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Symbiotic nitrogen (N)-fixing trees can provide large quantities of new N to ecosystems, but only if they are sufficiently abundant. The overall abundance and latitudinal abundance distributions of N-fixing trees are well characterised in the Americas, but less well outside the Americas. Here, we characterised the abundance of N-fixing trees in a network of forest plots spanning five continents, ~5,000 tree species and ~4 million trees. The majority of the plots (86%) were in America or Asia. In addition, we examined whether the observed pattern of abundance of N-fixing trees was correlated with mean annual temperature and precipitation. Outside the tropics, N-fixing trees were consistently rare in the forest plots we examined. Within the tropics, N-fixing trees were abundant in American but not Asian forest plots (~7% versus ~1% of basal area and stems). This disparity was not explained by mean annual temperature or precipitation. Our finding of low N-fixing tree abundance in the Asian tropics casts some doubt on recent high estimates of N fixation rates in this region, which do not account for disparities in N-fixing tree abundance between the Asian and American tropics. Synthesis. Inputs of nitrogen to forests depend on symbiotic nitrogen fixation, which is constrained by the abundance of N-fixing trees. By analysing a large dataset of ~4 million trees, we found that N-fixing trees were consistently rare in the Asian tropics as well as across higher latitudes in Asia, America and Europe. The rarity of N-fixing trees in the Asian tropics compared with the American tropics might stem from lower intrinsic N limitation in Asian tropical forests, although direct support for any mechanism is lacking. The paucity of N-fixing trees throughout Asian forests suggests that N inputs to the Asian tropics might be lower than previously thought.

ForestGEO is a network of scientists and long-term forest dynamics plots (FDPs) spanning the Earth's major forest types. ForestGEO's mission is to advance understanding of the diversity and dynamics of forests and to strengthen global capacity for forest science research. ForestGEO is unique among forest plot networks in its large-scale plot dimensions, censusing of all stems ≥1 cm in diameter, inclusion of tropical, temperate and boreal forests, and investigation of additional biotic (e.g., arthropods) and abiotic (e.g., soils) drivers, which together provide a holistic view of forest functioning. The 71 FDPs in 27 countries include approximately 7.33 million living trees and about 12,000 species, representing 20% of the world's known tree diversity. With >1300 published papers, ForestGEO researchers have made significant contributions in two fundamental areas: species coexistence and diversity, and ecosystem functioning. Specifically, defining the major biotic and abiotic controls on the distribution and coexistence of species and functional types and on variation in species' demography has led to improved understanding of how the multiple dimensions of forest diversity are structured across space and time and how this diversity relates to the processes controlling the role of forests in the Earth system. Nevertheless, knowledge gaps remain that impede our ability to predict how forest diversity and function will respond to climate change and other stressors. Meeting these global research challenges requires major advances in standardizing taxonomy of tropical species, resolving the main drivers of forest dynamics, and integrating plot-based ground and remote sensing observations to scale up estimates of forest diversity and function, coupled with improved predictive models. However, they cannot be met without greater financial commitment to sustain the long-term research of ForestGEO and other forest plot networks, greatly expanded scientific capacity across the world's forested nations, and increased collaboration and integration among research networks and disciplines addressing forest science.

A taxonomic description of all castes of Colobopsisexplodens Laciny & Zettel, sp. n. from Borneo, Thailand, and Malaysia is provided, which serves as a model species for biological studies on “exploding ants” in Southeast Asia. The new species is a member of the Colobopsiscylindrica (COCY) group and falls into a species complex that has been repeatedly summarized under the name Colobopsissaundersi (Emery, 1889) (formerly Camponotussaundersi). The COCY species group is known under its vernacular name “exploding ants” for a unique behaviour: during territorial combat, workers of some species sacrifice themselves by rupturing their gaster and releasing sticky and irritant contents of their hypertrophied mandibular gland reservoirs to kill or repel rivals. This study includes first illustrations and morphometric characterizations of males of the COCY group: Colobopsisexplodens Laciny & Zettel, sp. n. and Colobopsisbadia (Smith, 1857). Characters of male genitalia and external morphology are compared with other selected taxa of Camponotini. Preliminary notes on the biology of C.explodens Laciny & Zettel, sp. n. are provided. To fix the species identity of the closely related C.badia, a lectotype from Singapore is designated. The following taxonomic changes within the C.saundersi complex are proposed: Colobopsissolenobia (Menozzi, 1926), syn. n. and Colobopsistrieterica (Menozzi, 1926), syn. n. are synonymized with Colobopsiscorallina Roger, 1863, a common endemic species of the Philippines. Colobopsissaginata Stitz, 1925, stat. n., hitherto a subspecies of C.badia, is raised to species level.

Since the 1970s Southeast Asian peat swamp forests have been increasingly threatened by anthropogenic disturbance. Peat swamps act as refuge for many endangered species, and they may turn into a net producer of CO2 and greatly contribute to climate change if cleared and drained. As one of the main invertebrate decomposers in the tropics, termites are likely to play a major role in peat forests. In this paper, we used a grid-based sampling plot protocol to sample termites in Brunei. We sampled termite communities in pristine and selectively logged peat swamp forests, that we compared with termite communities sampled in heath and dipterocarp forests. More precisely, we determined: (i) termite species diversity in peat swamp forests, and (ii) how termites respond to peat swamp logging. We found that species richness was the highest in the mixed dipterocarp forest. Selective logging had limited impact on species richness in peat swamp forest, suggesting that termite communities are resilient to limited amount of perturbations. Further data are needed to better understand the impact peat swamp clearance has on termite populations and their contribution to climate change.

Functional effects of ants in rainforest canopies depend on difficult to characterize ant diets. In Bornean dipterocarp forests, certain diurnal, arboreal, territorial, and ecologically dominant ‘COCY’ ant species (Colobopsis cylindrica clade) grazed epiphytic biofilms on adaxial leaf surfaces, as well as on tree trunks and branches. Microscopic examination of worker buccal pellets revealed numerous (mainly ascomycete) fungal spores, together with insect appendages and cuticle. Direct observations, video-imaging, and δ15N isotope data rule out feeding by predation, but isotopes cannot separate fungi from plant and insect exudates as principal nitrogen sources. Lipid-rich products, extracted from pellets in situ, are hypothesized sources of essential sterols. Also present in pellets were colorful mandibular gland (MG) compounds unique to this ant clade and deployed, as a derived character state, in suicidal defense of foraging territories. Mildly antimicrobial and highly adhesive MG products also occur basally in the clade and may have first evolved for roles in microbial sterilization and food-gathering and processing. Proteomic studies of YG COCY ants detected 2% proteins in hypertrophied, product-filled MG reservoirs, but SDS-PAGE qualitative analysis revealed mostly low-molecular mass proteins and peptides (8–15 kDa), too small for enzymes but consistent with membrane-binding proteins and/or antimicrobial peptides. Breakdown of chitin and chitosan in pellets may occur with enzymes derived from molting fluids in insect cuticle (proteases and chitinases) and/or fungi and bacteria. To the extent that COCY workers collect and consume pathogenic and/or beneficial phyllosphere microbes, ant effects on plants may be mediated by these activities. Abstrak Sejauh mana terdapat kesan fungsi semut di kanopi hutan hujan berasaskan kesukaran mengkaji pemakanan semut tersebut. Di dalam hutan dipterokarp Borneo spesies tertentu dari klad Camponotus (Colobopsis) cylindricus (spesies ‘COCY’) yang giat pada siang, tinggal di pokok, mempertahankan kawasan dan berekologi dominan didapati memakan lapisan nipis (biofilem) hidupan epifit pada permukaan daun dan juga batang dan dahan pokok. Pemerhatian pelet yang masih dalam mulut pekerja semut yang belum lagi tukar kepada produk yang berlipid tinggi mendedahkan banyak spora kulat (kebanyakannya ‘ascomycete’) yang tercampur dengan bahagian badan dan kutikal serangga. Pemerhatian dan data isotop δ15N menunjukkan ketiadaan predasi tetapi tidak dapat menunjukkan bahawa kulat dan cecair dari tumbuhan merupakan punca nitrogen utama. Pelet juga mengandungi kompaun glanda mandibular (MG) yang unik kepada klad semut tersebut yang digunakan oleh spesies-spesies yang tertentu dalam pertahanan kawasan pemakanan sehingga mati, iaitu satu ciri yang kemudian muncul. Produk MG yang kuat melekat mungkin muncul untuk memainkan peranan dalam pemungutan bahan makanan dan mungkin juga merendahkan kesan mikrob semasa pemerosesan makanan untuk dapat N dan/atau lipid (misalnya sterol sejati). Kajian proteomik dapat mengesani protin sebanyak 2 peratus yang terkandung di dalam takungan MG yang diisikan dengan produk, juga penganalisis kualitatif SDS-PAGE menunjukkan protin dan peptid yang bermolekular rendah (8-15 kDa) yang terlalu halus bagi enzim tetapi boleh jadi protin pengikatan membran atau peptid antimikrob. Penghancuran kitin dan kitosan dalam bahan pelet mungkin berlaku berkesan dari enzim yang berpunca dari kutikal serangga (yang mengandungi kitinas) dan/atau kulat dan bakteria. Kesan semut ke atas tumbuh-tumbuhan kemungkinan dipengaruhi aktiviti tersebut sejauh mana pekerja COCY memungut dan makan mikrob filoplan yang patogenik dan/atau berguna.

Unlike most other fungi, molds of the genus Trichoderma (Hypocreales, Ascomycota) are aggressive parasites of other fungi and efficient decomposers of plant biomass. Although nutritional shifts are common among hypocrealean fungi, there are no examples of such broad substrate versatility as that observed in Trichoderma. A phylogenomic analysis of 23 hypocrealean fungi (including nine Trichoderma spp. and the related Escovopsis weberi) revealed that the genus Trichoderma has evolved from an ancestor with limited cellulolytic capability that fed on either fungi or arthropods. The evolutionary analysis of Trichoderma genes encoding plant cell wall-degrading carbohydrate-active enzymes and auxiliary proteins (pcwdCAZome, 122 gene families) based on a gene tree / species tree reconciliation demonstrated that the formation of the genus was accompanied by an unprecedented extent of lateral gene transfer (LGT). Nearly one-half of the genes in Trichoderma pcwdCAZome (41%) were obtained via LGT from plant-associated filamentous fungi belonging to different classes of Ascomycota, while no LGT was observed from other potential donors. In addition to the ability to feed on unrelated fungi (such as Basidiomycota), we also showed that Trichoderma is capable of endoparasitism on a broad range of Ascomycota, including extant LGT donors. This phenomenon was not observed in E. weberi and rarely in other mycoparasitic hypocrealean fungi. Thus, our study suggests that LGT is linked to the ability of Trichoderma to parasitize taxonomically related fungi (up to adelphoparasitism in strict sense). This may have allowed primarily mycotrophic Trichoderma fungi to evolve into decomposers of plant biomass., Author summary Individual fungi rely on particular host organisms or substrates for their nutrition. Therefore, the genomes of fungi feeding on plant biomass necessarily contain genes encoding plant cell wall-degrading enzymes, while animal parasites may depend on proteolytic activity. Molds in the genus Trichoderma (Ascomycota) display a unique nutritional versatility. They can feed on other fungi, attack animals, and degrade plant debris. The later property is so efficient that one species (T. reesei) is commercially used for the production of cellulolytic enzymes required for making biofuels and other industry. In this work, we have investigated the evolution of proteins required for plant cell wall degradation in nine Trichoderma genomes and found an unprecedented number of lateral gene transfer (LGT) events for genes encoding these enzymes. Interestingly, the transfers specifically occurred from Ascomycota molds that feed on plants. We detected no cases of LGT from other fungi (e.g., mushrooms or wood-rotting fungi from Basidiomycota) that are frequent hosts of Trichoderma. Therefore, we propose that LGT may be linked to the ability of Trichoderma to parasitize on related organisms. This is a characteristic ecological trait that distinguishes Trichoderma from other mycoparasitic fungi. In this report, we demonstrate that the lateral transfer of genes may result in a profound nutritional expansion and contribute to the emergence of a generalist capable of feeding on organic matter of any origin.

A supra-annual, community-level synchronous flowering prevails in several parts of the tropical forests of Southeast Asia and its evolution has been hypothesized to be linked to pollinator shifts. The aseasonal Southeast Asian lowland rainforests are dominated by Dipterocarpaceae, which exhibit great floral diversity, a range of pollination syndromes and include species with annual and supra-annual gregarious flowering. Phylogenetic relationships within this family are still unclear, especially in the tribe Shoreeae. Here, we develop a pipeline to maximize recovery of genome-wide SNPs from restriction-site associated DNA sequencing (RADseq) in non-model organisms across wide phylogenetic scales. We then infer phylogenomic relationships in the tribe Shoreeae using both traditional and coalescent analyses. The phylogenetic trees obtained with these methods are congruent to each other and highly resolved. They allow reconstructing the evolutionary patterns of floral traits (number of stamens, anther structure and anther/appendage size) in the group. Our inferences indicate that species with many stamens, but smaller, globose anthers and longer appendages and have evolved multiple times from species with fewer stamens, but larger, oblong anthers and shorter appendages. This could have happened in parallel to iterative shifts in pollinators across the uncovered phylogeny from larger, longer generation to smaller, shorter-generation insects that can quickly build up the necessary population sizes during mass flowering episodes.

Tropical forests play a critical role in mitigating climate change, and therefore, an accurate and precise estimation of tropical forest carbon (C) is needed. However, there are many uncertainties associated with C stock estimation in a tropical forest, mainly due to its large variations in biomass. Hence, we quantified C stocks in an intact lowland mixed dipterocarp forest (MDF) in Brunei, and investigated variations in biomass and topography. Tree, deadwood, and soil C stocks were estimated by using the allometric equation method, the line intersect method, and the sampling method, respectively. Understory vegetation and litter were also sampled. We then analyzed spatial variations in tree and deadwood biomass in relation to topography. The total C stock was 321.4 Mg C ha -1 , and living biomass, dead organic matter, and soil C stocks accounted for 67%, 11%, and 23%, respectively, of the total. The results reveal that there was a relatively high C stock, even compared to other tropical forests, and that there was no significant relationship between biomass and topography. Our results provide useful reference data and a greater understanding of biomass variations in lowland MDFs, which could be used for greenhouse gas emission-reduction projects.

Summary: Tropical forest above-ground wood production (AGWP) varies substantially along environmental gradients. Some evidence suggests that AGWP may vary between regions and specifically that Asian forests have particularly high AGWP. However, comparisons across biogeographic regions using standardized methods are lacking, limiting our assessment of pan-tropical variation in AGWP and potential causes. We sampled AGWP in NW Amazon (17 long-term forest plots) and N Borneo (11 plots), both with abundant year-round precipitation. Within each region, forests growing on a broad range of edaphic conditions were sampled using standardized soil and forest measurement techniques. Plot-level AGWP was 49% greater in Borneo than in Amazonia (9.73 ± 0.56 vs. 6.53 ± 0.34 Mg dry mass ha -1 a -1 , respectively; regional mean ± 1 SE). AGWP was positively associated with soil fertility (PCA axes, sum of bases and total P). After controlling for the edaphic environment, AGWP remained significantly higher in Bornean plots. Differences in AGWP were largely attributable to differing height-diameter allometry in the two regions and the abundance of large trees in Borneo. This may be explained, in part, by the greater solar radiation in Borneo compared with NW Amazonia. Trees belonging to the dominant SE Asian family, Dipterocarpaceae, gained woody biomass faster than otherwise equivalent, neighbouring non-dipterocarps, implying that the exceptional production of Bornean forests may be driven by floristic elements. This dominant SE Asian family may partition biomass differently or be more efficient at harvesting resources and in converting them to woody biomass. Synthesis. N Bornean forests have much greater AGWP rates than those in NW Amazon when soil conditions and rainfall are controlled for. Greater resource availability and the highly productive dipterocarps may, in combination, explain why Asian forests produce wood half as fast again as comparable forests in the Amazon. Our results also suggest that taxonomic groups differ in their fundamental ability to capture carbon and that different tropical regions may therefore have different carbon uptake capacities due to biogeographic history. North Bornean forests have much greater AGWP rates than those in north-western Amazon when soil conditions and rainfall are controlled for. Greater resource availability and the highly productive dipterocarps may, in combination, explain why these Asian forests produce wood half as fast again as comparable forests in the Amazon. Our results also suggest that taxonomic groups differ in their fundamental ability to capture carbon and that different tropical regions may therefore have different carbon uptake capacities due to biogeographic history.

Plant habitat associations are well documented in Bornean lowland tropical forests, but few studies contrast the prevalence of associations across sites. We examined habitat associations and community composition of Dipterocarpaceae trees in two contrasting Bornean lowland mixed dipterocarp forests separated by approximately 100 km: Andulau (uniform topography, lower altitudinal range, sandy soils) and Belalong (highly dissected topography, higher altitudinal range, clay-rich soils). Dipterocarpaceae trees ≥ 1 cm diameter at breast height (dbh) were censused in 20-m wide belt transects established along topographic gradients at each site. Dipterocarp density, evenness, species richness, and diversity were significantly higher at Andulau than Belalong. Significant site associations (with either Andulau or Belalong) were detected for 19 (52%) of the 37 dipterocarp species tested. Dipterocarpaceae community composition at Belalong correlated with soil nutrient concentrations as well as measures of vegetation and topographic structure, but community composition at Andulau correlated with fewer habitat variables. Within each site, dipterocarp density, species richness, and diversity were consistently higher on ridges than in slopes and valleys. Significant within-site associations to topographic habitats were less common at Andulau (10% of species tested) than at Belalong (15%). We conclude that edaphic and other environmental factors influence dipterocarp community composition at a local scale, and are more important drivers of community structure in the more variable environment at Belalong. Species richness and diversity of dipterocarps on small plots, however, were higher at Andulau, suggesting that factors other than environmental heterogeneity contribute to contrasts in dipterocarp tree species richness at small scales.

Davidson, D.W., Salim, K.A. and Billen, J. 2011. Histology of structures used in territorial combat by Borneo’s ‘exploding ants’. —Acta Zoologica (Stockholm) 00:1–5. Histological studies of the hypertrophied structures of Borneo’s ‘exploding ants’ confirm homology with mandibular glands; reservoirs (only) are hypertrophied as storage structures. The general anatomical organization and ultrastructure of the glands is similar to that in other ant species. Development of secretory structures is not unusually great, and secretory cells are apparently confined to the head. These results suggest gradual production of mandibular gland compounds and filling of reservoirs and are consistent with known and hypothesized roles of mandibular glands and their products.

Research was supported by NERC through the Tropical Biomes in Transition (TROBIT) Consortium; AMAZONICA; AfriTRON and the RAINFOR network with additional support from the Gordon and Betty Moore Foundation. LB was supported by a NERC PhD Studentship and Henrietta Hutton Grant (RGS-IBG); SLL was supported by a Royal Society University Research Fellowship; some African data was collected under a NERC New Investigator Award (AfriTRON); Additional support was provided by EScFund grant of the Malaysian Ministry of Science, Technology and Innovation (MOSTI); Tropenbos International and the European Commission (project B7-6201/98-13/FOR); Large-scale Biosphere Atmosphere Experiment in Amazonia (LBA) (ND-11, CD-37, CD-41, and TG-07) under the leadership of the Brazilian Ministry of Science and Technology (MCT); PELD-CNPq and PROCAD-CAPES; and, the Brazilian National Council for Scientific and Technological Development (CNPq). Tropical tree height-diameter (H:D) relationships may vary by forest type and region making large-scale estimates of above-ground biomass subject to bias if they ignore these differences in stem allometry. We have therefore developed a new global tropical forest database consisting of 39 955 concurrent H and D measurements encompassing 283 sites in 22 tropical countries. Utilising this database, our objectives were: 1. to determine if H:D relationships differ by geographic region and forest type (wet to dry forests, including zones of tension where forest and savanna overlap). 2. to ascertain if the H:D relationship is modulated by climate and/or forest structural characteristics (e.g. stand-level basal area, A). 3. to develop H:D allometric equations and evaluate biases to reduce error in future local-to-global estimates of tropical forest biomass. Annual precipitation coefficient of variation (P-V), dry season length (S-D), and mean annual air temperature (T-A) emerged as key drivers of variation in H:D relationships at the pantropical and region scales. Vegetation structure also played a role with trees in forests of a high A being, on average, taller at any given D. After the effects of environment and forest structure are taken into account, two main regional groups can be identified. Forests in Asia, Africa and the Guyana Shield all have, on average, similar H:D relationships, but with trees in the forests of much of the Amazon Basin and tropical Australia typically being shorter at any given D than their counterparts elsewhere. The region-environment-structure model with the lowest Akaike's information criterion and lowest deviation estimated stand-level H across all plots to within a median -2.7 to 0.9% of the true value. Some of the plot-to-plot variability in H:D relationships not accounted for by this model could be attributed to variations in soil physical conditions. Other things being equal, trees tend to be more slender in the absence of soil physical constraints, especially at smaller D. Pantropical and continental-level models provided less robust estimates of H, especially when the roles of climate and stand structure in modulating H:D allometry were not simultaneously taken into account. Publisher PDF

Although recent methodological advances have allowed the incorporation of rate variation in molecular dating analyses, the calibration procedure, performed mainly through fossils, remains resistant to improvements. One source of uncertainty pertains to the assignment of fossils to specific nodes in a phylogeny, especially when alternative possibilities exist that can be equally justified on morphological grounds. Here we expand on a recently developed fossil cross-validation method to evaluate whether alternative nodal assignments of multiple fossils produce calibration sets that differ in their internal consistency. We use an enlarged Crypteroniaceae-centered phylogeny of Myrtales, six fossils, and 72 combinations of calibration points, termed calibration sets, to identify (i) the fossil assignments that produce the most internally consistent calibration sets and (ii) the mean ages, derived from these calibration sets, for the split of the Southeast Asian Crypteroniaceae from their West Gondwanan sister clade (node X). We found that a correlation exists between s values, devised to measure the consistency among the calibration points of a calibration set (Near and Sanderson, 2004), and nodal distances among calibration points. By ranking all sets according to the percent deviation of s from the regression line with nodal distance, we identified the sets with the highest level of corrected calibration-set consistency. These sets generated lower standard deviations associated with the ages of node X than sets characterized by lower corrected consistency. The three calibration sets with the highest corrected consistencies produced mean age estimates for node X of 79.70, 79.14, and 78.15 My. These timeframes are most compatible with the hypothesis that the Crypteroniaceae stem lineage dispersed from Africa to the Deccan plate as it drifted northward during the Late Cretaceous.