Spatial and temporal soil moisture resource partitioning by trees and grasses in a temperate savanna, Arizona, USA

Stable isotope analysis was used to determine sources of water used by coexisting trees and grasses in a temperate savanna dominated by Quercus emoryi Torr. We predicted that (1) tree seedlings and bunchgrasses utilize shallow sources of soil water, (2) mature savanna trees use deeper sources of water, and (3) trees switch from shallow to deep water sources within 1 year of germination. We found that Q. emoryi trees, saplings, and seedlings (about 2 months, 1 year, and 2 years old), and the dominant bunchgrass [Trachypogon montufari (H.B.K.) Nees.] utilized seasonally available moisture from different depths within the soil profile depending on size/age relationships. Sapling and mature Q. emoryi acquired water from >50 cm deep, 2-month-old seedlings utilized water from <15 cm, and 1- and 2-year-old seedlings and grasses used water from between 20 cm and 35 cm. This suggests that very young seedlings are decoupled from grasses in this system, which may facilitate germination and early establishment of Q. emoryi within extant stands of native grasses. The potential for subsequent interaction between Q. emoryi and native grasses was evidenced by similar patterns of soil water use by 1- and 2-year-old seedlings and grasses. Q. emoryi seedlings did not switch from shallow to deep sources of soil water within 2 years of germination: water use by these seedlings apparently becomes independent of water use by grasses after 2 years of age. Finally, older trees (saplings, mature trees) use water from deeper soil layers than grasses, which may facilitate the stable coexistence of mature trees and grasses. Potential shifts in the seasonality of precipitation may alter interactions between woody plants and grasses within temperate savannas characterized by bimodal precipitation regimes: reductions in summer precipitation or soil moisture may be particularly detrimental to warm-season grasses and seedlings of Q. emoryi. Received: 21 November 1996 / Accepted: 2 May 1997     

Is there a temporal niche separation in the leaf phenology of savanna trees and grasses?

Aim It has been proposed that, in tropical savannas, trees deploy their leaves earlier in the growing season and grasses deploy their leaves later. This hypothesis implies a mechanism that facilitates the coexistence of trees and grasses in savannas. If true, this hypothesis would also allow algorithms to use differences in the phenological timing of grass and tree leaves to partition the relative contribution of grasses and trees to net primary production. In this study we examine whether a temporal niche separation between grasses and trees exists in savanna. Location A semi‐arid, subtropical savanna, Kruger National Park, South Africa. Methods We use a multi‐spectral camera to track through an entire growing season the normalized difference vegetation index (NDVI) of individual canopies of grasses and trees at eight sites arranged along a precipitation and temperature gradient. Results Among trees, we identified two distinct phenological syndromes: an early flushing syndrome and a late‐flushing syndrome. Leaf flush in the tree strategies appears to pre‐empt rainfall, whereas grass leaf flush follows the rain. The growing season of trees is 20 (late‐flushing trees) to 27 (early flushing trees) days longer than that of the grasses. Main conclusions We show that grasses and trees have different leaf deployment strategies. Trees deployed leaves at lower temperatures than grasses and retained them for longer at the end of the growing season. The timing of the increase in NDVI is, however, similar between grasses and late‐flushing trees and this complicates the separation of grass and tree signals from multi‐spectral satellite imagery.     

Root vascular anatomy predicts maximum growth rates in savanna trees and grasses

Root-based functional traits are relatively overlooked as drivers of savanna plant community dynamics, an important gap in water-limited ecosystems. Recent work has shed light on patterns of trait coordination in roots, but less is known about the relationship between root functional traits, water acquisition, and plant demographic rates. Here, we investigated how fine-root vascular and morphological traits are related in two dominant PFTs (C₃ trees and C₄ grasses from the savanna biome), whether root traits can predict plant relative growth rate (RGR), and whether root trait multivariate relationships differ in trees and grasses. We used root data from 21 tree and 18 grass species grown under greenhouse conditions, and quantified a suite of vascular and morphological root traits. We used a principal components analysis (PCA) to identify common axes of trait variation, compared trait correlation matrices between the two PFTs, and investigated the relationship between PCA axes and individual traits and RGR. We found that there was no clear single axis integrating vascular and morphological traits, but found that vascular anatomy predicted RGR in both trees and grasses. Trait correlation matrices differed in trees and grasses, suggesting potentially divergent patterns of trait coordination between the two functional types. Our results suggested that, despite differences in trait relationships between trees and grasses, root conductivity may constrain maximum growth rate in both PFTs, highlighting the critical role that water relations play in savanna vegetation dynamics and suggesting that root water transport capacity is an important predictor of plant performance in the savanna biome.     

Hyperspectral remote detection of niche partitioning among canopy trees driven by blowdown gap disturbances in the Central Amazon

Advanced recruitment and neutral processes play important roles in determining tree species composition in tropical forest canopy gaps, with few gaps experiencing clear secondary successional processes. However, most studies are limited to the relatively limited spatial scales provided by forest inventory plots, and investigations over the entire range of gap size are needed to better understand how ecological processes vary with tree mortality events. This study employed a landscape approach to test the hypothesis that tree species composition and forest structural attributes differ between large blowdown gaps and relatively undisturbed primary forest. Spectral mixture analysis on hyperspectral satellite imagery was employed to direct field sampling to widely distributed sites, and blowdown plots were compared with undisturbed primary forest plots. Tree species composition and forest structural attributes differed markedly between gap and non-gap sites, providing evidence of niche partitioning in response to disturbance across the region. Large gaps were dominated by classic Neotropical pioneer genera such as Cecropia and Vismia, and average tree size was significantly smaller. Mean wood density of trees recovering in large gaps (0.55 g cm⁻³) was significantly lower than in primary forest plots (0.71 g cm⁻³), a difference similar to that found when comparing less dynamic (i.e., tree recruitment, growth, and mortality) Central Amazon forests with more dynamic Western Amazon forests. Based on results, we hypothesize that the importance of neutral processes weaken, and niche processes strengthen, in determining community assembly along a gradient in gap size and tree mortality intensity. Over evolutionary time scales, pervasive dispersal among colonizers could result in the loss of tree diversity in the pioneer guild through competitive exclusion. Results also underscore the importance of considering disturbance processes across the landscape when addressing forest carbon balance.     

Seed dispersal, seedling establishment and gap partitioning among tropical pioneer trees

1 We examined the abundance and distribution patterns of pioneer seeds in the soil seed bank, and of pioneer seedlings in 53 recently formed gaps, in a 50‐ha forest dynamics plot on Barro Colorado Island (BCI), Panama. The aim was to assess the importance of dispersal limitation (failure of seeds to arrive at all sites suitable for their germination) and establishment limitation (failure of seeds having reached a site to germinate successfully and establish as seedlings) in determining patterns of gap occupancy.
2 The abundance of seeds in the soil seed bank was strongly negatively correlated with seed size, but was not correlated with the abundance of reproductive‐sized adult trees in the plot. In contrast, the abundance of pioneer seedlings > 10 cm height in natural gaps was strongly correlated with adult abundance, but was not correlated with seed size.
3 Seedlings were non‐randomly distributed among gaps, but seedling abundance was not directly related to gap size, and there was no evidence of partitioning of the light environment of gaps by small seedlings. Large differences in growth and mortality rates among species were observed after 1 year, and this may result in the gap size partitioning previously found in saplings of the same species.
4 Seedlings of most species, particularly those with large seeds, were relatively more abundant than expected in gaps close to their conspecific adults. Proximity to reproductives, and by inference dispersal limitation, therefore exerts some effect on seedling distribution. None the less, large differences between seed and seedling abundances for some species, and low seedling occupancy rates in some gaps close to adult conspecifics, suggest that seedling emergence probabilities and species‐specific establishment requirements may also be important determinants of local abundance.     

Niche partitioning of bacterial communities in biological crusts and soils under grasses,shrubs and trees in the Kalahari

The Kalahari of southern Africa is characterised by sparse vegetation interspersed with microbe-dominated biological soil crusts (BSC) which deliver a range of ecosystem services including soil stabilisation and carbon fixation. We characterised the bacterial communities of BSCs (0–1 cm depth) and the subsurface soil (1–2 cm depth) in an area typical of lightly grazed Kalahari rangelands, composed of grasses, shrubs, and trees. Our data add substantially to the limited amount of existing knowledge concerning BSC microbial community structure, by providing the first bacterial community analyses of both BSCs and subsurface soils of the Kalahari region based on a high throughput 16S ribosomal RNA gene sequencing approach. BSC bacterial communities were distinct with respect to vegetation type and soil depth, and varied in relation to soil carbon, nitrogen, and surface temperature. Cyanobacteria were predominant in the grass interspaces at the soil surface (0–1 cm) but rare in subsurface soils (1–2 cm depth) and under the shrubs and trees. Bacteroidetes were significantly more abundant in surface soils of all areas even in the absence of a consolidated crust, whilst subsurface soils yielded more sequences affiliated to Acidobacteria, Actinobacteria, Chloroflexi, and Firmicutes. The common detection of vertical stratification, even in disturbed sites, suggests a strong potential for BSC recovery after physical disruption, however severe depletion of Cyanobacteria near trees and shrubs may limit the potential for natural BSC regeneration in heavily shrub-encroached areas.     

Acoustic niche partitioning among seven cuckoo species in a forest in eastern China

Acoustic communication among birds plays an important role in attracting mates and defending territories. For the successful transmission of songs, individuals of different species often avoid singing at the same time to reduce acoustic interference from background noise and overlapping signals from heterospecifics. Such behavioural acoustic niche partitioning may occur especially among closely related species due to their ecological similarities. In this study, we recorded bird sounds in a subtropical forest in China in May–June 2019 and detected seven cuckoo species. Extracting characteristics of the cuckoo calls, we found that only four of the 21 pairs of species overlapped in frequency range, and 19 pairs were classified accurately using a linear discriminant analysis classifier based on their features. The remaining two species pairs could be separated based on temporal or spatial distribution patterns. We also analysed the temporal distribution patterns and overlap time of the calls, finding that the seven species exhibit partitioning in at least one of three acoustic dimensions (site, frequency, activity time). We conclude that the seven sympatric cuckoo species were strongly partitioned in acoustic signal space and minimally masked each other's signals.     

Root niche partitioning between shallow rooted succulents in a South African semi desert: implications for diversity

With a high percentage of endemics along the west coast of South Africa, especially in the family Aizoaceae, the region is considered one of the earth’s biodiversity hot spots. It has been suggested that the diversity and radiation of the Aizoaceae are coincident with low but predictable rainfall and lack of competition between species. In this study we examine the relationship between water source and the efficiency of PSII photochemistry for representative Aizoaceae and non-Aizoaceae. We do this to determine the extent to which the different genera are adapted to the frequent, low volume, precipitation characteristic of the region and to ascertain the extent to which there is competition for water. Our water isotope results show that the Aizoaceae use shallow surface water while the non-Aizoaceae use a deeper water source. We are however not able to show the extent to which the Aizoaceae utilize fog or dew. Our chlorophyll fluorescence results show that there are no differences in efficiency of PSII photochemistry between the species in the wet season. The decline from wet to dry season for the Aizoaceae is, however, more dramatic than that of the non-Aizoaceae reflecting the differences in rooting depth between the different families. These results suggest that, during the dry season, there is no competition for resources between families but there is competition between species. We conclude that the adaptation to using shallow water, coupled with susceptibility to drought of adult short lived Aizoaceae may be a, mechanism for the diversification of this family.     

Daily and seasonal courses of gas exchange and niche partitioning among coexisting tree species in a tropical montane forest

We investigated the seasonal gas exchange patterns of three different functional types of tropical afromontane trees, namely Podocarpus falcatus (Thunb.) Mirb. (evergreen gymnosperm), Prunus africana (Hook. f.) Kalkm. (broad-leaf evergreen), and Croton macrostachyus Hochst. ex Del. (broad-leaf deciduous) which grow side-by-side in the Munessa forest, southern Central Ethiopia. The hypothesis is that the trees can make different use of the environmental conditions which change seasonally and also from year to year. These changes can be understood as fluctuating niches, the utilization of which allows coexistence through balanced carbon gain. In this study, light and moisture were considered the two main fluctuating niches. Porometry was used to measure the daily and seasonal responses of the leaves to environmental variables under dry and wet season conditions in the course of the climatically differing years 2009 and 2010. Correlations of the patterns of these responses were analyzed by regression analysis. From daily integrated rates of irradiation, photosynthesis and transpiration, light and water use efficiencies of photosynthetic carbon gain and their mutual interdependences were determined. Except an extended dry season in 2009 when C. macrostachyus had shed its foliage, all three species continued photosynthetic net CO₂ uptake concomitant with transpiration at all seasons, although at varying rates. Ecophysiological performance of P. falcatus leaves is mainly light-driven and responds relatively little to a change in moisture conditions. Its carbon and water relations are striving for stability rather than flexibility. As of a typically subdominant species of the forest, P. africana leaves can efficiently use low light intensities but suffer from photoinhibition at full light. Their performance, showing more dynamic response to the environment than P. falcatus, P. africana appears driven more by moisture than by light. Compared to the two evergreens C. macrostachyus exhibited the highest flexibility in its leaves’ physiological responses to environmental conditions, in particular to the light climate which is additionally potentiated by the fast turnover of its foliage. This species optimizes its carbon gain during the wet season and during the early dry period when cloud cover is minimal. Our findings thus reveal that elasticity of the response to fluctuating environmental conditions is an additional aspect in the assessment of the utilization of temporally fluctuating niches by adult tree individuals.     

Strontium as a tracer for calcium: uptake,transport and partitioning within tomato plants

Plant and Soil - Calcium (Ca2+) is a major structural plant nutrient whose low mobility in the phloem causes deleterious nutritional disorders in non-transpiring organs. Since strontium (Sr2+) and...     

Interactions among fungal endophytes,grasses and herbivores

The interaction between two species often depends on the presence or absence of a third species. One widespread three-species interaction involves fungal endophytes infecting grasses and the herbivores that feed upon them. The endophytes are allied with the fungal family Clavicipitaceae and grow systemically in intercellular spaces in above-ground plant tissues including seeds. Like relatedClaviceps species, the endophytes produce a variety of alkaloids that make the host plants toxic or distasteful to herbivores. A large number of grass species are infected, especially cool-season grasses in temperate areas. Field and laboratory studies have shown that herbivores avoid infected plants in choice trials and suffer increased mortality and decreased growth on infected grasses in feeding experiments. Resistance to herbivores may provide a selective advantage to infected plants in competitive interactions with noninfected plants. Recent studies have shown that differential herbivory can reverse competitive hierarchies among plant species. Both endophyte-infected and noninfected tall fescue grass (Festuca arundinacea) are outcompeted by orchardgrass (Dactylis glomerata) in the absence of insect herbivory. However, when herbivores are present infected tall fescue outcompetes orchardgrass. These results suggest that the frequency of infection in grass species and grassland communities will increase over time. Several studies are reviewed illustrating increases in infection frequency within grass populations subject to herbivore pressure. Endophytic fungi may be important regulators of plant-herbivore interactions and so indirectly affect the structure and dynamics of plant communities.     

Water source partitioning among trees growing on shallow karst soils in a seasonally dry tropical climate

The sources of water used by woody vegetation growing on karst soils in seasonally dry tropical regions are little known. In northern Yucatan (Mexico), trees withstand 4–6 months of annual drought in spite of the small water storage capacity of the shallow karst soil. We hypothesized that adult evergreen trees in Yucatan tap the aquifer for a reliable supply of water during the prolonged dry season. The naturally occurring concentration gradients in oxygen and hydrogen stable isotopes in soil, bedrock, groundwater and plant stem water were used to determine the sources of water used by native evergreen and drought-deciduous tree species. While the trees studied grew over a permanent water table (9–20 m depth), pit excavation showed that roots were largely restricted to the upper 2 m of the soil/bedrock profile. At the peak of the dry season, the δ¹⁸O signatures of potential water sources for the vegetation ranged from 4.1 ± 1.1‰ in topsoil to −4.3 ± 0.1‰ in groundwater. The δ¹⁸O values of tree stem water ranged from −2.8 ± 0.3‰ in Talisia olivaeformis to 0.8 ± 1‰ in Ficus cotinifolia, demonstrating vertical partitioning of soil/bedrock water among tree species. Stem water δ¹⁸O values were significantly different from that of groundwater for all the tree species investigated. Stem water samples plotted to the right of the meteoric water line, indicating utilization of water sources subject to evaporative isotopic enrichment. Foliar δ¹³C in adult trees varied widely among species, ranging from −25.3 ± 0.3‰ in Enterolobium cyclocarpum to −28.7 ± 0.4‰ in T. olivaeformis. Contrary to initial expectations, data indicate that native trees growing on shallow karst soils in northern Yucatan use little or no groundwater and depend mostly on water stored within the upper 2–3 m of the soil/bedrock profile. Water storage in subsurface soil-filled cavities and in the porous limestone bedrock is apparently sufficient to sustain adult evergreen trees throughout the pronounced dry season.     

Variation partitioning as a tool to distinguish between niche and neutral processes

We assess the potential of different forms of variation partitioning to distinguish between environmental control and dispersal limitation in communities structured by combinations of niche and neutral processes. Simulation data reveal interactions between dispersal limitation, environmental control, and the spatial structure of environmental factors in the detected levels of variance fractions. The degree of dispersal limitation contributes to both the pure environmental and pure spatial variance partitions. This undermines the common practice of interpreting these partitions as direct expressions of niche and neutral processes, respectively. Furthermore, the proportion of variation attributed to environmental variation depends not only on the strength of environmental control, but also on the specific spatial configuration of the environmental variable. This has important implications for the interpretation of empirical studies. In particular, use of these analytical techniques to compare processes governing community structure among different study systems is unwarranted, as the results will reflect not only differences in the strength of the processes of interest, but also the influence of the unique spatial arrangement of the environmental variables in each system.     

Ontogenetic niche shifts and resource partitioning in a subarctic piscivore fish guild

The feeding ecology of three piscivorous fish species (perch (Perca fluviatilis), pike (Esox lucius) and burbot (Lota lota)), was studied in the subarctic Pasvik watercourse (69 °N), northern Norway and Russia. All three species primarily occupied the benthic habitats in the watercourse. Perch and burbot exhibited distinct ontogenetic niche shifts in food resource use, perch changing from a dominance of zooplankton to zoobenthos to fish, and burbot from zoobenthos to fish. Fish prey dominated the diet of all the investigated size-classes of pike, but small-sized pike (<20 cm) were not represented in the sample. Fish prey size was positively related to predator size in all three species. Whitefish (Coregonus lavaretus) was the dominant prey of pike and large-sized burbot and perch. Nine-spined sticklebacks (Pungitius pungitius) was also an important prey and appeared to be a dietary stepping-stone enhancing the transition from invertebrate feeding to consumption of large-sized whitefish prey for all three predators. A cluster analysis separated the different size groups of the three predator species into five functional feeding groups, most of them containing two or all three species. Within these feeding groups, and especially among the piscivorous size-classes, there was a strong and significant interspecific overlap in prey selection, and the dietary similarities between the species were in general much larger than the intraspecific similarities between ontogenetic stages. All three piscivorous species are important top predators in the aquatic food web of the watercourse, and their ontogenetic diet shifts and resource partitioning patterns generate a substantial food web complexity in this subarctic ecosystem.     

Stream microhabitat selectivity,resource partitioning,and niche shifts in grazing caddisfly larvae

Microhabitat selectivity, resource partitioning, and niche shifts in five species of grazing caddisfly larvae (Glossosoma califica, G. penitum, Dicosmoecus gilvipes, Neophylax rickeri, and N. splendens) were quantified by underwater measurement of microhabitat availability and utilization in three northern California streams. The microhabitat parameters water depth and velocity and rock size, roughness, and slope were measured. Comparisons of habitat available to habitat used revealed significant selection for at least two microhabitat parameters by each population, with depth and velocity being the most important. Comparisons of habitat used by different species showed significant partitioning of at least two microhabitat parameters at each site, with depth being partitioned at all sites. Non-parametric discriminant analysis revealed significant microhabitat partitioning on a multivariate level at two sites. Comparisons of habitat used at different sites quantified a major niche shift by D. gilvipes in its preference for riffles versus pools. Size-selective predation by dippers (Cinclus mexicanus) and steelhead (Salmo gairdneri gairdneri) is proposed as a hypothesis to explain the observed resource partitioning and niche shift.     

Root hydraulic conductivity and whole-plant water balance in tropical saplings following a shade-to-sun transfer

We hypothesized that pioneer and late successional species show different morphological and physiological responses in water use after gap formation. The magnitude of the responses was compared between two pioneer species (Macaranga gigantea and Trema orientalis) and four late successional species (Shorea sp.), in an experiment in which saplings were transferred from shade to sun. Although transpiration demand increased following the transfer, root hydraulic conductivity (Lp_r) decreased. Lp_r was sensitive to brief treatments with HgCl₂ (a specific inhibitor of aquaporins). This allows Lp_r to be divided into two components: cell-to-cell and apoplastic pathways. The Lp_r of cell-to-cell pathway decreased in all species following the transfer, relating to aquaporin depression in roots. Following the transfer, leaf osmotic potentials at full hydration decreased and both leaf mass per area [leaf mass/leaf area (LMA)] and fine-root surface area/leaf surface area (root SA/leaf SA) increased in almost all species, allowing saplings to compensate for the decrease in Lp_r. Physiologically, pioneer species showed larger decreases in Lp_r and more effective osmotic adjustment than late successional species, and morphologically, pioneer species showed larger increases in root SA/leaf SA and LMA. Water balance at the whole-plant level should be regulated by coupled responses between the aboveground and the belowground parts. Interspecific differences in responses after gap formation suggest niche differentiation in water use between pioneer and late successional species in accordance with canopy-gap size.     

Integrating phylogeny,geographic niche partitioning and secondary metabolite synthesis in bloom-forming Planktothrix

Toxic freshwater cyanobacteria form harmful algal blooms that can cause acute toxicity to humans and livestock. Globally distributed, bloom-forming cyanobacteria Planktothrix either retain or lose the mcy gene cluster (encoding the synthesis of the secondary metabolite hepatotoxin microcystin or MC), resulting in a variable spatial/temporal distribution of (non)toxic genotypes. Despite their importance to human well-being, such genotype diversity is not being mapped at scales relevant to nature. We aimed to reveal the factors influencing the dispersal of those genotypes by analyzing 138 strains (from Europe, Russia, North America and East Africa) for their (i) mcy gene cluster composition, (ii) phylogeny and adaptation to their habitat and (iii) ribosomally and nonribosomally synthesized oligopeptide products. Although all the strains from different species contained at least remnants of the mcy gene cluster, various phylogenetic lineages evolved and adapted to rather specific ecological niches (for example, through pigmentation and gas vesicle protein size). No evidence for an increased abundance of specific peptides in the absence of MC was found. MC and peptide distribution rather depended on phylogeny, ecophysiological adaptation and geographic distance. Together, these findings provide evidence that MC and peptide production are primarily related to speciation processes, while within a phylogenetic lineage the probability that strains differ in peptide composition increases with geographic distance.