Acacia trees with parasitic ants have fewer and less spacious spines than trees with mutualistic ants.

Obligate ant-defended plants provide food and shelter in exchange for protection against herbivores. Mesoamerican acacia trees have an obligate ant mutualism, but parasitic non-defending ants can also nest on the tree. We assessed whether rewards corresponded to ant defense within a plant species. As we expected, we found that parasite-inhabited trees had fewer swollen spines than ant-defended trees. Spine diameter was smaller in parasite-inhabited plants, but there were no differences in spine length, suggesting that spines serve as mechanical protection against herbivory. Parasite-inhabited plants may have reduced rewards because of plant differences when establishing, a plastic response to limited resources, or differential energy allocation when sensing the lack of defense.

Pollen micromorphological analysis of tribe Acacieae (Mimosaceae) with LM and SEM techniques.

The morphology of pollen grains of the 06 species of tribe Acacieae (Mimosaceae) belonging to one genus were investigated using scanning electron microscopic (SEM) and light microscopic (LM) approach. Lactic acid acetolysis method was used for (LM) while non-acetolysis for (SEM). The species were examined for the both qualitative and quantitative palynological features. Qualitatively the pollen was studied for the shape, exine sculpturing, pollen shape, and pollen type. Oblate spheroidal shape of pollen was observed in Accacia modesta. Subprolate shape in Accaca nilotica. In Accacia farnesiana, prolate pollen was examined while prolate spheroidal in Acacia tortilis and spheroidal pollen grains were studied in Acacia catechu, respectively. Pollen exine sculpturing of the studied tribe possesses novel variation. Foveolate, reticulate, foveolate-psilate, faint reticulate, fossulate, perforate, and scrabate exine sculpturing were noticed. Colpi were absent in all species. Excessive variation was observed in both qualitative and quantitative character of pollen. SPSS software was use for the quantitative measurement of the pollen grains. The current study will be helpful for the identification of the problematic species and phylogenetic studies of family Mimosaceae.

Changes in cambial activity are related to precipitation patterns in four tropical hardwood species grown in Indonesia.

PREMISE: Cambial activity in some tropical trees varies intra-annually, with the formation of xylem rings. Identification of the climatic factors that regulate cambial activity is important for understanding the growth of such species. We analyzed the relationship between climatic factors and cambial activity in four tropical hardwoods, Acacia mangium, Tectona grandis, Eucalyptus urophylla, and Neolamarckia cadamba in Yogyakarta, Java Island, Indonesia, which has a rainy season (November-June) and a dry season (July-October). METHODS: Small blocks containing phloem, cambium, and xylem were collected from main stems in January 2014, October 2015 and October 2016, and examined with light microscopy for cambial cell division, fusiform cambial cells, and expanding xylem cells as evidence of cambial activity. RESULTS: During the rainy season, when precipitation was high, cambium was active. By contrast, during the dry season in 2015, when there was no precipitation, cambium was dormant. However, in October 2016, during the so-called dry season, cambium was active, cell division was conspicuous, and a new xylem ring formation was initiated. The difference in cambial activity appeared to be related to an unusual pattern of precipitation during the typically dry months, from July to October, in 2016. CONCLUSIONS: Our results indicate that low or absent precipitation for 3 to 4 months induces cessation of cambial activity and temporal periodicity of wood formation in the four species studied. By contrast, in the event of continuing precipitation, cambial activity in the same trees may continue throughout the year. The frequency pattern of precipitation appears to be an important determinant of wood formation in tropical trees.

A standardization method to disentangle environmental information from axial trends of xylem anatomical traits.

Anatomical traits such as xylem conduit diameter and vessel connectivity are fundamental characteristics of the hydraulic architecture of vascular plants. Stem xylem conduits are narrow at the stem apex, and this confers resistance to embolisms that might otherwise be induced by large, negative water potentials at the top of tall trees. Below the apex, conduits progressively widen and this characteristic minimizes effects of path length on total hydraulic resistance. While interconnections among xylem vessels have been noted for decades, their role(s) are not fully clarified. For example, we do not know if they allow water to bypass embolized vessels, or increase the risk of spread of embolisms, or how their arrangement varies within a tree. Here we demonstrate the benefit of removing the independent effect of stem length on assessment of effects of external (e.g., climatic) factors on such xylem traits. We measured the hydraulic diameter (Dh) and vessel conductivity index (VCI) along the stem of 21 shrubs/trees of similar height (1.19 < H < 5.45 m) belonging to seven Acacia species, across a wide aridity gradient in Australia. All trees showed similar scaling exponents of Dh (b = 0.33) and VCI (b = 0.53) vs axial distance from the apex (L), thus conforming with general patterns in woody plants. After de-trending for L, neither Dh (P = 0.21) nor VCI (P = 0.109) differed across the aridity gradient. We found that across a wide gradient of aridity, climate had no effect on xylem anatomy of Acacia spp, which was instead dictated by axial distances from stem apices. We argue that the use of standardization procedures to filter out intrinsic patterns of vascular traits is an essential step in assessing climate-driven modifications of xylem architecture.

Intraspecific variation in spermoderm pattern of tribe Acacieae (Mimosoideae) using scanning electron microscopy techniques.

Current research carried out in Pakistan is the first report on spermoderm ornamentation of eight species of tribe Acacieae (Mimosoidae) by using scanning electron microscopic techniques representing two genera, Fedherbia and Acacia were examined. Different spermoderm ornamentation were observed, described and discussed for their taxonomic importance. Seeds surfaces of the studied tribe possess novel variations in macro and micro morphology. Great variations were observed in both qualitative and quantitative characters of seeds. Seeds shape was oblong, ovate to elliptical and spermoderm ornamentation was levigate, rugose, polygonal and discoid, colliculate, and papillose type. These variations in the spermoderm ornamentation can be used as an aid in identification and classification of the members of tribe Acacieae.

Critical analyses when modeling tree biomass to ensure additivity of its components.

It is presented the theme additivity of biomass of tree components. To evaluate and discuss this context, experimental information collected in forests of Acacia mearnsii De Wild. was used. Equations for components (stem and crown) and total biomass were fitted by means of two procedures: 1) generalized nonlinear least squares and 2) weighted-nonlinear seemingly unrelated regressions. Analyzing the performance of the estimators, it can be concluded that the two tested procedures are equivalent. On the other hand, this conclusion differs when evaluated the consistency and efficiency of the estimators. Fitting equations for the components and for the total biomass by an independent way is not realistic, because from a biological point of view the estimates of biomass are inconsistent, i.e., are not additive. The biomass estimates of the components and of the total, resulting from equations adjusted by means of systems of equations, provided narrower confidence intervals in relation to the equations adjusted independently, and is therefore more efficient. The second procedure presents better biological properties and statistics to estimate allometric equations for biomass of the components and for the total when compared with the independent estimation, thus it should be the method to be used.

Stem gravitropism and tension wood formation in Acacia mangium seedlings inclined at various angles.

Background and Aims: In response to a gravitational stimulus, angiosperm trees generally form tension wood on the upper sides of leaning stems in order to reorientate the stems in the vertical direction. It is unclear whether the angle of inclination from the vertical affects tension wood formation. This study was designed to investigate negative gravitropism, tension wood formation and growth eccentricity in Acacia mangium seedlings inclined at different angles. Methods: Uniform seedlings of A. mangium were artificially inclined at 30°, 45°, 60° and 90° from the vertical and harvested, with non-inclined controls, 3 months later. We analysed the effects of the angle of inclination on the stem recovery angle, the anatomical features of tension wood and radial growth. Key Results: Smaller inclination angles were associated with earlier stem recovery while stems subjected to greater inclination returned to the vertical direction after a longer delay. However, in terms of the speed of negative gravitopism towards the vertical, stems subjected to greater inclination moved more rapidly toward the vertical. There was no significant difference in terms of growth eccentricity among seedlings inclined at different angles. The 30°-inclined seedlings formed the narrowest region of tension wood but there were no significant differences among seedlings inclined at 45°, 60° and 90°. The 90°-inclined seedlings formed thicker gelatinous layers than those in 30°-, 45°- and 60°-inclined seedlings. Conclusion: Our results suggest that the angle of inclination of the stem influences negative gravitropism, the width of the tension wood region and the thickness of gelatinous layers. Larger amounts of gelatinous fibres and thicker gelatinous layers might generate the higher tensile stress required for the higher speed of stem-recovery movement in A. mangium seedlings.

Climate trends in the wood anatomy of Acacia sensu stricto (Leguminosae: Mimosoideae).

Background and Aims: This study investigates the structural diversity of the secondary xylem of 54 species of Acacia from four taxonomic sections collected across five climate regions along a 1200 km E-W transect from sub-tropical [approx. 1400 mm mean annual precipitation (MAP)] to arid (approx. 240 mm MAP) in New South Wales, Australia. Acacia sensu stricto ( s.s. ) is a critical group for understanding the effect of climate and phylogeny on the functional anatomy of wood. Methods: Wood samples were sectioned in transverse, tangential and radial planes for light microscopy and analysis. Key Results: The wood usually has thick-walled vessels and fibres, paratracheal parenchyma and uniseriate and biseriate rays, occasionally up to four cells wide. The greater abundance of gelatinous fibres in arid and semi-arid species may have ecological significance. Prismatic crystals in chambered fibres and axial parenchyma increased in abundance in semi-arid and arid species. Whereas vessel diameter showed only a small decrease from the sub-tropical to the arid region, there was a significant 2-fold increase in vessel frequency and a consequent 3-fold decrease in the vulnerability index. Conclusions: Although the underlying phylogeny determines the qualitative wood structure, climate has a significant influence on the functional wood anatomy of Acacia s.s. , which is an ideal genus to study the effect of these factors.

Ground level photosynthetically active radiation dynamics in stands of Acacia mearnsii De Wild.

The objective is to study the dynamics of photosynthetic radiation reaching the soil surface in stands of Acacia mearnsii De Wild and its influence on height growth in stands. This fact gives rise to the formulation of the following hypothesis for this study: "The reduction of the incidence of light inside the stand of black wattle will cause the inflection point in its height growth when this reaches 4 to 5 m in height, i.e. when the stand is between 2 and 3 years of age". The study was conducted in stands in the state of Rio Grande do Sul, Brazil, where diameters at breast height, total height and photosynthetically active radiation available at ground level were measured. The frequency tended to be more intense when the age of the stands increases. It was evident that a reduction of light incidence inside the forest occurred, caused by canopy closure. Consequently, closed canopy propitiated the competition of plants. This has affected the conditions for growth in diameter and height of this species, reason why it becomes possible to conceive the occurrence of an inflection point in the growth of these two variables, confirming the formulated hypothesis.

A model based on environmental factors for diameter distribution in black wattle in Brazil.

This article discusses the dynamics of a diameter distribution in stands of black wattle throughout its growth cycle using the Weibull probability density function. Moreover, the parameters of this distribution were related to environmental variables from meteorological data and surface soil horizon with the aim of finding a model for diameter distribution which their coefficients were related to the environmental variables. We found that the diameter distribution of the stand changes only slightly over time and that the estimators of the Weibull function are correlated with various environmental variables, with accumulated rainfall foremost among them. Thus, a model was obtained in which the estimators of the Weibull function are dependent on rainfall. Such a function can have important applications, such as in simulating growth potential in regions where historical growth data is lacking, as well as the behavior of the stand under different environmental conditions. The model can also be used to project growth in diameter, based on the rainfall affecting the forest over a certain time period.

Grasses and browsers reinforce landscape heterogeneity by excluding trees from ecosystem hotspots.

Spatial heterogeneity in woody cover affects biodiversity and ecosystem function, and may be particularly influential in savanna ecosystems. Browsing and interactions with herbaceous plants can create and maintain heterogeneity in woody cover, but the relative importance of these drivers remains unclear, especially when considered across multiple edaphic contexts. In African savannas, abandoned temporary livestock corrals (bomas) develop into long-term, nutrient-rich ecosystem hotspots with unique vegetation. In central Kenya, abandoned corral sites persist for decades as treeless 'glades' in a wooded matrix. Though glades are treeless, areas between adjacent glades have higher tree densities than the background savanna or areas near isolated glades. The mechanisms maintaining these distinctive woody cover patterns remain unclear. We asked whether browsing or interactions with herbaceous plants help to maintain landscape heterogeneity by differentially impacting young trees in different locations. We planted the mono-dominant tree species (Acacia drepanolobium) in four locations: inside glades, far from glades, at edges of isolated glades and at edges between adjacent glades. Within each location, we assessed the separate and combined effects of herbivore exclusion (caging) and herbaceous plant removal (clearing) on tree survival and growth. Both caging and clearing improved tree survival and growth inside glades. When herbaceous plants were removed, trees inside glades grew more than trees in other locations, suggesting that glade soils were favorable for tree growth. Different types of glade edges (isolated vs. non-isolated) did not have significantly different impacts on tree performance. This represents one of the first field-based experiments testing the separate and interactive effects of browsing, grass competition and edaphic context on savanna tree performance. Our findings suggest that, by excluding trees from otherwise favorable sites, both herbaceous plants and herbivores help to maintain functionally important landscape heterogeneity in African savannas.

Hydraulic conductance of Acacia phyllodes (foliage) is driven by primary nerve (vein) conductance and density.

We determined effects of venation traits on hydraulic conductance of phyllodes (foliage), using an array of Acacia s.str. species with diverse phyllode morphologies as the source of variation. Measurements were made on phyllodes from 44 species, grown in common gardens but originating from different positions along a precipitation gradient. K(phyllode) varied 18-fold and was positively correlated with primary nerve hydraulic conductance, and with primary nerve (vein) density but not with minor nerve density, in contrast with previous studies of true leaves in other dicotyledons. Phyllodes with higher primary nerve density also had greater mass per area (PMA) and larger bundle sheath extensions (BSEs) from their minor nerves. We suggest that higher primary nerve conductivity and density may decrease the distance travelled in the high-resistance extra-xylem pathways of the phyllode. Further, larger BSEs may increase the area available for dispersion of water from the xylem to the extra-xylem tissue. High PMA phyllodes were more common in acacias from areas receiving lower annual precipitation. Maximizing efficient water movement through phyllodes may be more important where rainfall is meagre and infrequent, explaining relationships between nerve patterns and the climates of origin in Australian phyllodinous Acacia.

Three-dimensional xylem networks and phyllode properties of co-occurring Acacia.

Reduced leaf size is often correlated to increased aridity, where smaller leaves demand less water via xylem conduits. However, it is unknown if differences in three-dimensional (3D) xylem connectivity reflect leaf-level adaptations. We used X-ray microtomography (micro-CT) to quantify 3D xylem connectivity in ∼5 mm diameter branch sections of co-occurring semi-arid Acacia species of varied phyllode size. We compared 3D connectivity to minimum branch water potential and two-dimensional (2D) vessel attributes derived from sections produced by micro-CT. 2D attributes included vessel area, density, vessel size to number ratio (S) and vessel lumen fraction (F). Trees with terete phyllodes had less negative water potentials than broad phyllode variants. 3D xylem connectivity was conserved across all trees regardless of phyllode type or minimum water potential. We also found that xylem connectivity was sensitive to vessel lumen fraction (F) and not the size to number ratio (S) even though F was consistent among species and phyllode variants. Our results demonstrate that differences in phyllode anatomy, and not xylem connectivity, likely explain diversity of drought tolerance among closely related Acacia species. Further analysis using our approach across a broader range of species will improve understanding of adaptations in the xylem networks of arid zone species.

Tree allometries reflect a lifetime of herbivory in an African savanna.

Theories of plant allometry provide a general description of allometric scaling that is supposedly applicable across a wide array of environmental conditions. Scaling theories, however, ignore disturbances such as herbivory in their derivation. Here we examine the influence of herbivores on the scaling of height and diameter of two common African savanna tree species. Using Bayesian piecewise regressions, we show that herbivores modify tree allometry. We also show that the pattern of allometric modification contains information regarding herbivore foraging behavior and the resultant alteration of plant architecture. Interpreting realized allometries in the light of theoretical predictions based on assumptions of zero disturbances may help reveal the degree of herbivore impacts. However, predictions of plant form and function that fail to include disturbances such as herbivory may struggle to find general applicability.

Decomposition of lignin and holocellulose on Acacia mangium leaves and twigs by six fungal isolates from nature.

This research was conducted in the aim of preventing wild fire through reducing potential energy source to become in situ fertilizer. To prevent forest fires by reducing wood waste using lignocellulose-degrading fungi, six fungal isolates were tested for lignin and cellulose-degrading activity with Acacia mangium leaves and twigs over a period of 1 to 3 months. The fungi degraded 8.9-27.1% of the lignin and 14-31% of the holocellulose. The degradation rate varied depending on the fungal species. An increase in incubation time tended to decrease the amounts of holocellulose and lignin. However, the hot water soluble tended to increase following a longer incubation period. From the results obtained here, more time was needed to degrade lignin rather than other components in the sample.

Turning over a new 'leaf': multiple functional significances of leaves versus phyllodes in Hawaiian Acacia koa.

Hawaiian endemic tree Acacia koa is a model for heteroblasty with bipinnately compound leaves and phyllodes. Previous studies suggested three hypotheses for their functional differentiation: an advantage of leaves for early growth or shade tolerance, and an advantage of phyllodes for drought tolerance. We tested the ability of these hypotheses to explain differences between leaf types for potted plants in 104 physiological and morphological traits, including gas exchange, structure and composition, hydraulic conductance, and responses to varying light, intercellular CO(2) , vapour pressure deficit (VPD) and drought. Leaf types were similar in numerous traits including stomatal pore area per leaf area, leaf area-based gas exchange rates and cuticular conductance. Each hypothesis was directly supported by key differences in function. Leaves had higher mass-based gas exchange rates, while the water storage tissue in phyllodes contributed to greater capacitance per area; phyllodes also showed stronger stomatal closure at high VPD, and higher maximum hydraulic conductance per area, with stronger decline during desiccation and recovery with rehydration. While no single hypothesis completely explained the differences between leaf types, together the three hypotheses explained 91% of differences. These findings indicate that the heteroblasty confers multiple benefits, realized across different developmental stages and environmental contexts.

Modelling changes in leaf shape prior to phyllode acquisition in Acacia mangium Willd. seedlings.

The aim of this study was to characterise changes in leaf shape prior to phyllode acquisition along the axes of Acacia mangium seedlings. The study area was located in North Lampung (South Sumatra, Indonesia), where these trees belong to a naturally regenerated stand. A total of 173 seedlings, less than three months old, were described node by node. Leaf shape and leaf length were recorded and the way in which one leaf type succeeded another was modelled using a hidden semi-Markov chain composed of seven states. The phyllotactical pattern was studied using another sample of forty 6-month-old seedlings. The results indicate (i) the existence of successive zones characterised by one or a combination of leaf types, and (ii) that phyllode acquisition seems to be accompanied by a change in the phyllotactical pattern. The concepts of juvenility and heteroblasty, as well as potential applications for taxonomy are discussed.

Breakdown of an ant-plant mutualism follows the loss of large herbivores from an African savanna.

Mutualisms are key components of biodiversity and ecosystem function, yet the forces maintaining them are poorly understood. We investigated the effects of removing large mammals on an ant-Acacia mutualism in an African savanna. Ten years of large-herbivore exclusion reduced the nectar and housing provided by plants to ants, increasing antagonistic behavior by a mutualistic ant associate and shifting competitive dominance within the plant-ant community from this nectar-dependent mutualist to an antagonistic species that does not depend on plant rewards. Trees occupied by this antagonist suffered increased attack by stem-boring beetles, grew more slowly, and experienced doubled mortality relative to trees occupied by the mutualistic ant. These results show that large mammals maintain cooperation within a widespread symbiosis and suggest complex cascading effects of megafaunal extinction.