Zdravko Baruch

The worldwide leaf economics spectrum

Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.

Leaf construction cost, nutrient concentration, and net CO2 assimilation of native and invasive species in Hawaii.

Abstract The effects of biological invasions are most evident in isolated oceanic islands such as the Hawaiian Archipelago, where invasive plant species are rapidly changing the composition and function of plant communities. In this study, we compared the specific leaf area (SLA), leaf tissue construction cost (CC), leaf nutrient concentration, and net CO2 assimilation (A) of 83 populations of 34 native and 30 invasive species spanning elevation and substrate age gradients on Mauna Loa volcano in the island of Hawaii. In this complex environmental matrix, where annual precipitation is higher than 1500 mm, we predicted that invasive species, as a group, will have leaf traits, such as higher SLA and A and lower leaf CC, which may result in more efficient capture of limiting resources (use more resources at a lower carbon cost) than native species. Overall, invasive species had higher SLA and A, and lower CC than native species, consistent with our prediction. SLA and foliar N and P were 22.5%, 30.5%, and 37.5% higher, respectively, in invasive species compared to native ones. Light-saturated photosynthesis was higher for invasive species (9.59 μmol m−2 s−1) than for native species (7.31 μmol m−2 s−1), and the difference was larger when A was expressed on a mass basis. Leaf construction costs, on the other hand, were lower for the invasive species (1.33 equivalents of glucose g−1) than for native species (1.37). This difference was larger when CC was expressed on an area basis. The trends in the above traits were maintained when groups of ecologically equivalent native and invasive species (i.e., sharing similar life history traits and growing in the same habitat) were compared. Foliar N and P were significantly higher in invasive species across all growth forms. Higher N may partially explain the higher A of invasive species. Despite relatively high N, the photosynthetic nitrogen use efficiency of invasive species was 15% higher than that of native species. These results suggest that invasive species may not only use resources more efficiently than native species, but may potentially demonstrate higher growth rates, consistent with their rapid spread in isolated oceanic islands.

African Grass Invasion in the Americas: Ecosystem Consequences and the Role of Ecophysiology

Clearing of natural vegetation for pastures and the deliberate introduction of African grasses constitute significant threats to the biological diversity of the tropics, subtropics, and warm temperate regions of the Americas. African grasses have escaped from cultivated pastures and revegetated rangeland sites and invaded natural areas at alarming rates. Invaded ecosystems tend to be biotically impoverished and differ markedly from adjacent non-invaded areas in structure and function. Effects of pasture creation and invasion by African grasses on ecosystem processes (transformation and flux of energy and matter) are primarily related to loss of woody species and changes in the fire regime. However, the ecophysiological attributes of the African grasses (e.g. high biomass allocation to leaves, high growth rate, and high leaf-level gas exchange rates) also have important consequences. Here we describe the extent of pasture creation with African grasses and their invasive spread in the New World and review ecological effects of these land-cover changes. We highlight a number of comparative ecophysiological studies within the context of mechanisms responsible for invasion by African grasses and resulting ecosystem change.

Responses to drought of five Brachiaria species. I. Biomass production, leaf growth, root distribution, water use and forage quality

The introduction of African grasses in Neotropical savannas has been a key factor to improve pasture productivity. We compared the response of five Brachiaria species to controlled drought (DT) in terms of biomass yield and allocation, pattern of root distribution, plant water use, leaf growth, nutrient concentration and dry matter digestibility. The perennial C4 forage grasses studied were B. brizantha (CIAT 6780), B. decumbens (CIAT 606), B. dictyoneura (CIAT 6133), B. humidicola (CIAT 679) and B. mutica. Two DT periods, which mimic short dry spells frequent in the rainy season, were imposed by suspending irrigation until wilting symptoms appeared. They appeared after 14 days in B. brizantha, B. decumbens and B. mutica, and after 28 days in B. humidicola and B. dictyoneura. The impossed drought stress was mild and only the largest grass, B. brizantha, showed reduced (23%) plant yield. The other grasses were able to adjust growth and biomass allocation in response to DT leaving total plant yield relatively unaffected. Brachiaria mutica, had a homogeneous root distribution throughout the soil profile. In the other species more than 80% of root biomass was allocated within the first 30 cm of the soil profile. Brachiaria brizantha and B. decumbens had the lowest proportion of roots below 50 cm. Drought caused a general reduction in root biomass. The shoot:root ratio in B. mutica and B. humidicola increased in response to DT at the expense of a reduction in root yield down to 50 cm depth. Although the total water volume utilized under DT was similar among grasses, the rate of water use was highest (0.25 l day−1) in B. brizantha, B. decumbens and B. mutica and lowest (0.13 l day−1) in B. humidicola and B. dictyoneura. In all species leaf expansion was reduced by DT but it was rapidly reassumed after rewatering. Drought increased specific leaf mass (SLM) only in B. brizantha compensating for leaf area reduction, but leaf area ratio (LAR) was unaffected in all species. In almost all grasses DT increased leaf N and K concentration and in vitro dry matter digestibility. The results indicate that B. brizantha, B. decumbens and to a lesser extent, B. mutica are better adapted to short dry periods, whereas B. humidicola and B. dictyoneura are better adapted to longer dry periods.

Responses to simulated herbivory and water stress in two tropical C4 grasses

SummaryThe African grass Hyparrhenia rufa has established itself successfully in South American savannas (Llanos) and displaced dominant native grasses such as Trachypogon plumosus from the wetter and more fertile habitats. Several ecophysiological traits have been related to the higher competitive capacity of H. rufa. To further analyze the behavior of both species, their growth, biomass allocation, physiological and architectural responses to defoliation and water stress were compared under controlled conditions. Although total, aerial and underground biomass decreased under defoliation in both grasses, increases in clipped-leaf biomass and area compensated for defoliation in H. rufa but not in T. plumosus. This difference was due mainly to a higher proportion of assimilates being directed to leaf and tiller production and a higher leaf growth rate in the African grass as compared to T. plumosus, which showed incrased senescence under frequent defoliation. In both species, water stress ameliorated the effects of defoliation. The ability to compensate for defoliated biomass in H. rufa is possibly related to its long coevolution with large herbivores in its original African habitat and is apparently one of the causes of its success in Neotropical savannas.

Responses to drought and flooding in tropical forage grasses

Seasonal drought and flooding severely limit pasture growth in tropical savannas. The objective of this study is to analyze and compare yield, biomass allocation, leaf growth rate and nutrient concentration of four important perennial C4 forage grasses to short term flooding and moderate drought under controlled conditions. The grasses studied were the tufted Andropogon gayanus (CIAT 621) and Hyparrhenia rufa and the stoloniferous Echinochloa polystachya and Brachiaria mutica.All grasses were able to adjust their growth and development in response to flooding and drought: leaf growth and total biomass decreased under both treatments but the specific responses to these treatments differed markedly. Considering only total yield and leaf area, A. gayanus and H. rufa were relatively more tolerant to and less affected by drought whereas B. mutica and E. polystachya were more flood tolerant.In A. gayanus and H. rufa, both treatments reduced the proportion of assimilates devoted to roots and culms while increasing that of leaves decreasing the root/shoot ratio. In contrast, in B. mutica and E. polystachya only the proportion devoted to culms or stolons increased under flooding but the root/shoot ratio remained relatively stable under both treatments. All grasses produced adventitious rootlets except A. gayanus which was the most affected by flooding. Waterlogging decreased leaf nutrient concentration in all grasses which contributed to growth reduction. All species were relatively tolerant to both stresses. The results confirm the empirical observation that stoloniferous species B. mutica and E. polystachya are more tolerant to flooding thanks to adaptations typical of wetland plants such as hollow stolons which enhance oxygen diffusion to the roots and the development of adventitious rootlets that promotes water and nutrient absorption.

Water relations of native and introduced C4 grasses in a neotropical savanna

Introduced African grasses are invading Neotropical savannas and displacing the native herbaceous community. This work, which is part of a program to understand the success of the African grasses, specifically investigates whether introduced and native grasses differ in their water relations. The water relations of the native Trachypogon plumosus and the successful invader Hyparrhenia rufa were studied in the field during two consecutive years in the seasonal savannas of Venezuela. The two C4 grasses differed clearly in their responses to water stress. H. rufa consistently had higher stomatal conductance, transpiration rate, leaf water and osmotic potential and osmotic adjustment than the native T. plumosus. Also, leaf senescence occurred much earlier during the dry season in H. rufa. Both grasses showed a combination of water stress evasion and tolerance mechanisms such as stomatal sensitivity to atmospheric or soil water stress, decreased transpiring area and osmotic adjustment. Evasion mechanisms are more conspicuous in H. rufa whereas T. plumosus is more drought tolerant and uses water more “conservatively”. The evasion mechanisms and oportunistic use of water by H. rufa, characteristic of invading species, contribute to, but only partially explain, the success of this grass in the Neotropical savannas where it displaces native plants from sites with better water and nutrient status. Conversely, the higher water stress tolerance of t. plumosus is consistent with its capacity to resist invasion by alien grasses on shallow soils and sites with poorer nutrient and water status.

Effects of fire and defoliation on the life history of native and invader C4 grasses in a Neotropical savanna

Abstract African grasses, introduced into Neotropical savannas to improve forage quality, have spread successfully and displaced native plants. To understand their competitive relationships, we compared biomass production and allocation, plant architecture and phenology, net photosynthesis (Pn), water relations, and nutrient content under fire and simulated herbivory between two C4 grasses, the native Trachypogon plumosus and the introduced Hyparrhenia rufa from a seasonal savanna in Venezuela. All variables were strongly influenced by the rainfall regime. Hyparrhenia produced bigger plants (in mass and size) with a large proportion of mass (>75%) allocated to leaves and culms. Its biomass production was more affected by fire than by defoliation. In contrast, Trachypogon was more affected by defoliation than by fire which promoted a flush of leaf growth even in the dry season. Fire caused up to 85% mortality in Hyparrhenia but none in Trachypogon where it increased inflorescence production. However, fire promoted abundant seed germination and fast seedling growth in Hyparrhenia, enabling it to colonize new areas. During the growing season Trachypogon had higher Pn and lower leaf water potential (Ψ) than Hyparrhenia but differences among treatments were not significant for either grass. Pn of Trachypogon ceased at a lower Ψ (−3.0 MPa) than in Hyparrhenia (−2.0 MPa), indicating its higher tolerance to water stress. During the dry season, Trachypogon leaves remained alive and retained low Pn. Leaf nutrient content was higher during the rainy season in both species. Differences in Pn could not explain the higher seasonal biomass production of Hyparrhenia. However, its water stress evasion strategy, larger biomass allocated to leaves, abundant germination and fast seedling growth appeared to be responsible for the success of Hyparrhenia as an invader of Neotropical savannas.

Ordination and classification of vegetation along an altitudinal gradient in the Venezuelan páramos

By means of ordination and classification techniques, the relationships between climate, soils, human activities and vegetation along an altitudinal gradient of the Venezuelan páramos are analyzed and interpreted. The altitudinal gradient chosen is characterized by decrease of temperature, precipitation, soil fertility, soil water-holding capacity, and plant cover as altitude increases. The ordination results suggest vegetation changes to be primarily related to environmental changes occurring with altitude, and secondly to disturbances caused mainly by grazing. Some results point toward a disjunction in the vegetational gradient occurring at ca. 3 500 m.a.s.l. and separating low and high páramo. This disjunction might have been caused by the glacial history of the páramos and the occurrence of frequent night-frosts. The soil samples were kindly analyzed by the Laboratorio de Edafologia, Centro Nacional de Investigaciones Agropecuarias. Help in plant identification was generously obtained from the specialists of Instituto Botánico, Instituto Nacional de Parques, Caracas.

Dynamics of energy and nutrient concentration and construction cost in a native and two alien C4 grasses from two neotropical savannas

In Venezuela, the alien grasses Melinis minutiflora Beauv. and Hyparrhenia rufa (Nees.) Stapf tend to displace the native savanna plant community dominated by Trachypogon plumosus (Humb. and Bonpl.) Nees. This occurs in either relatively wetter and fertile highland savannas or in drier and less fertile lowland savannas. Although the native and aliens are perennial C4 grasses, higher net assimilation leaf biomass per plant and germination rate of the latter are some causes for their higher growth rates and for their competitive success. The objective of this study is to compare seasonal tissue energy, N, P and K concentrations and the calculated construction costs (CC) between the native grass and either one of the alien grasses from lowland and highland savannas. We predict that, in order to out-compete native plants, alien grasses should be more efficient in resource use as evidenced by lower tissue energy and nutrient concentrations and CC.Tissue energy and nutrient concentration were measured throughout the year and compared between M. minutiflora and the co-occurring local population of T. plumosus in a highland savanna and between H. rufa and its neighbor local population of T. plumosus in a lowland savanna. CC was calculated from energy, N and ash concentrations considering ammonium as the sole N source. Differences between co-occurring species, T. plumosus populations, seasons, and organs were analyzed with ANOVA.Highland and lowland grasses differed in concentration and allocation of energy and nutrients whereas the differences between alien and native grasses were specific for each pair considered. Highland grasses had higher energy, N, P and CC than lowland grasses. These variables were always lowest in the culms. In the more stressed lowland site, tissue energy and nutrient concentrations decreased significantly during the dry season except in the roots of both grasses which had the highest energy and nutrients concentrations during the drought. This seasonal response was more marked in the local lowland population of T. plumosus in which maximum CC alternated seasonally between leaves and roots. Energy and nutrient concentrations and CC were the lowest in H. rufa. In the lowland savannas, the higher efficiency of resource use in the invader grass contributes to its higher competitive success through increased growth rate. In the highlands, overall tissue energy concentration and CC, but not N nor P concentration, were lower in the fast growing M. minutiflora but seasonal differences were lacking. The higher leaf CC in T. plumosus can be attributed to the higher proportion of sclerenchyma tissue which is more expensive to construct. Considering CC, both fast growing alien grasses are more efficient in resource use than the co-occurring native grass. However, the role of CC explaining the competitive success of the former, through higher growth rates, is more evident in the more stressful environment of the lowland savanna.