Opportunities to conduct large-scale field experiments are rare, but provide a unique opportunity to reveal the complex processes that operate within natural ecosystems. Here, we review the design of existing, large-scale forest fragmentation experiments. Based on this review, we develop a design for the Stability of Altered Forest Ecosystems (SAFE) Project, a new forest fragmentation experiment to be located in the lowland tropical forests of Borneo (Sabah, Malaysia). The SAFE Project represents an advance on existing experiments in that it: (i) allows discrimination of the effects of landscape-level forest cover from patch-level processes; (ii) is designed to facilitate the unification of a wide range of data types on ecological patterns and processes that operate over a wide range of spatial scales; (iii) has greater replication than existing experiments; (iv) incorporates an experimental manipulation of riparian corridors; and (v) embeds the experimentally fragmented landscape within a wider gradient of land-use intensity than do existing projects. The SAFE Project represents an opportunity for ecologists across disciplines to participate in a large initiative designed to generate a broad understanding of the ecological impacts of tropical forest modification.
In an earlier special issue of this journal, Marsh & Greer summarized forest land use in Sabah at that time and gave an introduction to the Danum Valley Conservation Area. Since that assessment, during the period 1990-2010, the forests of Sabah and particularly those of the ca 10 000 km(2) concession managed on behalf of the State by Yayasan Sabah (the Sabah Foundation) have been subject to continual, industrial harvesting, including the premature re-logging of extensive tracts of previously only once-logged forest and large-scale conversion of natural forests to agricultural plantations. Over the same period, however, significant areas of previously unprotected pristine forest have been formally gazetted as conservation areas, while much of the forest to the north, the south and the east of the Danum Valley Conservation Area (the Ulu Segama and Malua Forest Reserves) has been given added protection and new forest restoration initiatives have been launched. This paper analyses these forest-management and land-use changes in Sabah during the period 1990-2010, with a focus on the Yayasan Sabah Forest Management Area. Important new conservation and forest restoration and rehabilitation initiatives within its borders are given particular emphasis.
Global consciousness (GC), encompassing cosmopolitan orientation, global orientations (i.e. openness to multicultural experiences) and identification with all humanity, is a relatively stable individual difference that is strongly associated with pro-environmental attitudes and behaviours, less ingroup favouritism and prejudice, and greater pandemic prevention safety behaviours. Little is known about how it is socialized in everyday life. Using stratified samples from six societies, socializing institution factors correlating positively with GC were education, white collar work (and its higher income) and religiosity. However, GC also decreased with increasing age, contradicting a 'wisdom of elders' transmission of social learning, and not replicating typical findings that general prosociality increases with age. Longitudinal findings were that empathy-building, network-enhancing elements like getting married or welcoming a new infant, increased GC the most across a three-month interval. Instrumental gains like receiving a promotion (or getting a better job) also showed positive effects. Less intuitively, death of a close-other enhanced rather than reduced GC. Perhaps this was achieved through the ritualized management of meaning where a sense of the smallness of self is associated with growth of empathy for the human condition, as a more discontinuous or opportunistic form of culture-based learning. This article is part of the theme issue 'Evolution and sustainability: gathering the strands for an Anthropocene synthesis'.
Dynamics of the Pasoh forest in Peninsular Malaysia were assessed by drawing a comparison with a forest in Panama, Central America, whose dynamics have been thoroughly described. Census plots of 50 ha were established at both sites using standard methods. Tree mortality at Pasoh over an eight-year interval was 1.46% yr(-1) for all stems > or = 10 mm diameter at breast height (dbh), and 1.48% yr(-1) for stems > or = 100 mm dbh. Comparable figures at the Barro Colorado Island site in Panama (BCI) were 2.55% and 2.03%. Growth and recruitment rates were likewise considerably higher at BCI than at Pasoh. For example, in all trees 500-700 mm in dbh, mean BCI growth over the period 1985-1995 was 6 mm yr(-1), whereas mean Pasoh growth was about 3.5 mm yr(-1). Examining growth and mortality rates for individual species showed that the difference between the forests can be attributed to a few light-demanding pioneer species at BCI, which have very high growth and mortality; Pasoh is essentially lacking this guild. The bulk of the species in the two forests are shade-tolerant and have very similar mortality, growth and recruitment. The Pasoh forest is more stable than BCI's in another way as well: few of its tree populations changed much over the eight-year census interval. In contrast, at BCI, over 10% of the species had populations increasing or decreasing at a rate of >0.05 yr(-1) compared to just 2% of the species at Pasoh). The faster species turnover at BCI can probably be attributed to severe droughts that have plagued the forest periodically over the past 30 years; Pasoh has not suffered such extreme events recently. The dearth of pioneer species at Pasoh is associated with low-nutrient soil and slow litter breakdown, but the exact mechanisms behind this association remain poorly understood.
Large areas of tropical forest now exist as remnants scattered across agricultural landscapes, and so understanding the impacts of forest fragmentation is important for biodiversity conservation. We examined species richness and nestedness among tropical forest remnants in birds (meta-analysis of published studies) and insects (field data for fruit-feeding Lepidoptera (butterflies and moths) and ants). Species-area relationships were evident in all four taxa, and avian and insect assemblages in remnants typically were nested subsets of those in larger areas. Avian carnivores and nectarivores and predatory ants were more nested than other guilds, implying that the sequential loss of species was more predictable in these groups, and that fragmentation alters the trophic organization of communities. For butterflies, the ordering of fragments to achieve maximum nestedness was by fragment area, suggesting that differences among fragments were driven mainly by extinction. In contrast for moths, maximum nestedness was achieved by ordering species by wing length; species with longer wings (implying better dispersal) were more likely to occur at all sites, including low diversity sites, suggesting that differences among fragments were driven more strongly by colonization. Although all four taxa exhibited high levels of nestedness, patterns of species turnover were also idiosyncratic, and thus even species-poor sites contributed to landscape-scale biodiversity, particularly for insects.
This paper reports measurements of land-atmosphere fluxes of sensible and latent heat, momentum, CO(2), volatile organic compounds (VOCs), NO, NO(2), N(2)O and O(3) over a 30 m high rainforest canopy and a 12 m high oil palm plantation in the same region of Sabah in Borneo between April and July 2008. The daytime maximum CO(2) flux to the two canopies differs by approximately a factor of 2, 1200 mg C m(-2) h(-1) for the oil palm and 700 mg C m(-2) h(-1) for the rainforest, with the oil palm plantation showing a substantially greater quantum efficiency. Total VOC emissions are also larger over the oil palm than over the rainforest by a factor of 3. Emissions of isoprene from the oil palm canopy represented 80 per cent of the VOC emissions and exceeded those over the rainforest in similar light and temperature conditions by on average a factor of 5. Substantial emissions of estragole (1-allyl-4-methoxybenzene) from the oil palm plantation were detected and no trace of this VOC was detected in or above the rainforest. Deposition velocities for O(3) to the rainforest were a factor of 2 larger than over oil palm. Emissions of nitrous oxide were larger from the soils of the oil palm plantation than from the soils of the rainforest by approximately 25 per cent. It is clear from the measurements that the large change in the species composition generated by replacing rainforest with oil palm leads to profound changes in the net exchange of most of the trace gases measured, and thus on the chemical composition of the boundary layer over these surfaces.
The conversion of natural forest to oil palm plantation is a major current threat to the conservation of biodiversity in South East Asia. Most animal taxa decrease in both species richness and abundance on conversion of forest to oil palm, and there is usually a severe loss of forest species. The extent of loss varies significantly across both different taxa and different microhabitats within the oil palm habitat. The principal driver of this loss in diversity is probably the biological and physical simplification of the habitat, but there is little direct evidence for this. The conservation of forest species requires the preservation of large reserves of intact forest, but we must not lose sight of the importance of conserving biodiversity and ecosystem processes within the oil palm habitat itself. We urgently need to carry out research that will establish whether maintaining diversity supports economically and ecologically important processes. There is some evidence that both landscape and local complexity can have positive impacts on biodiversity in the oil palm habitat. By intelligent manipulation of habitat complexity, it could be possible to enhance not only the number of species that can live in oil palm plantations but also their contribution to the healthy functioning of this exceptionally important and widespread landscape.
Cooperation in food acquisition is a hallmark of the human species. Given that costs and benefits of cooperation vary among production regimes and work activities, the transition from hunting-and-gathering to agriculture is likely to have reshaped the structure of cooperative subsistence networks. Hunter-gatherers often forage in groups and are generally more interdependent and experience higher short-term food acquisition risk than horticulturalists, suggesting that cooperative labour should be more widespread and frequent for hunter-gatherers. Here we compare female cooperative labour networks of Batek hunter-gatherers of Peninsular Malaysia and Tsimane forager-horticulturalists of Bolivia. We find that Batek foraging results in high daily variation in labour partnerships, facilitating frequent cooperation in diffuse networks comprised of kin and non-kin. By contrast, Tsimane horticulture involves more restricted giving and receiving of labour, confined mostly to spouses and primary or distant kin. Tsimane women also interact with few individuals in the context of hunting/fishing activities and forage mainly with spouses and primary kin. These differences give rise to camp- or village-level networks that are more modular (have more substructure when partitioned) among Tsimane horticulturalists. Our findings suggest that subsistence activities shape the formation and extent of female social networks, particularly with respect to connections with other women and non-kin. We discuss the implications of restricted female labour networks in the context of gender relations, power dynamics and the adoption of farming in humans. This article is part of the theme issue 'Cooperation among women: evolutionary and cross-cultural perspectives'.
Investigations of land management impacts on hydrology are well developed in South-East Asia, having been greatly extended by national organizations in the last two decades. Regional collaborative efforts, such as the ASEAN-US watershed programme, have helped develop skills and long-running monitoring programmes. Work in different countries is significant for particular aspects: the powerful effects of both cyclones and landsliding in Taiwan, the significance of lahars in Java, of small-scale agriculture in Thailand and plantation establishment in Malaysia. Different aid programmes have contributed specialist knowledge such as British work on reservoir sedimentation, Dutch, Swedish and British work on softwood plantations and US work in hill-tribe agriculture. Much has been achieved through individual university research projects, including PhD and MSc theses. The net result is that for most countries there is now good information on changes in the rainfall-run-off relationship due to forest disturbance or conversion, some information on the impacts on sediment delivery and erosion of hillslopes, but relatively little about the dynamics and magnitude of nutrient losses. Improvements have been made in the ability to model the consequences of forest conversion and of selective logging and exciting prospects exist for the development of better predictions of transfer of water from the hillslopes to the stream channels using techniques such as multilevel modelling. Understanding of the processes involved has advanced through the detailed monitoring made possible at permanent field stations such as that at Danum Valley, Sabah.
Much of the forest remaining in South East Asia has been selectively logged. The processes promoting species coexistence may be the key to the recovery and maintenance of diversity in these forests. One such process is the Janzen-Connell mechanism, where specialized natural enemies such as seed predators maintain diversity by inhibiting regeneration near conspecifics. In Neotropical forests, anthropogenic disturbance can disrupt the Janzen-Connell mechanism, but similar data are unavailable for South East Asia. We investigated the effects of conspecific density (two spatial scales) and distance from fruiting trees on seed and seedling survival of the canopy tree Parashorea malaanonan in unlogged and logged forests in Sabah, Malaysia. The production of mature seeds was higher in unlogged forest, perhaps because high adult densities facilitate pollination or satiate pre-dispersal predators. In both forest types, post-dispersal survival was reduced by small-scale (1 m(2)) conspecific density, but not by proximity to the nearest fruiting tree. Large-scale conspecific density (seeds per fruiting tree) reduced predation, probably by satiating predators. Higher seed production in unlogged forest, in combination with slightly higher survival, meant that recruitment was almost entirely limited to unlogged forest. Thus, while logging might not affect the Janzen-Connell mechanism at this site, it may influence the recruitment of particular species.
Long-term (21-30 years) erosional responses of rainforest terrain in the Upper Segama catchment, Sabah, to selective logging are assessed at slope, small and large catchment scales. In the 0.44 km(2) Baru catchment, slope erosion measurements over 1990-2010 and sediment fingerprinting indicate that sediment sources 21 years after logging in 1989 are mainly road-linked, including fresh landslips and gullying of scars and toe deposits of 1994-1996 landslides. Analysis and modelling of 5-15 min stream-suspended sediment and discharge data demonstrate a reduction in storm-sediment response between 1996 and 2009, but not yet to pre-logging levels. An unmixing model using bed-sediment geochemical data indicates that 49 per cent of the 216 t km(-2) a(-1) 2009 sediment yield comes from 10 per cent of its area affected by road-linked landslides. Fallout (210)Pb and (137)Cs values from a lateral bench core indicate that sedimentation rates in the 721 km(2) Upper Segama catchment less than doubled with initially highly selective, low-slope logging in the 1980s, but rose 7-13 times when steep terrain was logged in 1992-1993 and 1999-2000. The need to keep steeplands under forest is emphasized if landsliding associated with current and predicted rises in extreme rainstorm magnitude-frequency is to be reduced in scale.
The primary objective of this project was to study the life cycle and ecology of Plasmodium pitheci, a malaria parasite of the orang-utan. The field work was based on the orang-utan rehabilitation centre in the Sepilok Forest Reserve of eastern Sabah. Two visits were made to Sepilok, the first in February and March, 1972, and the second (by W.P.) in January 1974. On the first visit two species of "surrogate host" were taken to Sabah, i.e. chimpanzees and Aotus monkeys for experimental work. The arboreal habitat of the orang-utan in the dipterocarp forests of eastern Sabah is described. In the Sepilok Forest Reserve dwell gibbons and leaf-monkeys, in addition to a small population of semi-domesticated and wild, free-ranging orang-utans of various ages. Although numerous species of anopheline mosquitoes have been collected in eastern Sabah, longitudinal studies are not available. Anopheles balabacensis was caught both attracted to orang-utans and to man at Sepilok. This species which is the main vector of human malaria in the north of Borneo, is suspected also of transmitting orang-utan malaria in this part of Sabah. Repeated blood examinations have been made on a number of orang-utans in the centre since 1966 and a high prevalence of infection was recorded with Plasmodium pitheci. In 1966 10 out of 19 animals had demonstrable parasitaemia. Detailed case histories are presented to show the course of parasitaemia in several orang-utans. Infections of P. pitheci were found to run a very chronic course. During the 1972 expedition a second, previously undescribed malaria parasite of the orang-utan was discovered, and was named P. silvaticum. The new parasite was successfully transmitted both by blood inoculation and, later, by sporozoite inoculation, into splenectomized chimpanzees. Although both species of malaria parasite may cause transitory signs of illness, orang-utans in general appear to be little discomforted by the infection. The animals do however suffer from other infectious diseases such as amoebic and balantidial dysentery, and melioidosis is a serious natural hazard which may have accounted for several deaths of wild orang-utans. An unidentified, intraerythrocytic structure that appeared in the blood of one chimpanzee, which had been inoculated with blood from an orang-utan, may have contributed to its death. Detailed descriptions and illustrations of P. pitheci and P. silvaticum are given. All stages of the life cycle of P. silvaticum are known (the tissue stages having been described in the liver of a "surrogate host", the chimpanzee) but only the blood and sporogonic stages of P. pitheci have been seen. This species was not infective to a chimpanzee, although there is an earlier report of a transient infection in this host by other workers. In the blood both parasites showed a tertian periodicity. From the appearance of the tissue schizonts on the seventh day it was estimated that the complete pre-erythrocytic cycle of P. silvaticum in the chimpanzee would occupy 8 days. P...
Community resistance to, and resilience from, perturbation will determine the trajectory of recovery from disturbance. Although selective timber extraction is considered a severe disturbance, fish communities from headwater streams around Danum Valley Field Centre, Sabah, Malaysia, showed few long-term changes in species composition or abundance. However, some species showed short-term (< 18 months) absence or decrease in abundance. These observations suggested that both resistance and resilience were important in maintaining long-term fish community structure. Resistance to perturbation was tested by monitoring fish communities before and after the creation of log-debris dams, while resilience was investigated by following the time-course of recolonization following complete removal of all fish. High community resistance was generally shown although the response was site-specific, dependent on the composition of the starting community, the size of the stream and physical habitat changes. High resilience was demonstrated in all recolonization experiments with strong correlations between pre- and post-defaunation communities, although there was a significant difference between pool and riffle habitats in the time-course of recovery. These differences can be explained by the movement characteristics of the species found in the different habitats. Resilience appeared to be a more predictable characteristic of the community than resistance and the implications of this for ensuring the long-term persistence of fish in the area are discussed.
Olfaction presents a particularly interesting arena to explore abstraction in language. Like other abstract domains, such as time, odours can be difficult to conceptualize. An odour cannot be seen or held, it can be difficult to locate in space, and for most people odours are difficult to verbalize. On the other hand, odours give rise to primary sensory experiences. Every time we inhale we are using olfaction to make sense of our environment. We present new experimental data from 30 Jahai hunter-gatherers from the Malay Peninsula and 30 matched Dutch participants from the Netherlands in an odour naming experiment. Participants smelled monomolecular odorants and named odours while reaction times, odour descriptors and facial expressions were measured. We show that while Dutch speakers relied on concrete descriptors, i.e. they referred to odour sources (e.g. smells like lemon), the Jahai used abstract vocabulary to name the same odours (e.g. musty). Despite this differential linguistic categorization, analysis of facial expressions showed that the two groups, nevertheless, had the same initial emotional reactions to odours. Critically, these cross-cultural data present a challenge for how to think about abstraction in language.This article is part of the theme issue 'Varieties of abstract concepts: development, use and representation in the brain'.
Changes in species composition in two 4-ha plots of lowland dipterocarp rainforest at Danum, Sabah, were measured over ten years (1986-1996) for trees > or = 10 cm girth at breast height (gbh). Each included a lower-slope to ridge gradient. The period lay between two drought events of moderate intensity but the forest showed no large lasting responses, suggesting that its species were well adapted to this regime. Mortality and recruitment rates were not unusual in global or regional comparisons. The forest continued to aggrade from its relatively (for Sabah) low basal area in 1986 and, together with the very open upper canopy structure and an abundance of lianas, this suggests a forest in a late stage of recovery from a major disturbance, yet one continually affected by smaller recent setbacks. Mortality and recruitment rates were not related to population size in 1986, but across subplots recruitment was positively correlated with the density and basal area of small trees (10-< 50cm gbh) forming the dense understorey. Neither rate was related to topography. While species with larger mean gbh had greater relative growth rates (rgr) than smaller ones, subplot mean recruitment rates were correlated with rgr among small trees. Separating understorey species (typically the Euphorbiaceae) from the overstorey (Dipterocarpaceae) showed marked differences in change in mortality with increasing gbh: in the former it increased, in the latter it decreased. Forest processes are centred on this understorey quasi-stratum. The two replicate plots showed a high correspondence in the mortality, recruitment, population changes and growth rates of small trees for the 49 most abundant species in common to both. Overstorey species had higher rgrs than understorey ones, but both showed considerable ranges in mortality and recruitment rates. The supposed trade-off in traits, viz slower rgr, shade tolerance and lower population turnover in the understorey group versus faster potential growth rate, high light responsiveness and high turnover in the overstorey group, was only partly met, as some understorey species were also very dynamic. The forest at Danum, under such a disturbance-recovery regime, can be viewed as having a dynamic equilibrium in functional and structural terms. A second trade-off in shade-tolerance versus drought-tolerance is suggested for among the understorey species. A two-storey (or vertical component) model is proposed where the understorcy-overstorey species' ratio of small stems (currently 2:1) is maintained by a major feedback process. The understorey appears to be an important part of this forest, giving resilience against drought and protecting the overstorey saplings in the long term. This view could be valuable for understanding forest responses to climate change where drought frequency in Borneo is predicted to intensify in the coming decades.
Stable carbon isotope (δ(13)C) series were developed from analysis of sequential radial wood increments from AD 1850 to AD 2009 for four mature primary rainforest trees from the Danum and Imbak areas of Sabah, Malaysia. The aseasonal equatorial climate meant that conventional dendrochronology was not possible as the tree species investigated do not exhibit clear annual rings or dateable growth bands. Chronology was established using radiocarbon dating to model age-growth relationships and date the carbon isotopic series from which the intrinsic water-use efficiency (IWUE) was calculated. The two Eusideroxylon zwageri trees from Imbak yielded ages of their pith/central wood (±1 sigma) of 670 ± 40 and 759 ± 40 years old; the less dense Shorea johorensis and Shorea superba trees at Danum yielded ages of 240 ± 40 and 330 ± 40 years, respectively. All trees studied exhibit an increase in the IWUE since AD 1960. This reflects, in part, a response of the forest to increasing atmospheric carbon dioxide concentration. Unlike studies of some northern European trees, no clear plateau in this response was observed. A change in the IWUE implies an associated modification of the local carbon and/or hydrological cycles. To resolve these uncertainties, a shift in emphasis away from high-resolution studies towards long, well-replicated time series is proposed to develop the environmental data essential for model evaluation. Identification of old (greater than 700 years) ringless trees demonstrates their potential in assessing the impacts of climatic and atmospheric change. It also shows the scientific and applied value of a conservation policy that ensures the survival of primary forest containing particularly old trees (as in Imbak Canyon and Danum).
A synthesis is presented of sampling work conducted under a UK government-funded Darwin Initiative grant undertaken predominantly within the Danum Valley Conservation Area (DVCA), Sabah, East Malaysia. The project concerned the assemblage structure, gas physiology and landscape gas fluxes of termites in pristine and two ages of secondary, dipterocarp forest. The DVCA termite fauna is typical of the Sunda region, dominated by Termes-group soil-feeders and Nasutitermitinae. Selective logging appears to have relatively little effect on termite assemblages, although soil-feeding termites may be moderately affected by this level of disturbance. Species composition changes, but to a small extent when considered against the background level of compositional differences within the Sunda region. Physiologically the assemblage is very like others that have been studied, although there are some species that do not fit on the expected body size-metabolic rate curve. As elsewhere, soil-feeders and soil-wood interface-feeders tend to produce more methane. As with the termite assemblage characteristics, gross gas and energy fluxes do not differ significantly between logged and unlogged sites. Although gross methane fluxes are high, all the soils at DVCA were methane sinks, suggesting that methane oxidation by methanotrophic bacteria was a more important process than methane production by gut archaea. This implies that methane production by termites in South-East Asia is not contributing significantly to the observed increase in levels of methane production worldwide. Biomass density, species richness, clade complement and energy flow were much lower at DVCA than at a directly comparable site in southern Cameroon. This is probably due to the different biogeographical histories of the areas.
Relatively, little is known about the relationship between biodiversity and ecosystem functioning in forests, especially in the tropics. We describe the Sabah Biodiversity Experiment: a large-scale, long-term field study on the island of Borneo. The project aims at understanding the relationship between tree species diversity and the functioning of lowland dipterocarp rainforest during restoration following selective logging. The experiment is planned to run for several decades (from seed to adult tree), so here we focus on introducing the project and its experimental design and on assessing initial conditions and the potential for restoration of the structure and functioning of the study system, the Malua Forest Reserve. We estimate residual impacts 22 years after selective logging by comparison with an appropriate neighbouring area of primary forest in Danum Valley of similar conditions. There was no difference in the alpha or beta species diversity of transect plots in the two forest types, probably owing to the selective nature of the logging and potential effects of competitive release. However, despite equal total stem density, forest structure differed as expected with a deficit of large trees and a surfeit of saplings in selectively logged areas. These impacts on structure have the potential to influence ecosystem functioning. In particular, above-ground biomass and carbon pools in selectively logged areas were only 60 per cent of those in the primary forest even after 22 years of recovery. Our results establish the initial conditions for the Sabah Biodiversity Experiment and confirm the potential to accelerate restoration by using enrichment planting of dipterocarps to overcome recruitment limitation. What role dipterocarp diversity plays in restoration only will become clear with long-term results.
We report measurements of atmospheric composition over a tropical rainforest and over a nearby oil palm plantation in Sabah, Borneo. The primary vegetation in each of the two landscapes emits very different amounts and kinds of volatile organic compounds (VOCs), resulting in distinctive VOC fingerprints in the atmospheric boundary layer for both landscapes. VOCs over the Borneo rainforest are dominated by isoprene and its oxidation products, with a significant additional contribution from monoterpenes. Rather than consuming the main atmospheric oxidant, OH, these high concentrations of VOCs appear to maintain OH, as has been observed previously over Amazonia. The boundary-layer characteristics and mixing ratios of VOCs observed over the Borneo rainforest are different to those measured previously over Amazonia. Compared with the Bornean rainforest, air over the oil palm plantation contains much more isoprene, monoterpenes are relatively less important, and the flower scent, estragole, is prominent. Concentrations of nitrogen oxides are greater above the agro-industrial oil palm landscape than over the rainforest, and this leads to changes in some secondary pollutant mixing ratios (but not, currently, differences in ozone). Secondary organic aerosol over both landscapes shows a significant contribution from isoprene. Primary biological aerosol dominates the super-micrometre aerosol over the rainforest and is likely to be sensitive to land-use change, since the fungal source of the bioaerosol is closely linked to above-ground biodiversity.