Phylogenetic community patterns suggest Central Indian tropical dry forests are structured by montane climate refuges

Aim

We used an eco-phylogenetic approach to investigate the diversity and assembly patterns of tropical dry forests (TDFs) in Central India. We aimed at informing conservation and restoration practices in these anthropogenically disturbed forests by identifying potential habitats of conservation significance and elements of regional biodiversity most vulnerable to human impact and climate change.

Location

Tropical dry forests of Madhya Pradesh, Central India.

Methods

We analysed the species richness, stem density, basal area and phylogenetic structure (standardized effect size of MNTD, MPD, PD and community evolutionary distinctiveness cED) of 117 tree species assemblages distributed across a ~230 to ~940 m elevational gradient. We examined how these community measures and taxonomic (Sørensen) and phylogenetic (UniFrac) beta diversity varied with elevation, precipitation, temperature and climatic stress.

Results

Species richness, phylogenetic diversity, stem density and basal area were positively correlated with elevation, with high-elevation plots exhibiting cooler temperatures, higher precipitation and lower stress. High-elevation assemblages also trended towards greater phylogenetic dispersion, which diminished at lower elevations and in drier, more stressful plots. Phylogenetic turnover was observed across the elevation gradient, and species evolutionary distinctiveness increased at lower elevations and under harsher abiotic conditions.

Main Conclusions

Harsher abiotic conditions at low elevations may act as a selective filter on plant lineages, leading to phylogenetically clustered low-diversity assemblages. These assemblages contained more evolutionarily distinct species that may contribute disproportionately to biodiversity. Conversely, milder abiotic conditions at high elevations may serve as refuges for drought-sensitive species, resulting in more diverse assemblages. Conservation practices that prioritize both high- and low-elevation habitats could promote the persistence of evolutionarily distinct species and areas of high biodiversity within the Central Indian landscape. Establishing connectivity between these habitats may provide a range of climatic conditions for species to retreat to or persist within as climates change.     

Phylogenetic structure of angiosperm communities during tropical forest succession

The phylogenetic structure of ecological communities can shed light on assembly processes, but the focus of phylogenetic structure research thus far has been on mature ecosystems. Here, I present the first investigation of phylogenetic community structure during succession. In a replicated chronosequence of 30 sites in northeastern Costa Rica, I found strong phylogenetic overdispersion at multiple scales: species present at local sites were a non-random assemblage, more distantly related than chance would predict. Phylogenetic overdispersion was evident when comparing the species present at each site with the regional species pool, the species pool found in each age category to the regional pool or the species present at each site to the pool of species found in sites of that age category. Comparing stem size classes within each age category, I found that during early succession, phylogenetic overdispersion is strongest in small stems. Overdispersion strengthens and spreads into larger size classes as succession proceeds, corroborating an existing model of forest succession. This study is the first evidence that succession leaves a distinct signature in the phylogenetic structure of communities.     

Phylogenetic tests of community assembly across regional to continental scales in tropical and subtropical rain forests

Aim To measure and quantify community phylogenetic structure to evaluate how evolutionary, ecological and biogeographic processes have shaped the distributions and assemblage of tropical and subtropical rain forest tree species across local, regional and continental scales. Location Australia. Methods We used 596 assemblage‐level samples and 1137 woody species in rain forest vegetation sampled across two latitude regions (tropics and sub‐tropics) and five distinct areas. Based on this dataset, we obtained and analysed species‐level trait values (for leaf size, seed size, wood density and maximum height at maturity), measures of community phylogenetic structure and species turnover across space (beta) and evolutionary time (phylobeta). Results Phylobeta values showed that at continental scales (i.e. across the latitude regions combined) species replacement, as turnover in assemblages through time, was by more phylogenetically distant (i.e. less closely related) taxa. Within latitude regions replacement was by more closely related taxa. Assemblages of species were more phylogenetically clustered across the whole phylogeny (net relatedness index) and with respect to more recent divergences (nearest related taxon index) where the effects of historic disturbance (climatic oscillations) had been greater, and less clustered in long‐term stable (refugial) locations. Local species composition in the stable wet tropics showed significant phylogenetic evenness, but there was no corresponding evenness in distributions of the ecological traits measured. Main conclusions Despite a shared evolutionary and biogeographic history, the two regions diverged from each other before the development of internal divergences. Phylogenetic evenness is more evident in long‐term stable habitats (refugia) where species interact in conserved niches. Phylogenetic clustering is more evident where recolonization of more highly disturbed areas from historically reduced species pools reflects filtering of species into phylogenetically preferred habitats.     

Phylogenetic isolation of host trees affects assembly of local Heteroptera communities

A host may be physically isolated in space and then may correspond to a geographical island, but it may also be separated from its local neighbours by hundreds of millions of years of evolutionary history, and may form in this case an evolutionarily distinct island. We test how this affects the assembly processes of the host''s colonizers, this question being until now only invoked at the scale of physically distinct islands or patches. We studied the assembly of true bugs in crowns of oaks surrounded by phylogenetically more or less closely related trees. Despite the short distances (less than 150 m) between phylogenetically isolated and non-isolated trees, we found major differences between their Heteroptera faunas. We show that phylogenetically isolated trees support smaller numbers and fewer species of Heteroptera, an increasing proportion of phytophages and a decreasing proportion of omnivores, and proportionally more non-host-specialists. These differences were not due to changes in the nutritional quality of the trees, i.e. species sorting, which we accounted for. Comparison with predictions from meta-community theories suggests that the assembly of local Heteroptera communities may be strongly driven by independent metapopulation processes at the level of the individual species. We conclude that the assembly of communities on hosts separated from their neighbours by long periods of evolutionary history is qualitatively and quantitatively different from that on hosts established surrounded by closely related trees. Potentially, the biotic selection pressure on a host might thus change with the evolutionary proximity of the surrounding hosts.     

Endemic birds of the Atlantic Forest: traits,conservation status,and patterns of biodiversity

The bird fauna of the Brazilian Atlantic Forest is exceptionally diverse and threatened, with high levels of endemism. Available lists of the endemic birds of the Atlantic Forest were generated before recent taxonomic revisions lumped or split species and before the recent increase in species occurrence records. Our objective, therefore, was to compile a new list of the endemic birds of the Atlantic Forest, characterize these species in terms of conservation status and natural history traits, and map remaining vegetation and protected areas. We combined GIS analysis with a literature search to compile a list of endemic species and, based on the phylogeny and distribution of these species, characterized areas in terms of species richness, phylogenetic diversity, and endemism. We identified 223 species of birds endemic to the Atlantic Forest, including 12 species not included in previous lists. In addition, 14 species included in previous lists were not considered endemic, either because they occur outside the Atlantic Forest biome or because they are not considered valid species. The typical Atlantic Forest endemic bird is a small forest‐dependent invertivore. Of the species on our list, 31% are considered threatened or extinct. Only ~ 34% of the spatial analysis units had > 10% forest cover, and protected area coverage was consistently low (< 1%). In addition, we found spatial incongruity among the different measures of biodiversity (species richness, relative phylogenetic diversity, restricted‐range species, and irreplaceability). Each of these measures provides information concerning different aspects of biological diversity. However, regardless of which aspect(s) of biodiversity might be considered most important, preservation of the remaining areas of remnant vegetation and further expansion of protected areas are essential if we are to conserve the many endemic species of birds in the Atlantic Forest.     

The importance of well protected forests for the conservation genetics of West African colobine monkeys

In tropical forests, anthropogenic activities are major drivers of the destruction and degradation of natural habitats, causing severe biodiversity loss. African colobine monkeys (Colobinae) are mainly folivore and strictly arboreal primates that require large forests to subsist, being among the most vulnerable of all nonhuman primates. The Western red colobus Piliocolobus badius and the King colobus Colobus polykomos inhabit highly fragmented West African forests, including the Cantanhez Forests National Park (CFNP) in Guinea-Bissau. Both species are also found in the largest and best-preserved West African forest—the Taï National Park (TNP) in Ivory Coast. Colobine monkeys are hunted for bushmeat in both protected areas, but these exhibit contrasting levels of forest fragmentation, thus offering an excellent opportunity to investigate the importance of well-preserved forests for the maintenance of evolutionary potential in these arboreal primates. We estimated genetic diversity, population structure, and demographic history by using microsatellite loci and mitochondrial DNA. We then compared the genetic patterns of the colobines from TNP with the ones previously obtained for CFNP and found contrasting genetic patterns. Contrary to the colobines from CFNP that showed very low genetic diversity and a strong population decline, the populations in TNP still maintain high levels of genetic diversity and we found no clear signal of population decrease in Western red colobus and a limited decrease in King colobus. These results suggest larger and historically more stable populations in TNP compared to CFNP. We cannot exclude the possibility that the demographic effects resulting from the recent increase of bushmeat hunting are not yet detectable in TNP using genetic data. Nevertheless, the fact that the TNP colobus populations are highly genetically diverse and maintain large effective population sizes suggests that well-preserved forests are crucial for the maintenance of populations, species, and probably for the evolutionary potential in colobines.     

Structure and function in two tropical gallery forest communities: implications for forest conservation in fragmented systems

1. Composition, growth and turnover of trees in two species-rich tropical gallery forests were examined to evaluate what community reorganization may be needed to transform recently created tropical forest fragments into stable refugia for regional forest biotas.
2. Rates of tree growth and turnover over a 5-year interval were comparable to those recorded in continuous forests and in both communities there had been some tree species turnover in the measured stem size classes during the 5-year interval.
3. The more abundant tree species in both communities formed three functional groups along gradients between streams and forest edges: edge-concentrators, core-concentrators and generalists.
4. Soil fertility showed no consistent increase close to streams and neither tree growth nor recruitment rates were increased in this zone. In contrast, forest edge zones exhibited increased rates of tree growth and recruitment indicating that growth processes in these forests are light-limited rather than soil-limited, and that forest edge zones are generally favourable habitats for tree populations.
5. Both communities showed signs of past fire incursions, and the tendency of a subset of tree species to concentrate in the more growth-limited core habitats is attributed to their fire sensitivity.
6. Rapid development of an edge zone of fire-insensitive tree species is considered to be essential to the survival of forest community fragments in the fire-prone landscapes of the tropics, and the edges of gallery forests are recommended as potential sources of species with which to fashion these protective ecotones.
7. Preservation of a diverse forest biota in the fire-protected interiors of fragments will require natural or artificially enhanced immigration rates that are sufficient to offset local extinctions.     

Phylogenetic alpha and beta diversity in tropical tree assemblages along regional-scale environmental gradients in northwest South America

Aims Environmental gradients are drivers of species diversity; however, we know relatively little about the evolutionary processes underlying these relationships. A potentially powerful approach to studying diversity gradients is to quantify the phylogenetic structure within and between assemblages arrayed along broad spatial and environmental gradients. Here, we evaluate the phylogenetic structure of plant assemblages along an environmental gradient with the expectation that the habitat specialization of entire lineages is an important evolutionary pattern influencing the structure of tree communities along environmental gradients.Methods We evaluated the effect of several environmental variables on the phylogenetic structure of plant assemblages in 145 plots distributed in northwestern South America that cover a broad environmental gradient. The phylogenetic alpha diversity was quantified for each plot and the phylogenetic beta diversity between each pair of plots was also quantified. Both the alpha and beta diversity measures were then related to spatial and environmental gradients in the study system.Important findings We found that gradients in temperature and potential evapotranspiration have a strong relationship with the phylogenetic alpha diversity in our study system, with phylogenetic overdispersion in low temperatures and phylogenetic clustering at higher temperatures. Further, the phylogenetic beta diversity between two plots increases with an increasing difference in temperature, whereas annual precipitation was not a significant predictor of community phylogenetic turnover. We also found that the phylogenetic structure of the plots in our study system was related to the degree of seasonal flooding and seasonality in precipitation. In particular, more stressful environments such as dry forests and flooded forests showed phylogenetic clustering. Finally, in contrast with previous studies, we find that phylogenetic beta diversity was not strongly related to the spatial distance separating two forest plots, which may be the result of the importance of the three independent mountain ranges in our study system, which generate a high degree of environmental variation over very short distances. In conclusion, we found that environmental gradients are important drivers of both phylogenetic alpha and phylogenetic beta diversities in these forests over spatial distance.     

Resilience of tropical rain forests: tree community reassembly in secondary forests

Understanding the recovery dynamics of ecosystems presents a major challenge in the human-impacted tropics. We tested whether secondary forests follow equilibrium or non-equilibrium dynamics by evaluating community reassembly over time, across different successional stages, and among multiple life stages. Based on long-term and static data from six 1-ha plots in NE Costa Rica, we show that secondary forests are undergoing reassembly of canopy tree and palm species composition through the successful recruitment of seedlings, saplings, and young trees of mature forest species. Such patterns were observed over time within sites and across successional stages. Floristic reassembly in secondary forests showed a clear convergence with mature forest community composition, supporting an equilibrium model. This resilience stems from three key factors co-occurring locally: high abundance of generalist species in the regional flora, high levels of seed dispersal, and local presence of old-growth forest remnants.     

Phylogenetic turnover in tropical tree communities: impact of environmental filtering,biogeography and mesoclimatic niche conservatism

Aim We addressed the roles of environmental filtering, historical biogeography and evolutionary niche conservatism on the phylogenetic structure of tropical tree communities with the following questions. (1) What is the impact of mesoclimatic gradients and dispersal limitation on phylogenetic turnover and species turnover? (2) How does phylogenetic turnover between continents compare in intensity with the turnover driven by climatic gradients at a regional scale? (3) Are independent phylogenetic reconstructions of the mesoclimatic niche of clades congruent between continents? Location Panama Canal Watershed and Western Ghats (India), two anciently divergent biogeographic contexts but with comparable rainfall gradients. Methods Using floristic data for 50 1‐ha plots in each region, independent measures of phylogenetic turnover (Π_ST) and species turnover (Jaccard) between plots were regressed on geographic and ecological distances. Mesoclimatic niches were reconstructed for each node of the phylogeny and compared between the two continents. Results (1) The phylogenetic turnover within each region is best explained by mesoclimatic differences (environmental filtering), while species turnover depends both on mesoclimatic differences and geographic distances (dispersal limitation). (2) The phylogenetic turnover between continents (Π_ST = 0.009) is comparable to that caused by mesoclimatic gradients within regions (Π_ST = 0.010) and both effects seem cumulative. (3) Independent phylogenetic reconstructions of the mesoclimatic niches were strongly correlated between the two continents (r = 0.61), despite the absence of shared species. Main conclusions Our results demonstrate a world‐wide deep phylogenetic signal for mesoclimatic niche within a biome, indicating that positive phylogenetic turnover at a regional scale reflects environmental filtering in plant communities.     

Phylogenetic and phenotypic structure among Banksia communities in south‐western Australia

Aim Phylogenetic and phenotypic patterns among coexisting banksias (Banksia, Proteaceae) in the infertile, fire‐prone landscapes of south‐western Australia were examined for evidence of community structuring. It was expected that closely related species would be spatially clustered (underdispersed) as a consequence of widespread recent speciation, strong edaphic fidelity and low dispersability. We also expected that edaphic filtering would result in phenotypic clustering of traits related to habitat specialization and that competitive exclusion among closely related species with similar regeneration biology and growth form would result in phenotypic overdispersion of these latter traits. Location Southwest Australian Floristic Region (SWAFR). Methods Based on published data for coexistence (richness and frequency) of Banksia species at 40 sites in the three floristic provinces, phylogenetic, soil type and morphological mean pairwise distance and mean nearest taxon distance were calculated for each site and compared with null communities. Patterns of co‐occurrence were examined at the local and subregional (provincial) scales. Results Of the 40 sites assessed, 21–30 displayed phylogenetic clustering of Banksia species (5–11 significantly) such that, overall, co‐occurring taxa were more closely related than expected by chance. Banksias in the Transitional Rainfall and Southeast Coastal Provinces were more likely to display phylogenetic clustering than in the High Rainfall Province. A significant trend for phylogenetic clustering associated with edaphic specialization (27–30 sites) was observed, as well as a significant trend for phenotypic overdispersion associated with growth form (25–28 sites). Results for regeneration biology depended on the metric used. Main conclusions We demonstrate spatial clustering of closely related banksias at the local and provincial scales, consistent with their restricted distribution (recent widespread speciation, patchy habitat availability and limited dispersability) in this geologically old and stable region. The clustering of closely related species may also be a consequence of habitat filtering linked to edaphic fidelity in the SWAFR flora, while overdispersion in growth form suggests that functional divergence favours coexistence in Banksia communities.     

Survival and growth of tree seedlings in anthropogenically disturbed Mexican montane rain forests

Abstract. The montane rain forest in Chiapas, Mexico has been subject to continuous human disturbance, mainly by wood extraction, cattle grazing and fire. To address the effect of this kind of disturbance regime on regeneration, survival and growth of naturally established tree seedlings were evaluated during a 3‐yr period in three forest stands which varied in frequency and intensity of human disturbance. Frequency of cattle browsing on transplanted seedlings of Cornus disciflora, Liquidambar styraciflua and Oreopanax xalapensis planted on different facing slopes was evaluated over an 18‐mo period, their survival and growth was also determined. The density of seedlings, number of species and survival of naturally recruited tree seedlings decreased with increased intensity of disturbance. Only seedlings of Pinus and Quercus species were recorded in heavily disturbed conditions. Seedlings grew more vigorously under moderately disturbed than in undisturbed or heavily disturbed conditions. More extreme microclimatic variation (air temperature, relative humidity and solar radiation) was recorded in heavily vs moderately disturbed forest. Survival and growth of transplanted tree seedlings differed with species. Grazing damage varied among seedlings located on different facing slopes. Plants located on NE facing slopes survived better than those on other slope positions. The mean proportion of surviving individuals of L. styraciflua (80%) and C. disciflora (78%) was higher than those for O. xalapensis (60%). A higher number of C. disciflora and O. xalapensis seedlings were grazed than L. styraciflua in any slope orientation treatment. The results suggest that human disturbance negatively affects natural recruitment, reduces richness of tree seedlings and favours a small number of tree species.     

Community assembly after long-term fragmentation: a case study of tropical rainforest in Xishuangbanna,south-west China

Background: Tropical rainforests represent the most species-rich and at the same time the most fragmented terrestrial biome on Earth. Fragmentation of tropical rainforests is having wide-ranging consequences for the maintenance of local species diversity and community assembly patterns.

Aims: To examine floristic changes and changes in community phylogenetic structure in the forest fragment over the past five decades.

Methods: A new taxonomic diversity algorithm (within-family diversity) was developed to assess floristic changes in the forest fragment. Community phylogenetic structure was then compared before and after fragmentation.

Results: Taxonomic diversity changed greatly among families, with changes occurring randomly across the phylogeny. The forest fragment had higher phylogenetic diversity, higher mean pair-wise phylogenetic distance, but lower mean nearest-neighbour distance. The community phylogenetic structure has changed significantly from clustering to dispersion.

Conclusions: High species turnover occurred in the forest fragment. While shade-tolerant species have been lost, and ruderal and alien species have been added, overall phylogenetic diversity has increased with species being more phylogenetically distant. Competitive exclusion, which was related to the relatively drier conditions in the forest after fragmentation, led the plant community phylogenetic structure to be more dispersed.     

Evolutionary diversity gradients in neotropical tree assemblages: New insights from Non-Flooded Evergreen forests

We investigated spatial patterns of evolutionary diversity along Neotropical Non-Flooded Evergreen Forests (NEF). We addressed the following questions: (i) What are the main NEF evolutionary groups? (ii) How evolutionary diversity varies across NEF environmental gradients? Based on a phylogeny of 1248 tree genera distributed over 1824 NEF assemblages, we examined the evolutionary differentiation using UPGMA and evopca. We measured lineage diversity (ses.PD) and structure (ses.MPD and ses.MNTD) and tested their response to environmental gradients using linear models. Phylogenetic dissimilarity segregated NEF into 12 evolutionary groups that largely confirm groups obtained in our previous work based on floristic similarity. However, one discrepancy was the amalgamation of Amazon and northern Atlantic Forest assemblages, while the southern Atlantic Forest remained an isolated group. Furthermore, Mesoamerica, which had been recognized as a single group, here split into six evolutionary groups. We found greater lineage diversity as altitude and latitude increased and temperature decreased. Evolutionary groups with the highest mean values of lineage diversity were those composed of Mesoamerican cloud forests, which harbor a mixture of tropical and temperate lineages representing a confluence of South and North American floras. We found that variations in phylogenetic diversity in NEF are primarily related to the coexistence of lineages of temperate and tropical climates in the mountain and nebular environments of NEF, indicating the strong contribution of extratropical niche conservatism in structuring evolutionary diversity.