Disentangling phylogenetic from non‐phylogenetic functional structure of bird assemblages in a tropical dry forest

Understanding the factors driving assembling structure of ecological communities remains a fundamental problem in ecology, especially when focusing on ecological and evolutionary relatedness among species rather than on their taxonomic identity. It remains critical though to separate the patterns and drivers of phylogenetic and functional structures, because traits are phylogenetically constrained, but phylogeny alone does not fully reflect trait variability among species. Using birds from the Brazilian dry forest as a study case, we employed two different approaches to decompose functional structure into its components that are shared and non‐shared with the phylogenetic structure. We investigated the spatial pattern and environmental hypotheses for these phylogenetically constrained and unconstrained aspects of functional structure, including climate‐induced physiological constraints, historical climatic stability, resource availability and habitat partitioning. We found only partial congruence between the two methods of structure decomposition. Still, we found a differential effect of factors on specific components of functional structure of bird assemblages. While climate affects phylogenetically constrained traits through endurance, habitat partitioning (especially forest cover) affects the functional structure that is independent of phylogeny. With this strategy, we were able to decompose the patterns and drivers of the functional structure of birds along a semiarid gradient and showed that the decomposition of the functional structure into its phylogenetic and non‐phylogenetic counterparts can offer a more complete portrait of the assembling rules in ecological communities. We claim for a further development and use of this sort of strategy to investigate assembling rules in ecological communities.     

Functional organization of woody plant assemblages along precipitation and human disturbance gradients in a seasonally dry tropical forest

Chronic anthropogenic disturbances (CAD) and rainfall are important drivers of plant community assembly, but little is known about the role played by inter‐ and intraspecific trait variation as communities respond to these pervasive forces. Here, we examined the hypothesis that lower precipitation and higher CAD reduce both intra‐ and interspecific trait variation in Caatinga dry forests. We sampled woody plants across 15 plots along precipitation and CAD gradients and measured resource‐use traits. The effects of precipitation and CAD on RaoQ functional diversity were decomposed into species turnover and intraspecific variability. We used “T‐statistics” to assess the trait sorting from the regional pool to local communities (i.e., external filtering), and within‐community forces leading to low trait overlap (i.e., internal filtering) at individual and species levels. Intraspecific variability explained at least one‐third of the total trait variation and 46% of variation in multitrait diversity across communities. Increasing disturbance reduced multitrait diversity, while precipitation affected some particular traits, such as wood density. Overall, precipitation determined species sorting across communities, while disturbance relaxed internal filters, leading to higher trait overlap within communities due to higher intraspecific variability. Our results suggest that the woody Caatinga flora contains a substantial amount of both inter‐ and intraspecific trait variation. This variation is not randomly distributed within and across communities, but varies according to rainfall conditions and disturbance intensity. These findings reinforce the emerging idea that human disturbances can reorganize plant communities at multiple scales and highlight trait variability as a key biological asset for the resilience of dry forests.     

Functional diversity of macromycete communities along an environmental gradient in a Mexican seasonally dry tropical forest

Macromycetes are important for ecosystem functioning due to their role in the nutrient cycling, and their function as pathogens and mutualists. Diversity metrics based on functional traits are robust predictors of ecosystem functionality since they incorporate an evolutionary and ecologic background. We examined diversity patterns of macrofungi using functional trait-based metrics of diversity along an altitudinal gradient in a seasonally dry tropical forest in southern Mexico. Our findings show that: (1) functional diversity varies with elevation, relating more to climatic variables than to vegetation structure; (2) functional diversity indexes exhibited contrasting patterns, so measures reflecting heterogeneity on trait abundance and niche complementarity tend to increase with elevation, whereas the measure of trait evenness decreases; and (3) functional diversity patterns depend on the type of functional trait considered and how they respond to environmental conditions. Our results indicate that functional diversity analyses help understanding of how macrofungal communities respond to environmental variation.     

Effects of seasonal precipitation change on soil respiration processes in a seasonally dry tropical forest

Precipitation is projected to change intensity and seasonal regime under current global projections. However, little is known about how seasonal precipitation changes will affect soil respiration, especially in seasonally dry tropical forests. In a seasonally dry tropical forest in South China, we conducted a precipitation manipulation experiment to simulate a delayed wet season (DW) and a wetter wet season (WW) over a three‐year period. In DW, we reduced 60% throughfall in April and May to delay the onset of the wet season and irrigated the same amount water into the plots in October and November to extend the end of the wet season. In WW, we irrigated 25% annual precipitation into plots in July and August. A control treatment (CT) receiving ambient precipitation was also established. Compared with CT, DW significantly increased soil moisture by 54% during October to November, and by 30% during December to April. The treatment of WW did not significantly affect monthly measured soil moisture. In 2015, DW significantly increased leaf area index and soil microbial biomass but decreased fine root biomass. In contrast, WW significantly decreased fine root biomass and forest floor litter stocks. Soil respiration was not affected by DW, which could be attributed to the increased microbial biomass offsetting the decrease in fine root biomass. In contrast, WW significantly increased soil respiration from 3.40 to 3.90 μmol m^?2 s^?1 in the third year, mainly due to the increased litter decomposition and soil pH (from 4.48 to 4.68). The present study suggests that both a delayed wet season and a wetter wet season will have significant impacts on soil respiration‐associated ecosystem components. However, the ecosystem components can respond in different directions to the same change in precipitation, which ultimately affected soil respiration.     

Influence of rainfall and grazing on herbage dynamics in a seasonally dry tropical savanna

Species composition and herbage dynamics in relation to rainfall variability and cattle grazing were studied in permanently protected, grazed, and temporarily fenced treatments on three sites in a seasonally dry tropical savanna. Permanently protected sites, established between 1979 and 1984, were 55–79% similar with each other in species composition, and 14–25% similar with grazed sites during the period 1986–1988. Similarity among grazed sites was only 36–43%. Number of species was greater in the grazed treatment than in the permanently protected treatment. The percentages of annual grasses and non-leguminous forbs were greater in grazed savanna than in permanently protected savanna. Species diversity was higher in grazed savanna than in the corresponding permanently protected savanna. Species the two annual cycles studied, peak live shoot biomass was 614 g m^-2 in permanently protected savanna, 109 g m^-2 in grazed savanna, and 724 g m^-2 in temporarily fenced savanna. Live shoot biomass in temporarily fenced savanna was 18 to 44% greater than in permanently protected savanna. Peak canopy biomass ranged from 342 to 700 g m^-2 in permanently protected savanna. It was related with total rainy season rainfall, and was particularly sensitive to late rainy season rainfall. On the other hand, peak canopy biomass in grazed savanna ranged from 59 to 169 g m^-2 and was related to grazing intensity rather than either total rainy season rainfall or late rainy season rainfall. Coefficient of variation of green biomass in permanently protected savanna was related with rainfall variability indicating it to be a pulsed system which responds quickly to rainfall events. Biomass of woody species ranged from 2466 to 5298 g m^-2 in permanently protected savanna and from 744 to 1433 g m^–2 in the grazed savanna. Green foliage biomass was 3.7 to 6.4% of the woody biomass in permanently protected and 5.6 to 5.9% in grazed savanna, and supplements substantially the fodder resource during the dry periods of the year.     

Woody plant diversity in relation to environmental factors in a seasonally dry tropical forest landscape

Questions

Water availability is known to be a first‐order driver of plant diversity; yet water also affects fire regimes and soil fertility, which, in turn, affect plant diversity. We examined how precipitation, fire and soil properties jointly determine woody plant diversity. Specifically, we asked how woody plant diversity varies along a sharp precipitation gradient (about 600–1,800 mm mean annual precipitation [MAP ]within a ~45‐km distance) exhibiting considerable variation in long‐term fire burn frequency and soil fertility, in a southern Indian seasonally dry tropical forest (SDTF ) landscape.

Location

Mudumalai, Western Ghats, India.

Methods

Woody plants ≥1‐cm DBH were enumerated in 19 1‐ha permanent plots spanning a range of tropical vegetation types from dry thorn forest, through dry and moist deciduous forest to semi‐evergreen forest. Burn frequencies were derived from annual fire maps. Six measures of surface soil properties – total exchangeable bases (Ca + Mg + K), organic carbon (OC ), total N, pH , plant available P and micronutrients (Fe + Cu + Zn + Mn) were used in the analyses. Five measures of diversity – species richness, Shannon diversity, the rarefied/extrapolated versions of these two measures, and Fisher's α – were modelled as functions of MAP , annual fire burn frequency and the principal components of soil properties.

Results

Most soil nutrients and OC increased with MAP , except in the wettest sites. Woody productivity increased with MAP , while fire frequency was highest at intermediate values of MAP . Woody plant diversity increased with MAP but decreased with increasing fire frequency, resulting in two local diversity maxima along the MAP gradient – in the semi‐evergreen and dry thorn forest – separated by a low‐diversity central region in dry deciduous forest where fire frequency was highest. Soil variables were, on the whole, less strongly correlated with diversity than MAP .

Conclusions

Although woody plant diversity in this landscape, representative of regional SDTF s, is primarily limited by water availability, our study emphasizes the role of fire as a potentially important second‐order driver that acts to reduce diversity in this landscape.

     

Floral traits and pollination systems in the Caatinga, a Brazilian tropical dry forest

BACKGROUND AND AIMS: Pollination is a critical stage in plant reproduction and thus in the maintenance and evolution of species and communities. The Caatinga is the fourth largest ecosystem in Brazil, but despite its great extent and its importance few studies providing ecological information are available, with a notable lack of work focusing on pollination biology. Here, general data are presented regarding the frequency of pollination systems within Caatinga communities, with the aim of characterizing patterns related to floral attributes in order to make possible comparisons with data for plant communities in other tropical areas, and to test ideas about the utility of syndromes. This paper also intends to provide a reference point for further studies on pollination ecology in this threatened ecosystem. METHODS: The floral traits and the pollination systems of 147 species were analysed in three areas of Caatinga vegetation in northeastern Brazil, and compared with world-wide studies focusing on the same subject. For each species, floral attributes were recorded as form, size, colour, rewards and pollination units. The species were grouped into 12 guilds according to the main pollinator vector. Analyses of the frequencies of the floral traits and pollination systems were undertaken. KEY RESULTS: Nectar and pollen were the most common floral resources and insect pollination was the most frequent, occurring in 69.9 % of the studied species. Of the entomophilous species, 61.7 % were considered to be melittophilous (43.1 % of the total). Vertebrate pollination occurred in 28.1 % of the species (ornithophily in 15.0 % and chiropterophily in 13.1 %), and anemophily was recorded in only 2.0 %. CONCLUSIONS: The results indicated that the pollination systems in Caatinga, despite climatic restrictions, are diversified, with a low percentage of generalist flowers, and similar to other tropical dry and wet forest communities, including those with high rainfall levels.     

Water uptake and transport in lianas and co-occurring trees of a seasonally dry tropical forest

Water uptake and transport were studied in eight liana species in a seasonally dry tropical forest on Barro Colorado Island, Panama. Stable hydrogen isotope composition (D) of xylem and soil water, soil volumetric water content (_v), and basal sap flow were measured during the 1997 and 1998 dry seasons. Sap flow of several neighboring trees was measured to assess differences between lianas and trees in magnitudes and patterns of daily sap flow. Little seasonal change in _v was observed at 90–120 cm depth in both years. Mean soil water D during the dry season was –19 at 0–30 cm, –34 at 30–60 cm, and –50 at 90–120 cm. Average values of xylem D among the liana species ranged from –28 to –44 during the middle of the dry season, suggesting that water uptake was restricted to intermediate soil layers (30–60 cm). By the end of the dry season, all species exhibited more negative xylem D values (–41 to –62), suggesting that they shifted to deeper water sources. Maximum sap flux density in co-occurring lianas and trees were comparable at similar stem diameter (DBH). Furthermore, lianas and trees conformed to the same linear relationship between daily sap flow and DBH. Our observations that lianas tap shallow sources of soil water at the beginning of the dry season and that sap flow is similar in lianas and trees of equivalent stem diameter do not support the common assumptions that lianas rely primarily on deep soil water and that they have higher rates of sap flow than co-occurring trees of similar stem size.     

Plant functional types broadly describe water use strategies in the Caatinga,a seasonally dry tropical forest in northeast Brazil

1. In seasonally dry tropical forests, plant functional type can be classified as deciduous low wood density, deciduous high wood density, or evergreen high wood density species. While deciduousness is often associated with drought‐avoidance and low wood density is often associated with tissue water storage, the degree to which these functional types may correspond to diverging and unique water use strategies has not been extensively tested.
2. We examined (a) tolerance to water stress, measured by predawn and mid‐day leaf water potential; (b) water use efficiency, measured via foliar δ¹³C; and (c) access to soil water, measured via stem water δ¹⁸O.
3. We found that deciduous low wood density species maintain high leaf water potential and low water use efficiency. Deciduous high wood density species have lower leaf water potential and variable water use efficiency. Both groups rely on shallow soil water. Evergreen high wood density species have low leaf water potential, higher water use efficiency, and access alternative water sources. These findings indicate that deciduous low wood density species are drought avoiders, with a specialized strategy for storing root and stem water. Deciduous high wood density species are moderately drought tolerant, and evergreen high wood density species are the most drought tolerant group.
4. Synthesis. Our results broadly support the plant functional type framework as a way to understand water use strategies, but also highlight species‐level differences.

     

Taxonomic and functional ant diversity along a secondary successional gradient in a tropical forest

The taxonomic diversity (TD) of tropical flora and fauna tends to increase during secondary succession. This increase may be accompanied by changes in functional diversity (FD), although the relationship between TD and FD is not well understood. To explore this relationship, we examined the correlations between the TD and FD of ants and forest age in secondary forests at the α‐ and β‐diversity levels using single‐ and multi‐trait‐based approaches. Our objectives were to understand ant diversity patterns and to evaluate the role of secondary forests in the conservation of biodiversity and in the resilience of tropical forests. Ant assemblages were sampled across a chronosequence in the Lacandon region, Mexico. All species were characterized according to 12 functional ecomorphological traits relevant to their feeding behavior. We found that TD and FD were related to forest age at the alpha level, but not at the beta level. α‐functional richness and divergence increased linearly with species richness and diversity, respectively. Also, the relationship between taxonomic and functional turnover was linear and positive. Our results indicated that functional traits were complementary across the chronosequence. The increase in FD was mainly driven by the addition of rare species with relevant traits. The older secondary forests did not recover all of the functions of old growth forest but did show a tendency to recovery. Because older successional stages support more TD and FD, we suggest developing agriculture and forestry management practices that facilitate rapid post‐agricultural succession and thereby better preserve the functionality of tropical forests.     

Plant sexual systems and a review of the breeding system studies in the Caatinga, a Brazilian tropical dry forest

BACKGROUNDS AND AIMS: The reproductive biology of a community can provide answers to questions related to the maintenance of the intraspecific pollen flow and reproductive success of populations, sharing and competition for pollinators and also questions on conservation of natural habitats affected by fragmentation processes. This work presents, for the first time, data on the occurrence and frequency of plant sexual systems for Caatinga communities, and a review of the breeding system studies of Caatinga species. METHODS: The sexual systems of 147 species from 34 families and 91 genera occurring in three Caatinga areas in north-eastern Brazil were analysed and compared with worldwide studies focusing on reproductive biology of different tropical communities. KEY RESULTS: The frequency of hermaphrodite species was 83.0 % (122 species), seven of these (or 4.8 % of the total) being heterostylous. Monoecy occurred in 9.5 % (14) of the species, and andromonoecy in 4.8 % (seven). Only 2.7 % (four) of the species were dioecious. A high percentage of hermaphrodite species was expected and has been reported for other tropical ecosystems. With respect to the breeding system studies with species of the Caatinga, the authors' data for 21 species and an additional 18 species studied by others (n = 39) revealed a high percentage (61.5 %) of obligatory self-incompatibility. Agamospermy was not recorded among the Caatinga studied species. CONCLUSIONS: The plant sexual systems in the Caatinga, despite the semi-arid climate, are similar to other tropical dry and wet forest communities, including those with high rainfall levels, except for the much lower percentage of dioecious species. The high frequency of self-incompatible species is similar to that reported for Savanna areas in Brazil, and also for dry (deciduous and semideciduous) and humid tropical forest communities.     

Response of soil respiration to simulated N deposition in a disturbed and a rehabilitated tropical forest in southern China

Responses of soil respiration (CO₂ emission) to simulated N deposition were studied in a disturbed (reforested forest with previous understory and litter harvesting) and a rehabilitated (reforested forest with no understory and litter harvesting) tropical forest in southern China from October 2005 to September 2006. The objectives of the study were to test the following hypotheses: (1) soil respiration is higher in rehabilitated forest than in disturbed forest; (2) soil respiration in both rehabilitated and disturbed tropical forests is stimulated by N additions; and (3) soil respiration is more sensitive to N addition in disturbed forest than in rehabilitated forest due to relatively low soil nutrient status in the former, resulting from different previous human disturbance. Static chamber and gas chromatography techniques were employed to quantify the soil respiration, following different N treatments (Control, no N addition; Low-N, 5 g N m⁻² year⁻¹; Medium-N, 10 g N m⁻² year⁻¹), which had been applied continuously for 26 months before the respiration measurement. Results showed that soil respiration exhibited a strong seasonal pattern, with the highest rates observed in the hot and wet growing season (April–September) and the lowest rates in winter (December–February) in both rehabilitated and disturbed forests. Soil respiration rates exhibited significant positive exponential relationship with soil temperature and significant positive linear relationship with soil moisture. Soil respiration was also significantly higher in the rehabilitated forest than in the disturbed forest. Annual mean soil respiration rate in the rehabilitated forest was 20% lower in low-N plots (71 ± 4 mg CO₂-C m⁻² h⁻¹) and 10% lower in medium-N plots (80 ± 4 mg CO₂-C m⁻² h⁻¹) than in the control plots (89 ± 5 mg CO₂-C m⁻² h⁻¹), and the differences between the control and low-N or medium-N treatments were statistically significant. In disturbed forest, annual mean soil respiration rate was 5% lower in low-N plots (63 ± 3 mg CO₂-C m⁻² h⁻¹) and 8% lower in medium-N plots (61 ± 3 mg CO₂-C m⁻² h⁻¹) than in the control plots (66 ± 4 mg CO₂-C m⁻² h⁻¹), but the differences among treatments were not significant. The depressed effects of experimental N deposition occurred mostly in the hot and wet growing season. Our results suggest that response of soil respiration to elevated N deposition in the reforested tropical forests may vary depending on the status of human disturbance. Responsible Editor: Hans Lambers.     

Variations in soil carbon sequestration and their determinants along a precipitation gradient in seasonally dry tropical forest ecosystems

The effect of precipitation regime on the C cycle of tropical forests is poorly understood, despite the existence of models that suggest a drier climate may substantially alter the source‐sink function of these ecosystems. Along a precipitation regime gradient containing 12 mature seasonally dry tropical forests growing under otherwise similar conditions (similar annual temperature, rainfall seasonality, and geological substrate), we analyzed the influence of variation in annual precipitation (1240 to 642 mm) and duration of seasonal drought on soil C. We investigated litterfall, decomposition in the forest floor, and C storage in the mineral soil, and analyzed the dependence of these processes and pools on precipitation. Litterfall decreased slightly – about 10% – from stands with 1240 mm yr^?1 to those with 642 mm yr^?1, while the decomposition decreased by 56%. Reduced precipitation strongly affected C storage and basal respiration in the mineral soil. Higher soil C storage at the drier sites was also related to the higher chemical recalcitrance of litter (fine roots and forest floor) and the presence of charcoal across sites, suggesting an important indirect influence of climate on C sequestration. Basal respiration was controlled by the amount of recalcitrant organic matter in the mineral soil. We conclude that in these forest ecosystems, the long‐term consequences of decreased precipitation would be an increase in organic layer and mineral soil C storage, mainly due to lower decomposition and higher chemical recalcitrance of organic matter, resulting from changes in litter composition and, likely also, wildfire patterns. This could turn these seasonally dry tropical forests into significant soil C sinks under the predicted longer drought periods if primary productivity is maintained.     

Soil water, nutrient availability and sapling survival under organic and polyethylene mulch in a seasonally dry tropical forest

We examine the effect of mulches on the soil volumetric water content (SVWC), pH, carbon (C), total and mineral (NH₄ and NO₃) nitrogen (N), total and bicarbonate phosphorus (P), and on the survival and relative growth rate of three species, Ipomea wolcottiana Rose, Lonchocarpus eriocarinalis Micheli and Caesalpinia eriostachys Benth, in a degraded seasonally dry tropical forest (SDTF) area. Our study year was unusually dry, with only half of the mean annual rainfall. Sixteen plots (5 × 6 m) for each of our four treatments, mulches with alfalfa (Medicago sativa L.) straw, forest litter (SDTF litter), polyethylene and bare soil (control), were used. In each plot, 20 tree saplings were planted of each species. The SVWC was higher in plots mulched with polyethylene than in bare soil plots. The soil pH did not change with mulching, and there were no differences between treatments in the concentrations of soil organic C, total N, NO₃ and total P. However, soil concentrations of NH₄ were highest in plots with alfalfa straw and of bicarbonate P in plots with polyethylene. Sapling survival was higher in polyethylene mulch plots than in other mulching treatments, in the order I.␣wolcottiana > C. eriostachys > L. eriocarinalis. Sapling survival under organic mulches, alfalfa straw and forest litter were similar, and lowest in bare soil. The relative growth rate followed the order L. eriocarinalis < C. eriostachys < I. wolcotiana, and the growth rate of all species was greatest under polyethylene mulch. We conclude that a combination of polyethylene mulch with species of high growth rate is best for restoring seasonally dry tropical areas.     

Environment differentially affects the functional and phylogenetic structures of plant communities in a dry evergreen Afromontane tropical forest

Testing how local environmental conditions influence plant community assembly is important to understand the underlying mechanisms that promote and/or maintain biodiversity. Functional traits are used to find the broad spectrum of resource use strategies that plants use to respond to environmental variation. The patterns and drivers of plant community assembly through the lens of traits and phylogeny; however, remain to be studied in a uniquely biodiversity rich but poorly known fragmented dry Afromontane forest of Ethiopia. Here, we combined trait and community phylogenetic data from thirty sampling plots of 20 × 20 m size to determine the functional and phylogenetic structures and their drivers in a fragmented, human-dominated dry evergreen Afromontane forest. We found phylogenetic and functional clustering of plants in which the effect of environment was found to be trait specific. A weak phylogenetic signal for traits was detected suggesting that species resource use strategies may not be inferred using species phylogenetic distance. Additionally, we found functional traits to be weak in predicting species abundance distribution. Overall, while this study shows a non-random community assembly pattern, it also highlights the importance of deterministic processes being trait specific.     

Leaf litter fungi in a Central Amazonian forest: the influence of rainfall,soil and topography on the distribution of fruiting bodies

Fungi are important components of tropical ecosystems, especially in the recycling of nutrients. However, there is little information on how fungal diversity is structured at scales suitable to plan their conservation. We tested if the distribution of fruiting bodies of litter fungi was random in the landscape (over 25 km²) in a tropical evergreen forest in Central Amazonia. We used linear regressions to evaluate the influence of rainfall, soil characteristics and topography on morphospecies richness and composition. Fungi were collected twice in thirty 0.25 × 250 m plots. Short-term rainfall was represented by the cumulative rainfall in the three days before each plot was surveyed. Plots were classified in two groups based on cumulative rainfall. Clay content in soil and rainfall influenced morphospecies richness, but responses to edaphic factors depended on rainfall. Wetter periods apparently decreased limiting moisture conditions in higher areas, allowing fungal activity and fruiting body production. Morphospecies composition was influenced by clay content, but influence on fungi was probably indirect as clay content was correlated with altitude, plant community and nitrogen availability. Our results suggest that the species of litter fungi are not randomly distributed in the landscape. Furthermore, they indicate that it is viable to conduct mesoscale evaluations of fungal diversity, if the temporal and spatial variation and their interaction are taken into account.