ectomycorrhizal plant. When the nests were abandoned, new root growth into the nest offered opportunities for new mycorrhizal associations to develop. Thus, the patches created by LCAs appear to be crucial sites for the establishment and survival of shifting mycorrhizal plant-fungal associations, in turn facilitating the high diversity of these communities. A better understanding of the interactions of organisms, including cross-kingdom and ant-mycorrhizal fungal interactions, would improve our understanding of how these ecosystems might tolerate environmental change.

Functional trait diversity of Cyclanthaceae and its convergent evolution with Araceae in Neotropical forests.

The Cyclanthaceae comprise a relatively small family of about 230 species and 12 genera in the Pandanales that is widespread in wet Neotropical forests. The great majority of species can be divided into three growth forms (understory herbs, epiphytes, and root-climbing hemiepiphytes) that share functional traits with similar growth forms present in the Araceae, a member of the Alismatales and not closely related. Our objectives were first to characterize the diversity, functional growth forms, and ecological traits of Cyclanthaceae at the La Selva Biological Station. Specific functional leaf and canopy traits of terrestrial herbs and epiphytes are very similar and associated with ecological success in both families. We further examined the functional traits of root-climbing hemiepiphytes, a specialized growth form that links the two families but rare in other families and argue that their specialized functional traits allow them to be considered as a distinct functional growth form. A key trait in distinguishing hemiepiphytes which are rare outside of the Cyclanthaceae and Araceae is the severance of the main stem hydraulic connection to the soil early in plant development. We used field data to examine the possible evolutionary pathways of developmental and ecological transition from terrestrial to hemiepiphyte growth forms. The broader ecological success of hemiepiphytic Araceae compared to Cyclanthaceae is hypothesized to result from the presence of heteroblasty in developing stems and leaves which allows more efficient utilization of complex canopy light environments of wet tropical forests.

Plasticity in Compensatory Growth to Artificial Defoliation and Light Availability in Four Neotropical Understory and Forest Edge Herb Species.

Defoliation from falling branches is a major factor in the survival of understory herbs in tropical rainforests. Experimental studies of defoliation under three levels of light environment compared responses to partial and total defoliation in four species of tropical rainforest understory herbs. We predicted that elevated levels of light availability would help compensate for damage to through compensatory growth in both understory and forest edge species and that forest edge species would more effectively compensate under high light conditions than shade-tolerant species from the forest understory All species showed a high tolerance to defoliation under high and intermediate light conditions. Under low-light conditions survival differed dramatically with minimal mortality in forest-edge species compared to high mortality in completely defoliated understory species. Defoliation, and light × defoliation interactions, impacted multiple growth traits in understory species. In contrast, forest-edge species showed no effect of defoliation except on total biomass, and only one light × defoliation interaction was observed. Our results indicate that differences in biomass allocation, leaf ecophysiology, and other growth parameters between forest understory and edge species may be structuring post-damage response in understory and forest edge herbs.

Functional traits of broad-leaved monocot herbs in the understory and forest edges of a Costa Rican rainforest.

Broad-leaved monocot herbs are widespread and dominant components of the shaded understories of wet neotropical forests. These understory habitats are characterized by light limitation and a constant threat of falling branches. Many shaded understory herb species have close relatives that occupy forest edges and gaps, where light availability is higher and defoliation threat is lower, creating an opportunity for comparative analysis of functional traits in order to better understand the evolutionary adaptations associated with this habitat transition. We documented ecological, morphological and ecophysiological traits of multiple herb species in six monocot families from each of these two habitats in the wet tropical rainforest at the La Selva Biological Station, Costa Rica. We found that a mixture of phylogenetic canalization and ecological selection for specific habitats helped explain patterns of functional traits. Understory herbs were significantly shorter and had smaller leaves than forest edge species. Although the mean number of leaves per plant and specific leaf area did not differ between the two groups, the larger leaf size of forest edge species gave them more than three times the mean plant leaf area. Measures of leaf water content and nitrogen content varied within both groups and mean values were not significantly different. Despite the high leaf nitrogen contents, the maximum photosynthetic rates of understory herbs were quite low. Measures of δ 13C as an analog of water use efficiency found significantly lower (more negative) values in understory herbs compared to forest edge species. Clonality was strongly developed in several species but did not show strong phylogenetic patterns. This study highlights many functional traits that differ between broad-leaved monocot species characteristic of understory and forest edge habitats, as well as traits that vary primarily by phylogenetic relatedness. Overall, plant functional traits do not provide a simple explanation for the relative differences in abundance for individual understory and forest edge species with some occurring in great abundance while others are relatively rare.

Invasive annuals respond more negatively to drought than native species.

In his foundational list of 'ideal weed' characteristics, Baker (1965) proposed that weedy plants maximize reproductive output under high resource availability. Since then, the idea that invasive plant species are more responsive to fluctuating resources compared with native or noninvasive species has gained considerable traction, although few studies extend this hypothesis to include reproductive output. We revisit Baker's hypothesis in the context of invasion and drought in California grasslands, exploring whether invasives show greater growth and reproductive responses to water availability compared with the native wildflowers they displace. In an outdoor potted study, we grew eight native and eight invasive species of annuals commonly found in southern California grasslands to reproductive maturity under both well-watered and drought conditions. While drought negatively impacted plant performance overall, invasives showed more negative responses for growth and reproductive traits. Invasives also grew larger than native species, especially under well-watered conditions, and produced seed with higher rates of germination. Invasives may be more negatively impacted by drought compared with natives, but they are also able to capitalize on high resource conditions and greatly increase reproductive output. Such opportunistic responses exhibited by invasives might explain previously observed fluctuations in their abundance under variable precipitation.

Fire and Plant Diversification in Mediterranean-Climate Regions.

Despite decades of broad interest in global patterns of biodiversity, little attention has been given to understanding the remarkable levels of plant diversity present in the world's five Mediterranean-type climate (MTC) regions, all of which are considered to be biodiversity hotspots. Comprising the Mediterranean Basin, California, central Chile, the Cape Region of South Africa, and southwestern Australia, these regions share the unusual climatic regime of mild wet winters and warm dry summers. Despite their small extent, covering only about 2.2% of world land area, these regions are home to approximately one-sixth of the world vascular plant flora. The onset of MTCs in the middle Miocene brought summer drought, a novel climatic condition, but also a regime of recurrent fire. Fire has been a significant agent of selection in assembling the modern floras of four of the five MTC regions, with central Chile an exception following the uplift of the Andes in the middle Miocene. Selection for persistence in a fire-prone environment as a key causal factor for species diversification in MTC regions has been under-appreciated or ignored. Mechanisms for fire-driven speciation are diverse and may include both directional (novel traits) and stabilizing selection (retained traits) for appropriate morphological and life-history traits. Both museum and nursery hypotheses have important relevance in explaining the extant species richness of the MTC floras, with fire as a strong stimulant for diversification in a manner distinct from other temperate floras. Spatial and temporal niche separation across topographic, climatic and edaphic gradients has occurred in all five regions. The Mediterranean Basin, California, and central Chile are seen as nurseries for strong but not spectacular rates of Neogene diversification, while the older landscapes of southwestern Australia and the Cape Region show significant components of both Paleogene and younger Neogene speciation in their diversity. Low rates of extinction suggesting a long association with fire more than high rates of speciation have been key to the extant levels of species richness.

Size class structure, growth rates, and orientation of the central Andean cushion Azorella compacta.

Azorella compacta (llareta; Apiaceae) forms dense, woody, cushions and characterizes the high elevation rocky slopes of the central Andean Altiplano. Field studies of an elevational gradient of A. compacta within Lauca National Park in northern Chile found a reverse J-shape distribution of size classes of individuals with abundant small plants at all elevations. A new elevational limit for A. compacta was established at 5,250 m. A series of cushions marked 14 years earlier showed either slight shrinkage or small degrees of growth up to 2.2 cm yr(-1). Despite their irregularity in growth, cushions of A. compacta show a strong orientation, centered on a north-facing aspect and angle of about 20° from horizontal. This exposure to maximize solar irradiance closely matches previous observations of a population favoring north-facing slopes at a similar angle. Populations of A. compacta appear to be stable, or even expanding, with young plants abundant.

Land use compounds habitat losses under projected climate change in a threatened California ecosystem.

Given the rapidly growing human population in mediterranean-climate systems, land use may pose a more immediate threat to biodiversity than climate change this century, yet few studies address the relative future impacts of both drivers. We assess spatial and temporal patterns of projected 21(st) century land use and climate change on California sage scrub (CSS), a plant association of considerable diversity and threatened status in the mediterranean-climate California Floristic Province. Using a species distribution modeling approach combined with spatially-explicit land use projections, we model habitat loss for 20 dominant shrub species under unlimited and no dispersal scenarios at two time intervals (early and late century) in two ecoregions in California (Central Coast and South Coast). Overall, projected climate change impacts were highly variable across CSS species and heavily dependent on dispersal assumptions. Projected anthropogenic land use drove greater relative habitat losses compared to projected climate change in many species. This pattern was only significant under assumptions of unlimited dispersal, however, where considerable climate-driven habitat gains offset some concurrent climate-driven habitat losses. Additionally, some of the habitat gained with projected climate change overlapped with projected land use. Most species showed potential northern habitat expansion and southern habitat contraction due to projected climate change, resulting in sharply contrasting patterns of impact between Central and South Coast Ecoregions. In the Central Coast, dispersal could play an important role moderating losses from both climate change and land use. In contrast, high geographic overlap in habitat losses driven by projected climate change and projected land use in the South Coast underscores the potential for compounding negative impacts of both drivers. Limiting habitat conversion may be a broadly beneficial strategy under climate change. We emphasize the importance of addressing both drivers in conservation and resource management planning.

Comparative patterns of plant invasions in the Mediterranean biome.

The objective of this work was to compare and contrast the patterns of alien plant invasions in the world's five mediterranean-climate regions (MCRs). We expected landscape age and disturbance history to have bearing on levels of invasion. We assembled a database on naturalized alien plant taxa occurring in natural and semi-natural terrestrial habitats of all five regions (specifically Spain, Italy, Greece and Cyprus from the Mediterranean Basin, California, central Chile, the Cape Region of South Africa and Southwestern - SW Australia). We used multivariate (hierarchical clustering and NMDS ordination) trait and habitat analysis to compare characteristics of regions, taxa and habitats across the mediterranean biome. Our database included 1627 naturalized species with an overall low taxonomic similarity among the five MCRs. Herbaceous perennials were the most frequent taxa, with SW Australia exhibiting both the highest numbers of naturalized species and the highest taxonomic similarity (homogenization) among habitats, and the Mediterranean Basin the lowest. Low stress and highly disturbed habitats had the highest frequency of invasion and homogenization in all regions, and high natural stress habitats the lowest, while taxonomic similarity was higher among different habitats in each region than among regions. Our analysis is the first to describe patterns of species characteristics and habitat vulnerability for a single biome. We have shown that a broad niche (i.e. more than one habitat) is typical of naturalized plant species, regardless of their geographical area of origin, leading to potential for high homogenization within each region. Habitats of the Mediterranean Basin are apparently the most resistant to plant invasion, possibly because their landscapes are generally of relatively recent origin, but with a more gradual exposure to human intervention over a longer period.

Phenotypic plasticity and differentiation in fitness-related traits in invasive populations of the Mediterranean forb Centaurea melitensis (Asteraceae).

PREMISE OF THE STUDY: Biological invasions threaten global biodiversity, resulting in severe ecological and economic costs. Phenotypic plasticity and differentiation in fitness-related traits after introduction can contribute to increased performance in invasive populations of plants. We determined whether postintroduction evolution in trait means or in their plasticity, or inherent species-wide phenotypic plasticity has promoted invasiveness in a European annual forb. • METHODS: In a common greenhouse, we compared several fitness-related traits and the phenotypic plasticity of those traits under four levels of nutrients among native and invasive populations of Centaurea melitensis. We tested 18 populations from three regions of similar mediterranean climate type: the native range (southern Spain) and two invaded ranges (California and central Chile). • KEY RESULTS: Centaurea melitensis possesses overall phenotypic plasticity, which is a trait that promotes invasiveness. Invasive populations were differentiated from native plants for several trait means and their levels of phenotypic plasticity in directions that enhance competitive ability and success. Invasive plants flowered earlier and grew faster in the early stages of growth phases, important features for invasiveness. • CONCLUSIONS: Phenotypic plasticity, its evolution postinvasion, and the evolution of fitness-related trait means in invasive populations have potentially contributed to the invasion of C. melitensis in California and Chile. Along with an overall wide range of tolerance to growing conditions, C. meltiensis populations that have colonized habitats in California and Chile have undergone rapid evolution in several life history traits and the plasticities of those traits in directions that would promote invasiveness in mediterranean ecosystems.

Fire as an evolutionary pressure shaping plant traits.

Traits, such as resprouting, serotiny and germination by heat and smoke, are adaptive in fire-prone environments. However, plants are not adapted to fire per se but to fire regimes. Species can be threatened when humans alter the regime, often by increasing or decreasing fire frequency. Fire-adaptive traits are potentially the result of different evolutionary pathways. Distinguishing between traits that are adaptations originating in response to fire or exaptations originating in response to other factors might not always be possible. However, fire has been a factor throughout the history of land-plant evolution and is not strictly a Neogene phenomenon. Mesozoic fossils show evidence of fire-adaptive traits and, in some lineages, these might have persisted to the present as fire adaptations.

Environmental sensor networks in ecological research.

Environmental sensor networks offer a powerful combination of distributed sensing capacity, real-time data visualization and analysis, and integration with adjacent networks and remote sensing data streams. These advances have become a reality as a combined result of the continuing miniaturization of electronics, the availability of large data storage and computational capacity, and the pervasive connectivity of the Internet. Environmental sensor networks have been established and large new networks are planned for monitoring multiple habitats at many different scales. Projects range in spatial scale from continental systems designed to measure global change and environmental stability to those involved with the monitoring of only a few meters of forest edge in fragmented landscapes. Temporal measurements have ranged from the evaluation of sunfleck dynamics at scales of seconds, to daily CO2 fluxes, to decadal shifts in temperatures. Above-ground sensor systems are partnered with subsurface soil measurement networks for physical and biological activity, together with aquatic and riparian sensor networks to measure groundwater fluxes and nutrient dynamics. More recently, complex sensors, such as networked digital cameras and microphones, as well as newly emerging sensors, are being integrated into sensor networks for hierarchical methods of sensing that promise a further understanding of our ecological systems by revealing previously unobservable phenomena.

The influence of life form on carbon and nitrogen relationships in tropical rainforest ferns.

Tropical ferns are characterized by a high diversity of plant life forms, yet there have been few large-scale studies on the functional ecology of these different forms. We examined epiphytic, hemiepiphytic, and terrestrial ferns, and asked whether there are differences in the mineral nutrition and water relations across different growth forms of a diverse assemblage of species. We measured specific leaf area, leaf nitrogen concentrations, and natural abundance of the stable isotopes delta(15)N and delta(13)C of 48 fern species from 36 genera across a wide range of habitats at La Selva Biological Station in Costa Rica. We found that epiphytes were significantly different in all measured variables from hemiepiphytic and terrestrial species, and that terrestrial and soil-rooted hemiepiphytes were indistinguishable in all variables excluding SLW. A multivariate analysis revealed that aspects of N nutrition were the most reliable at separating epiphytic species from other life forms. Our study demonstrates that the natural abundance of both C and N as well as N relations and leaf morphology are useful when segregating different plant life forms, and that the N cycle of epiphytic and terrestrial habitats function independently from each other.

Partitioning of soil water among tree species in a Brazilian Cerrado ecosystem.

Source water used by woody perennials in a Brazilian savanna (Cerrado) was determined by comparing the stable hydrogen isotope composition (deltaD) of xylem sap and soil water at different depths during two consecutive dry seasons (1995 and 1996). Plant water status and rates of water use were also determined and compared with xylem water deltaD values. Overall, soil water deltaD decreased with increasing depth in the soil profile. Mean deltaD values were -35 per thousand for the upper 170 cm of soil and -55 per thousand between 230 and 400 cm depth at the end of the 1995 dry season. Soil water content increased with depth, from 18% near the surface to about 28% at 400 cm. A similar pattern of decreasing soil water deltaD with increasing depth was observed at the end of the 1996 dry season. Patterns consistent with hydraulic lift were observed in soil profiles sampled in 1995 and 1997. Concurrent analyses of xylem and soil water deltaD values indicated a distinct partitioning of water resources among 10 representative woody species (five deciduous and five evergreen). Among these species, four evergreen and one deciduous species acquired water primarily in the upper soil layers (above 200 cm), whereas three deciduous and one evergreen species tapped deep sources of soil water (below 200 cm). One deciduous species exhibited intermediate behavior. Total daily sap flow was negatively correlated with xylem sap deltaD values indicating that species with higher rates of water use during the dry season tended to rely on deeper soil water sources. Among evergreen species, minimum leaf water potentials were also negatively correlated with xylem water deltaD values, suggesting that access to more readily available water at greater depth permitted maintenance of a more favorable plant water status. No significant relationship between xylem water deltaD and plant size was observed in two evergreen species, suggesting a strong selective pressure for small plants to rapidly develop a deep root system. The degree of variation in soil water partitioning, leaf phenology and leaf longevity was consistent with the high diversity of woody species in the Cerrado.

Conopy architecture of Larrea tridentata (DC.) Cov., a desert shrub: foliage orientation and direct beam radiation interception.

At sites in the United States, creosote bushes (Larrea tridentata (DC.) Cov.) orient foliage clusters predominantly toward the southeast. Foliage of bushes at the southernmost distribution extreme in Mexico shows no predominant orientation. Clusters at all sites are inclined between 33° and 71° from the horizontal. Inclinations are steeper in the drier and hotter Mojave Desert than in the Chihuahuan Desert. Individual leaflets, though not measured, appear more randomly oriented than foliage clusters. In several populations studied, branches were shorter in the southeastern sectors of the crown, reducing self-shading early in the morning. Measurements of direct beam radiation interception by detached branches, using digital image processing, indicated that foliage clusters oriented toward the southeast exhibited less self-shading during spring mornings than clusters oriented northeast. This effect was not apparent at the summer solstice. This type of canopy architecture may tend to minimize self-shading during the morning hours when conditions are more favorable for photosynthesis, resulting in an improved daily water use efficiency.

Nitrogen utilization efficiencies in mediterranean-climate shrubs of California and Chile.

Comparative studies of nitrogen utilization efficiencies (g assimilation mg-1 leaf nitrogen expended) for dominant chaparral shrubs of California and matorral shrubs of central Chile indicate varying relationships of these efficiencies to total shrub productivity and community dominance. In California where nitrogen is limiting for shrub growth, increasing nitrogen utilization efficiency is correlated with increased productivity, and species with the highest values are dominant on polar (north-facing) slopes. In Chile where soil nutrients are not strongly limiting, increased nitrogen utilization efficiency beyond a low threshold has little or no effect on productivity. This may be an important aspect of the relative ecological success of deciduous shrubs in Chile. High nitrogen utilization efficiency in Chile is associated with broad ecological amplitude rather than increased competitive ability through higher productivity.

Parallel evolution of leaf pubescence in Encelia in coastal deserts of North and South America.

Members of the shrub genus Encelia occur in the arid coastal regions of western North America and South America along gradients of precipitation extending from less than 50 mm annually to somewhat over 350 mm. At moist ends of the gradient species possess glabrate, green leaves. Proceeding to drier regions, the species possess progressively more pubescent, whiter leaves. Leaf absorptance to solar radiation decreases with precipitation in an identical pattern along gradients in both North and South America.