Identity and relationships of the Arboreal Caatinga among other floristic units of seasonally dry tropical forests (SDTFs) of north-eastern and Central Brazil

The tree species composition of seasonally dry tropical forests (SDTF) in north-eastern and central Brazil is analyzed to address the following hypotheses: (1) variations in species composition are related to both environment (climate and substrate) and spatial proximity; (2) SDTF floristic units may be recognized based on peculiar composition and environment; and (3) the Arboreal Caatinga, a deciduous forest occurring along the hinterland borders of the Caatinga Domain, is one of these units and its flora is more strongly related to the caatinga vegetation than to outlying forests. The study region is framed by the Brazilian coastline, 50th meridian west and 21st parallel south, including the Caatinga Domain and extensions into the Atlantic Forest and Cerrado Domains. Multivariate and geostatistic analyses were performed on a database containing 16,226 occurrence records of 1332 tree species in 187 georeferenced SDTF areas and respective environmental variables. Tree species composition varied significantly with both environmental variables and spatial proximity. Eight SDTF floristic units were recognized in the region, including the Arboreal Caatinga. In terms of species composition, its tree flora showed a stronger link with that of the Cerrado Dry Forest Enclaves. On the other hand, in terms of species frequency across sample areas, the links were stronger with two other units: Rock Outcrops Caatinga and Agreste and Brejo Dry Forests. There is a role for niche-based control of tree species composition across the SDTFs of the region determined primarily by the availability of ground water across time and secondarily by the amount of soil mineral nutrients. Spatial proximity also contributes significantly to the floristic cohesion of SDTF units suggesting a highly dispersal-limited tree flora. These units should be given the status of eco-regions to help driving the conservation policy regarding the protection of their biodiversity.     

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.

     

Dynamics of viable nitrifier community, N-mineralization and nitrification in seasonally dry tropical forests and savanna

The study was conducted in Vindhyan region, to assess the N-mineralization, nitrification and size of viable community of ammonium- and nitrite-oxidizing bacteria as affected by different sites and seasons. Six different ecosystems (four forests and two savannas), which differ in terms of topography, vegetation and moisture status, were selected for the present study. The soils of the study sites differ significantly in its physico-chemical properties. The savanna site had significantly higher pH (7.2), bulk density (1.37 g cm(-3)) and silt content (67.80%) but lower water holding capacity (1.37%), total-C (16,356 microg g(-1) dry soil), N (1090 microg g(-1) dry soil) and P (213 microg g(-1) dry soil) than forest sites. The soil moisture content, N-mineralization, nitrification rates and numbers of ammonium- and nitrite-oxidizing bacteria were highest in the wet season and lowest in dry season, while the size of mineral-N (NH4(+)-N and NO3(-)-N) showed a reverse trend at the sites. The N-mineralization, nitrification and nitrifier population size differ significantly across the site and season. The numbers of free-living cells of ammonium- and nitrite-oxidizing bacteria were significantly related to each other and to N-mineralization, nitrification, soil moisture and mineral-N components. The N-mineralization, nitrification and the viable number of nitrifying cells were consistently higher for forest soils compared to savanna sites. It was concluded that soil microbial process (N-mineralization and nitrification) and nitrifier population size were dependent on site topography, vegetation cover and soil moisture status.     

Genetic diversity of the rain tree (Albizia saman) in Colombian seasonally dry tropical forest for informing conservation and restoration interventions

Albizia saman is a multipurpose tree species of seasonally dry tropical forests (SDTFs) of Mesoamerica and northern South America typically cultivated in silvopastoral and other agroforestry systems around the world, a trend that is bound to increase in light of multimillion hectare commitments for forest and landscape restoration. The effective conservation and sustainable use of A. saman requires detailed knowledge of its genetic diversity across its native distribution range of which surprisingly little is known to date. We assessed the genetic diversity and structure of A.saman across twelve representative locations of SDTF in Colombia, and how they may have been shaped by past climatic changes and human influence. We found four different genetic groups which may be the result of differentiation due to isolation of populations in preglacial times. The current distribution and mixture of genetic groups across STDF fragments we observed might be the result of range expansion of SDTFs during the last glacial period followed by range contraction during the Holocene and human‐influenced movement of germplasm associated with cattle ranching. Despite the fragmented state of the presumed natural A. saman stands we sampled, we did not find any signs of inbreeding, suggesting that gene flow is not jeopardized in humanized landscapes. However, further research is needed to assess potential deleterious effects of fragmentation on progeny. Climate change is not expected to seriously threaten the in situ persistence of A. saman populations and might present opportunities for future range expansion. However, the sourcing of germplasm for tree planting activities needs to be aligned with the genetic affinity of reference populations across the distribution of Colombian SDTFs. We identify priority source populations for in situ conservation based on their high genetic diversity, lack or limited signs of admixture, and/or genetic uniqueness.     

Holocene southward expansion in seasonally dry tropical forests in South America: phylogeography of Ficus bonijesulapensis (Moraceae)

We conducted a phylogeographical and niche modelling study of the tree Ficus bonijesulapensis, endemic to Brazilian seasonally dry tropical forests (SDTFs), in order to evaluate the effects of Quaternary climatic fluctuations on population dynamics. The trnQ–5′rps16 region of plastid DNA was sequenced from 15 populations. Three phylogeographical groups were identified by the median‐joining algorithm network and spatial analysis of molecular variance (SAMOVA) (F_CT = 0.591): a central‐west, a central‐east and a scattered group. The central groups had higher total haplotype and nucleotide diversities than the scattered group. Ecological niche modelling suggested that, since the Last Interglacial (130 kyr bp ), the central and north regions have been relatively stable, whereas the southern region of the species distribution has been less stable. The phylogeographical groups showed concordance with the floristic units described for SDTFs. The low genetic diversity, unimodal mismatch distribution and unfavourable climatic conditions in the southern region suggest a recent southward expansion of the range of the species during the Holocene, supporting the hypothesis of the southward expansion of SDTFs during this period. The central and northern regions of the current distribution of F. bonijesulapensis, which are consistent with arboreal caatinga and rock outcrop floristic units, were potential refugia during Quaternary climatic fluctuations. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 189–201.     

Insights into the historical construction of species-rich Mesoamerican seasonally dry tropical forests: the diversification of Bursera (Burseraceae, Sapindales)

? Mesoamerican arid biomes epitomize neotropical rich and complex biodiversity. To document some of the macroevolutionary processes underlying the vast species richness of Mesoamerican seasonally dry tropical forests (SDTFs), and to evaluate specific predictions about the age, geographical structure and niche conservatism of SDTF-centered woody plant lineages, the diversification of Bursera is reconstructed. ? Using a nearly complete Bursera species-level phylogeny from nuclear and plastid genomic markers, we estimate divergence times, test for phylogenetic and temporal diversification heterogeneity, test for geographical structure, and reconstruct habitat shifts. ? Bursera became differentiated in the earliest Eocene, but diversified during independent early Miocene consecutive radiations that took place in SDTFs. The late Miocene average age of Bursera species, the presence of phylogenetic geographical structure, and its strong conservatism to SDTFs conform to expectations derived from South American SDTF-centered lineages. ? The diversification of Bursera suggests that Mesoamerican SDTF richness derives from high speciation from the Miocene onwards uncoupled from habitat shifts, during a period of enhanced aridity resulting mainly from global cooling and regional rain shadows.     

A global overview of the conservation status of tropical dry forests

Aim To analyse the conservation status of tropical dry forests at the global scale, by combining a newly developed global distribution map with spatial data describing different threats, and to identify the relative exposure of different forest areas to such threats. Location Global assessment. Methods We present a new global distribution map of tropical dry forest derived from the recently developed MODIS Vegetation Continuous Fields (VCF) product, which depicts percentage tree cover at a resolution of 500 m, combined with previously defined maps of biomes. This distribution map was overlaid with spatial data to estimate the exposure of tropical dry forests to a number of different threats: climate change, habitat fragmentation, fire, human population density and conversion to cropland. The extent of tropical dry forest currently protected was estimated by overlaying the forest map with a global data set of the distribution of protected areas. Results It is estimated that 1,048,700 km² of tropical dry forest remains, distributed throughout the three tropical regions. More than half of the forest area (54.2%) is located within South America, the remaining area being almost equally divided between North and Central America, Africa and Eurasia, with a relatively small proportion (3.8%) occurring within Australasia and Southeast Asia. Overall, c. 97% of the remaining area of tropical dry forest is at risk from one or more of the threats considered, with highest percentages recorded for Eurasia. The relative exposure to different threats differed between regions: while climate change is relatively significant in the Americas, habitat fragmentation and fire affect a higher proportion of African forests, whereas agricultural conversion and human population density are most influential in Eurasia. Evidence suggests that c. 300,000 km² of tropical dry forest now coincide with some form of protected area, with 71.8% of this total being located within South America. Main conclusions Virtually all of the tropical dry forests that remain are currently exposed to a variety of different threats, largely resulting from human activity. Taking their high biodiversity value into consideration, this indicates that tropical dry forests should be accorded high conservation priority. The results presented here could be used to identify which forest areas should be accorded highest priority for conservation action. In particular, the expansion of the global protected area network, particularly in Mesoamerica, should be given urgent consideration.     

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.     

Bird-keeping in the Caatinga,NE Brazil

Brazil has one of the greatest bird diversities in the world, with around 1,800 species. However, many species are now threatened by illegal capture and trade that is largely driven by the culture of keeping wild birds as pets. This study documents the nature and purpose of bird capture and trade in the semi-arid Caatinga biome in northeastern Brazil, which has around 510 bird species and a population of around 25 million people. Data were obtained through observation in the homes of bird keepers and at open air markets, as well as through semi-structured and open-ended interviews and informal conversations with 52 wild bird keepers. We recorded 38 species of birds kept as pets in this study. The family with the largest number of species recorded was the Emberizidae (29%), followed by the Columbidae (16%) and the Psittacidae (16%). The most frequently kept species were: Paroaria dominicana, Aratinga cactorum, Icterus jamacaii, Sporophila albogularis, Turdus rufiventris and Cyanocompsa brissonii, which are all native to this region. Among the recorded species, only Pintassilgo (Carduelis yarrellii) is listed as an endangered species in Brazil. Of the people interviewed, 32% observed a reduction in abundance, or disappearance, of several bird species in the region. The need for measures aimed at the sustainability of wild avifauna use is therefore evident. These include law enforcement, commercial breeding and environmental education, of which the last is considered the most likely to be effective.     

Plant biodiversity inventory and conservation of two tropical dry evergreen forests on the Coromandel coast, south India

Species diversity, population structure, abundance and dispersion patterns of all woody plants 10cm gbh were inventoried in two 1-ha plots of tropical dry evergreen (sacred grove or temple) forests at Kuzhanthaikuppam (KK) and Thirumanikkuzhi (TM) on the Coromandel coast of south India. Site KK is a stunted forest (average tree height ca 6 m) and TM a tall forest (average tree height ca 10 m). A total of 54 species (in 47 genera and 31 families) were recorded. Species richness and stand density were 42 and 38 species and 1367 and 974 individuals ha–1 respectively for the sites KK and TM. About 50% of the total species were common to both the sites. Site TM is twofold more voluminous (basal area 29.48 m2 ha–1) than KK (basal area 15.44 m2 ha–1). Nearly one third of the individuals are multi-stemmed in the low-statured site KK whereas one fourth of the tree density is multi-stemmed in TM. Species abundance pattern varied between the two sites. The abundance of three species in KK and two species in TM is pronounced. Memecylon umbellatum, the most abundant species contributing to one third of total stand density in KK, is least represented in TM. Species richness, density and diversity indices decreased with increasing girth threshold. Most species exhibited clumped dispersion of individuals both at 0.25 and 1-ha scales. Population structure for girth frequency is an expanding one for both the sites, except for basal area distribution in KK. Variations in plant diversity and abundance are related to site attributes and human impacts. In the light of habitat uniqueness, species richness and sacred grove status, the need for conservation is emphasized.     

Hydraulic and photosynthetic co-ordination in seasonally dry tropical forest trees

In the present study the linkage between hydraulic, photosynthetic and phenological properties of tropical dry forest trees were investigated. Seasonal patterns of stem‐specific conductivity (K_SP) described from 12 species, including deciduous, brevi‐deciduous and evergreen species, indicated that only evergreen species were consistent in their response to a dry‐to‐wet season transition. In contrast, K_SP in deciduous and brevi‐deciduous species encompassed a range of responses, from an insignificant increase in K_SP following rains in some species, to a nine‐fold increase in others. Amongst deciduous species, the minimum K_SP during the dry season ranged from 6 to 56% of wet season K_SP, indicating in the latter case that a significant portion of the xylem remained functional during the dry season. In all species and all seasons, leaf‐specific stem conductivity (K_L) was strongly related to the photosynthetic capacity of the supported foliage, although leaf photosynthesis became saturated in species with high K_L. The strength of this correlation was surprising given that much of the whole‐plant resistance appears to be in the leaves. Hydraulic capacity, defined as the product of K_L and the soil–leaf water potential difference, was strongly correlated with the photosynthetic rate of foliage in the dry season, but only weakly correlated in the wet season.     

The environmental filtering and the conservation of tropical dry forests in mountains in a global change scenario

Tropical dry forests (TDFs) are ecosystems that present stresses for woody species, and these stresses are expected to filter out many lineages that do not adapt to these stressful environments. In most cases, in the Andes, only small fragments of these forests remain with different conservation conditions in a gradient of altitude and stresses. This work aimed to answer whether taxonomic and phylogenetic diversities increase or decrease with increasing altitude in the Andes TDFs. The working hypothesis was that the variation in altitude determines the taxonomic and phylogenetic diversity of the Andes TDFs. The study was carried out through a floristic census in seven different locations in an Andean slope with altitudes ranging from 282 up to 799 m ASL. In each location, 20 plots of 25-m?×?4-m were established, totaling 140 plots. Taxonomic and phylogenetic diversity indices were estimated. We also used principal component analysis to describe the influence of climatic variables at the seven study sites. We found the influence of environmental filtering in the studied sites. As climatic seasonality increases and altitude decreases, taxonomic and phylogenetic diversity decrease and phylogenetic clustering increases. The major concern in a global warming scenario is that increasing the effects of high temperatures and droughts could extinct species and phylogenetic lineages as TDF fragmentation prevents species from migrating upwards in the mountains. Therefore, to promote connectivity is a priority besides stop habitat loss of TDFs.

     

Survival and growth of dominant tree seedlings in seasonally tropical dry forests of Yucatan: site and fertilization effects

Aims Seasonally tropical dry forests of the Yucatan Peninsula are typically found in sites with nutrient-poor soils because of the recent geological origin of the region. The landscape is dominated by extensive karstic plates that shape environments where vegetation regeneration through seed germination may be limited by the availability of suitable microsites. In this study, we documented the survival and growth of seedlings from three dominant tree species (Bursera simaruba, Piscidia piscipula and Lysiloma latisiliquum) in seasonally tropical dry forests in Yucatan. Specifically, we evaluated the effect of nutrient addition (N and P, separately and in combination) on seedling survival and growth across three sites with differing levels of precipitation.Methods We conducted a nutrient addition experiment, whereby we established 12 plots of dimensions 10×10 m (100 m 2) at each site, from which three plots were randomly selected to receive one of four treatments: N addition, P addition, N and P addition and no nutrient addition (controls). Prior to treatment application, in each plot, we planted 10 seedlings of each species in October 2010 and subsequently conducted surveys of plant growth and survival every 20 days from November 2010 to April 2011.Important findings Overall, nutrient addition increased seedling survival and the magnitude of this effect was similar among sites. We did not observe an additive effect of the N + P treatment on survival. Similarly, we observed a positive effect of nutrient addition on seedling growth, but this effect was contingent upon site; regarding survival, the effects of N and P on seedling growth were not additive. These results suggest that seedling recruitment and growth in the three dominant species of trees in Yucatan are limited by nutrient availability but that the magnitude of this effect, particularly on seedling growth, is specific for species and site.     

Mycorrhizal benefits on native plants of the Caatinga,a Brazilian dry tropical forest

In the terrestrial ecosystems, soil is an important component, characterized by holding high diversity of microorganisms which play a key role for productivity and vegetal composition. The group of symbionts microorganisms stands out for contributing directly to the growth and plant nutrition, and among them, the arbuscular mycorrhizal fungi form one of the oldest and well established associations. In order to increase the knowledge and contribute for further research with AMF and plants of Caatinga, in this review we compile data from previous studies on the effects of symbiosis between arbuscular mycorrhizal fungi (AMF) and plants of the Caatinga, a type of dry tropical forest found in the northeast of Brazil. These studies collected data under various experimental conditions, emphasizing fungal efficiency and host responsiveness in soils with varied fertility. From our analysis we conclude that in general the symbiotic efficiency on these plants depends on many factors, such as the plant-fungi combination, fertility and soil type. Furthermore, in leguminosae the impact of a joint inoculation with nitrogen fixing bacteria must be taken into account. Claroideoglomus etunicatum was the most tested AMF species benefiting almost all plants tested. Approximately 30 plant species were studied regarding possible benefits provided by AMF and of these only Hymenea courbaril and Aspidosperma pyrifolium did not respond to mycorrhization. Higher efficiency of the mycorrhizal symbiosis can be obtained in soils with low P levels, emphasizing the essential role of these microorganisms in the growth and survival of plant species from the Caatinga biome.     

Species delimitation in annual killifishes from the Brazilian Caatinga,the Hypsolebias flavicaudatus complex (Cyprinodontiformes: Rivulidae): implications for taxonomy and conservation

The organisms investigated are members of the Hypsolebias flavicaudatus species complex; a clade of morphologically similar annual killifishes endemic to the semi-arid savannah of north-eastern Brazil in the São Francisco River basin, which is a tropical area under intensive process of habitat loss. A taxonomic revision was conducted combining two different approaches for species delimitation – a tree-based approach using mt-DNA (cytochrome b) and a character-based approach using morphological variation (colour patterns, fin morphology, meristic and morphometric data). Nine species are recognised, of which five are new species: H. flagellatus (Costa), H. flavicaudatus (Costa & Brasil), H. gilbertobrasili sp. nov., H. guanambi (Costa & Amorim), H. janaubensis (Costa), H. nitens sp. nov., H. pterophyllus sp. nov., H. radiseriatus sp. nov. and H. sertanejo sp. nov. On the basis of 17 years of field studies, preliminary conservation status assessments indicate that two species are highly threatened with extinction, at least one is endangered and another is vulnerable, whereas the others are not endangered. This result deeply contrasts with a previous assessment 10 years ago, when the whole H. flavicaudatus complex was considered as a single species of no conservation concern. Discrepancies in the assessment of conservation among closely related species living in similar habitats are a consequence of heterogeneous human occupation and disproportionate recent development throughout the São Francisco River basin.     

Differences in climate-growth relationships between two tree species that co-occur in a seasonally dry tropical forest in Northeastern Brazil

Seasonally dry tropical forests are an important global climate regulator and represent one of main drivers of carbon sink dynamics. However, projections of climate change suggest future productivity losses and negative impacts on forest functioning. Understanding the interaction between climate variability and tree growth responses between species with different growth strategies represents a crucial challenge to forecast ecosystem functioning in the future. Here we used tree ring chronology to evaluate changes in growth and climate sensitivity of two tropical tree species that co-occur in a seasonally dry tropical forest in Brazil: Cedrela odorata and Ceiba glaziovii. Using Pearson correlations and linear regressions we explored how growth variability is correlated with local (precipitation, temperature) and global (ocean temperature and El Niño Southern Oscillation - ENSO) climatic factors. Tree growth was closely related with precipitation in C. odorata (r = 0.59) and C. glaziovii (0.24). Differences were found at monthly level, which C. odorata showing greater sensitivity in the beginning of rainy season. The South Atlantic Temperature was positively correlated with C. odorata, while ENSO was negatively correlated. Our results showed a dominant effect of precipitation on tree growth and suggest that are different growth strategies among species, which C. odorata being the most sensitive to drought and C. glaziovii more adapted with parenchyma in trunk. Therefore, C. odorata is probably more vulnerable to the deleterious effects of future climate change than C. glaziovii. Our findings highlight the importance of understanding the climate sensitivity of different seasonally dry tropical forest species, which is critical to predicting carbon dynamics in tropical regions. These also reveal that differences in sensitivity must be considered when prioritizing conservation measures for seasonally dry tropical forests.