Thresholds for adding degraded tropical forest to the conservation estate.

Logged and disturbed forests are often viewed as degraded and depauperate environments compared with primary forest. However, they are dynamic ecosystems1 that provide refugia for large amounts of biodiversity2,3, so we cannot afford to underestimate their conservation value4. Here we present empirically defined thresholds for categorizing the conservation value of logged forests, using one of the most comprehensive assessments of taxon responses to habitat degradation in any tropical forest environment. We analysed the impact of logging intensity on the individual occurrence patterns of 1,681 taxa belonging to 86 taxonomic orders and 126 functional groups in Sabah, Malaysia. Our results demonstrate the existence of two conservation-relevant thresholds. First, lightly logged forests (<29% biomass removal) retain high conservation value and a largely intact functional composition, and are therefore likely to recover their pre-logging values if allowed to undergo natural regeneration. Second, the most extreme impacts occur in heavily degraded forests with more than two-thirds (>68%) of their biomass removed, and these are likely to require more expensive measures to recover their biodiversity value. Overall, our data confirm that primary forests are irreplaceable5, but they also reinforce the message that logged forests retain considerable conservation value that should not be overlooked.

Forest Cover Change and Ecosystem Services: A Case Study of Community Forest in Mechinagar and Buddhashanti Landscape (MBL), Nepal.

The community-based forest management program has been successful in the conservation of forest cover in Nepal. We investigated forest cover change for the Mechinagar and Buddhashanti Landscape (MBL) area within the Jhapa district, Nepal, during 1990-2019 using Landsat images and GIS tools and valuated the major ecosystem services (ES) of Kalika Community-managed Forest (KCF) within the MBL landscape using the economic approach-market price method (revealed price). Land cover analysis of the MBL area indicated that over the study period, there were increases in urban/built-up areas, forest, and tea plantations, and declines in cultivated land, shrub, barren land, water body, and grassland areas. In particular, forest cover increased by 8.6% from 41.5 sq.km to 45.1 sq.km, due to the conversion of 3.9 sq.km cultivated land and 2.4 sq.km shrub into forest cover. KCF, the selected case study area for ES valuation, has been successful in providing noticeable economic benefits from provisioning and cultural services. Of the provisioning services, wood and timber make the largest contribution with an average revenue collection of Nepali Rupees (NRs) 3091.4 thousands followed by non-timber forest products (NRs 883.1 thousands) and firewood (NRs 524.3 thousands), respectively, while ecotourism-based income is also important (NRs 458.4 thousands) and is increasing in later years. As monitoring forest cover in a rapidly changing landscape and evaluating the ES of the community-managed forest cover are imperative for sustainable environmental planning and policy formulation, these research outputs are expected to be a significant benchmark for planners, policy makers, and future researchers.

Forest restoration and support for sustainable ecosystems in the Gandaki Basin, Nepal.

Restoring degraded forest is essential if we are to reduce human pressure on natural ecosystems and their biodiversity. Forests were nationalized in 1957 in Nepal and as a consequence, forest cover declined from 45% in 1964 to just 29% in 1994. However, as its response, sectoral plans and policies, particularly introduction of community-based forest management programs since the 1980s and conservation activities resulted in large scale forest cover restoration. Here, we examined the forest cover change in the Gandaki River Basin (GRB), the catchment with the largest altitudinal variation (ranging from ± 93 to 8167 m) and environmental and ecological significance. To see how forests have changed since then, we analyzed snapshots of spatiotemporal, ecological and physiographic changes in forest cover, and forest type at decadal intervals from 1996 to 2016 using Landsat 5 and 8 satellite images. We observed an overall gain in forest cover of 207 km2, from 7571 km2 (34.4% of the total area) in 1996 to 7778 km2 (35.3%) in 2016. Of the 21 forest cover types identified, the greatest forest coverage during 2016 was of Schima-Castanopsis forest (25.9%) and hill sal forests (16.4%). In terms of physiographic zones, land below 500 m (Tarai) where most people live, witnessed gradual declines in forest cover, in contrast to large increases in forests above 500 m. Historical examination of forest cover at ecological and physiographic scales helps to identify the elevation-wise distribution of forest resources, vegetation composition, ecosystem characteristics, anthropogenic pressure upon vegetation, and hence the overall influence of LULC upon the environment. These outputs will assist planners, policy makers, and researchers in their formulation of effective basin wide plans and policies to ensure the protection of basin level biodiversity and ecosystem function.

Land use/land cover change and ecosystem services in the Bagmati River Basin, Nepal.

Delivery of ecosystem services is strongly affected by changes in the land use/land cover (LULC) of an area. In this study, we analyze spatiotemporal changes in LULC of the rapidly changing Bagmati River Basin (BRB) of Nepal during 1988-2018 using Landsat satellite images. We also quantify carbon storage in different physiographic regions and LULC classes using the Integrated Valuation of Ecosystem Services and Trade-offs (InVEST) model and assess economic valuation of carbon using the benefit transfer method. According to our analysis, there were increases in urban/built-up (247.5%), barren land (109.5%), shrub land (32.4%), and declines in forest cover (- 6.2%), cultivated land (- 4.1%), waterbodies (- 30.3%), sand (- 29.2%), and grass cover (- 10.6%) during the study period. As a result of these changes in LULC, carbon storage declined from 31.4 million tons year-1 in 1988 (worth 157.0 million USD) to 30.8 million tons year-1 (154.1 million USD) in 2018 with the total loss of 2.9 million USD. The largest decline in stored carbon was observed in Tarai and Dun valleys, from 6.8 to 6.5 million tons (- 1.4 million USD) followed by Churia, from 7.8 to 7.6 million tons (- 1.1 million USD). Increases in carbon storage were observed in urban/built-up and shrub land areas and declines in cultivated land, forest, barren land, waterbodies and grass land. The results of LULC change and estimated carbon stock in BRB provides a baseline for planners and policy makers to formulate appropriate plans to sustainably manage the region's land cover and to mitigate carbon loss.

Quantifying the drivers of urban expansion in Nepal.

The Tarai region of Nepal is regarded as the food bowl of Nepal, and yet urban areas have increased in size at an average annual rate of 12% for the 30 years since 1988/1989, largely at the expense of prime agricultural land. Nepal is recognized internationally as highly sensitive to food security with 40% of its population undernourished. To aid future planning and reduce potential further loss of agricultural land and consequent increased food insecurity, we here investigated the previously unknown factors underlying this rapid urban expansion. We achieved this through analyses of land use and land cover (LULC) data, population, and climatic data, in association with focus group discussions and questionnaire surveys. We found that socioeconomic factors were perceived to have made the highest (62%) contribution to urbanization, particularly migration-led population growth and the economic opportunities offered by urban areas, followed by political factors (14.5%), physical factors (12%), and planning and policy factors (11.5%). In addition, climate and physiographic features make the area attractive for urban development along with favorable government plans and policies. Accelerated urban expansion during this period was particularly driven by mass migration due to political upheaval in the country resulting in rapid population and urban center growth. Of the total 293 urban centers in the country, the Tarai region includes 150 (51.2%) of which 77 (26.3%) are located in province 2 alone and accommodate 17.2% of Nepal's households. This increasing urbanization trend is expected to continue in the future due to current socioeconomic and demographic factors. We hope our results which show what has driven past urbanization will aid future urban planning and management of the Tarai as well as other similar regions elsewhere in the world. We also identified that such rapid urban growth is largely at the cost of populations in rural areas with rural depopulation resulting in agriculture being abandoned in some areas. Given Nepal's sensitivity to food security and lower food production, this will be an increasing problem for the future.

Simulating urban expansion in a rapidly changing landscape in eastern Tarai, Nepal.

Understanding the spatiotemporal dynamics of urbanization and predicting future growth is now essential for sustainable urban planning and policy making. This study explores future urban expansion in the rapidly growing region of eastern lowland Nepal. We used the hybrid cellular automata-Markov (CA-Markov) model, which utilizes historical land use and land cover (LULC) maps and several biophysical change driver variables to predict urban expansion for the years 2026 and 2036. Transitional area matrices were generated based on historical LULC data from 1996 to 2006, from 2006 to 2016, and from 1996 to 2016. The approach was validated by cross comparing the actual and simulated maps for 2016. Evaluation gave satisfactory values of Kno (0.89), Kstandard (0.84), and Klocation (0.89) which verifies the accuracy of the model. Hence, the CA-Markov model was utilized to simulate the LULC map for the years 2026 and 2036. The study area experienced rapid peri/urban expansion and sharp decline in area of cultivated land during 1989-2016. Built-up area increased by 110.90 km2 over a period of 27 years at the loss of 87.59 km2 cultivated land. Simulation analysis indicates that urban expansion will continue with urban cover increasing to 230 km2 (8.95%) and 318.51 km2 (12.45%) by 2026 and 2036, respectively, with corresponding declines in cultivated land to 1453.83 km2 (56.86%) and 1374.93 km2 (53.77%) for the same years. The alarming increase in urban areas coupled with loss of cultivated land will have negative implications for food security and environmental equilibrium in the region.

How Many Species of Insects and Other Terrestrial Arthropods Are There on Earth?

In the last decade, new methods of estimating global species richness have been developed and existing ones improved through the use of more appropriate statistical tools and new data. Taking the mean of most of these new estimates indicates that globally there are approximately 1.5 million, 5.5 million, and 7 million species of beetles, insects, and terrestrial arthropods, respectively. Previous estimates of 30 million species or more based on the host specificity of insects to plants now seem extremely unlikely. With 1 million insect species named, this suggests that 80% remain to be discovered and that a greater focus should be placed on less-studied taxa such as many families of Coleoptera, Diptera, and Hymenoptera and on poorly sampled parts of the world. DNA tools have revealed many new species in taxonomically intractable groups, but unbiased studies of previously well-researched insect faunas indicate that 1-2% of species may be truly cryptic.

New approaches narrow global species estimates for beetles, insects, and terrestrial arthropods.

It has been suggested that we do not know within an order of magnitude the number of all species on Earth [May RM (1988) Science 241(4872):1441-1449]. Roughly 1.5 million valid species of all organisms have been named and described [Costello MJ, Wilson S, Houlding B (2012) Syst Biol 61(5):871-883]. Given Kingdom Animalia numerically dominates this list and virtually all terrestrial vertebrates have been described, the question of how many terrestrial species exist is all but reduced to one of how many arthropod species there are. With beetles alone accounting for about 40% of all described arthropod species, the truly pertinent question is how many beetle species exist. Here we present four new and independent estimates of beetle species richness, which produce a mean estimate of 1.5 million beetle species. We argue that the surprisingly narrow range (0.9-2.1 million) of these four autonomous estimates--derived from host-specificity relationships, ratios with other taxa, plant:beetle ratios, and a completely novel body-size approach--represents a major advance in honing in on the richness of this most significant taxon, and is thus of considerable importance to the debate on how many species exist. Using analogous approaches, we also produce independent estimates for all insects, mean: 5.5 million species (range 2.6-7.8 million), and for terrestrial arthropods, mean: 6.8 million species (range 5.9-7.8 million), which suggest that estimates for the world's insects and their relatives are narrowing considerably.

Estimating global arthropod species richness: refining probabilistic models using probability bounds analysis.

A key challenge in the estimation of tropical arthropod species richness is the appropriate management of the large uncertainties associated with any model. Such uncertainties had largely been ignored until recently, when we attempted to account for uncertainty associated with model variables, using Monte Carlo analysis. This model is restricted by various assumptions. Here, we use a technique known as probability bounds analysis to assess the influence of assumptions about (1) distributional form and (2) dependencies between variables, and to construct probability bounds around the original model prediction distribution. The original Monte Carlo model yielded a median estimate of 6.1 million species, with a 90 % confidence interval of [3.6, 11.4]. Here we found that the probability bounds (p-bounds) surrounding this cumulative distribution were very broad, owing to uncertainties in distributional form and dependencies between variables. Replacing the implicit assumption of pure statistical independence between variables in the model with no dependency assumptions resulted in lower and upper p-bounds at 0.5 cumulative probability (i.e., at the median estimate) of 2.9-12.7 million. From here, replacing probability distributions with probability boxes, which represent classes of distributions, led to even wider bounds (2.4-20.0 million at 0.5 cumulative probability). Even the 100th percentile of the uppermost bound produced (i.e., the absolutely most conservative scenario) did not encompass the well-known hyper-estimate of 30 million species of tropical arthropods. This supports the lower estimates made by several authors over the last two decades.

Feeding guild structure of beetles on Australian tropical rainforest trees reflects microhabitat resource availability.

1. We tested the hypotheses that feeding guild structure of beetle assemblages changed with different arboreal microhabitats and that these differences were consistent across rainforest tree species. 2. Hand collection and beating techniques were used from the gondola of the Australian Canopy Crane to collect beetles from five microhabitats (mature leaves, flush leaves, flowers, fruit and suspended dead wood) within the rainforest canopy. A simple randomization procedure was implemented to test whether the abundances of each feeding guild on each microhabitat were different from that expected based on a null hypothesis of random distribution of individuals across microhabitats. 3. Beetles from different feeding guilds were not randomly distributed, but congregated on those microhabitats that are likely to provide the highest concentrations of their preferred food sources. Herbivorous beetles, in particular, were over-represented on flowers and flush foliage and under-represented on mature leaves and dead wood. Proportional numbers of species within each feeding guild were remarkably uniform across tree species for each microhabitat, but proportional abundances of feeding guilds were all significantly non-uniformly distributed between host tree species, regardless of microhabitat, confirming patterns previously found for arthropods in trees in temperate and tropical forests. 4. These results show that the canopy beetle community is partitioned into discrete assemblages between microhabitats and that this partitioning arises because of differences in feeding guild structure as a function of the diversity and the temporal and spatial availability of resources found on each microhabitat.

Quantifying uncertainty in estimation of tropical arthropod species richness.

There is a bewildering range of estimates for the number of arthropods on Earth. Several measures are based on extrapolation from species specialized to tropical rain forest, each using specific assumptions and justifications. These approaches have not provided any sound measure of uncertainty associated with richness estimates. We present two models that account for parameter uncertainty by replacing point estimates with probability distributions. The models predict medians of 3.7 million and 2.5 million tropical arthropod species globally, with 90% confidence intervals of [2.0, 7.4] million and [1.1, 5.4] million, respectively. Estimates of 30 million or greater are predicted to have <0.00001 probability. Sensitivity analyses identified uncertainty in the proportion of canopy arthropod species that are beetles as the most influential parameter, although uncertainties associated with three other parameters were also important. Using the median estimates suggests that in spite of 250 years of taxonomy and around 855,000 species of arthropods already described, approximately 70% await description.

How do herbivorous insects respond to drought stress in trees?

Increased frequency and severity of drought, as a result of climate change, is expected to drive critical changes in plant-insect interactions that may elevate rates of tree mortality. The mechanisms that link water stress in plants to insect performance are not well understood. Here, we build on previous reviews and develop a framework that incorporates the severity and longevity of drought and captures the plant physiological adjustments that follow moderate and severe drought. Using this framework, we investigate in greater depth how insect performance responds to increasing drought severity for: (i) different feeding guilds; (ii) flush feeders and senescence feeders; (iii) specialist and generalist insect herbivores; and (iv) temperate versus tropical forest communities. We outline how intermittent and moderate drought can result in increases of carbon-based and nitrogen-based chemical defences, whereas long and severe drought events can result in decreases in plant secondary defence compounds. We predict that different herbivore feeding guilds will show different but predictable responses to drought events, with most feeding guilds being negatively affected by water stress, with the exception of wood borers and bark beetles during severe drought and sap-sucking insects and leaf miners during moderate and intermittent drought. Time of feeding and host specificity are important considerations. Some insects, regardless of feeding guild, prefer to feed on younger tissues from leaf flush, whereas others are adapted to feed on senescing tissues of severely stressed trees. We argue that moderate water stress could benefit specialist insect herbivores, while generalists might prefer severe drought conditions. Current evidence suggests that insect outbreaks are shorter and more spatially restricted in tropical than in temperate forests. We suggest that future research on the impact of drought on insect communities should include (i) assessing how drought-induced changes in various plant traits, such as secondary compound concentrations and leaf water potential, affect herbivores; (ii) food web implications for other insects and those that feed on them; and (iii) interactions between the effects on insects of increasing drought and other forms of environmental change including rising temperatures and CO2 levels. There is a need for larger, temperate and tropical forest-scale drought experiments to look at herbivorous insect responses and their role in tree death.

The potential for species conservation in tropical secondary forests.

In the wake of widespread loss of old-growth forests throughout the tropics, secondary forests will likely play a growing role in the conservation of forest biodiversity. We considered a complex hierarchy of factors that interact in space and time to determine the conservation potential of tropical secondary forests. Beyond the characteristics of local forest patches, spatial and temporal landscape dynamics influence the establishment, species composition, and persistence of secondary forests. Prospects for conservation of old-growth species in secondary forests are maximized in regions where the ratio of secondary to old-growth forest area is relatively low, older secondary forests have persisted, anthropogenic disturbance after abandonment is relatively low, seed-dispersing fauna are present, and old-growth forests are close to abandoned sites. The conservation value of a secondary forest is expected to increase over time, as species arriving from remaining old-growth forest patches accumulate. Many studies are poorly replicated, which limits robust assessments of the number and abundance of old-growth species present in secondary forests. Older secondary forests are not often studied and few long-term studies are conducted in secondary forests. Available data indicate that both old-growth and second-growth forests are important to the persistence of forest species in tropical, human-modified landscapes.

Vulnerability and resilience of tropical forest species to land-use change.

We provide a cross-taxon and historical analysis of what makes tropical forest species vulnerable to extinction. Several traits have been important for species survival in the recent and distant geological past, including seed dormancy and vegetative growth in plants, small body size in mammals, and vagility in insects. For major past catastrophes, such as the five mass extinction events, large range size and vagility or dispersal were key to species survival. Traits that make some species more vulnerable to extinction are consistent across time scales. Terrestrial organisms, particularly animals, are more extinction prone than marine organisms. Plants that persist through dramatic changes often reproduce vegetatively and possess mechanisms of die back. Synergistic interactions between current anthropogenic threats, such as logging, fire, hunting, pests and diseases, and climate change are frequent. Rising temperatures threaten all organisms, perhaps particularly tropical organisms adapted to small temperature ranges and isolated by distance from suitable future climates. Mutualist species and trophic specialists may also be more threatened because of such range-shift gaps. Phylogenetically specialized groups may be collectively more prone to extinction than generalists. Characterization of tropical forest species' vulnerability to anthropogenic change is constrained by complex interactions among threats and by both taxonomic and ecological impediments, including gross undersampling of biotas and poor understanding of the spatial patterns of taxa at all scales.

How do beetle assemblages respond to cyclonic disturbance of a fragmented tropical rainforest landscape?

There are surprisingly few studies documenting effects of tropical cyclones (including hurricanes and typhoons) on rainforest animals, and especially insects, considering that many tropical forests are frequently affected by cyclonic disturbance. Consequently, we sampled a beetle assemblage inhabiting 18 upland rainforest sites in a fragmented landscape in north-eastern Queensland, Australia, using a standardised sampling protocol in 2002 and again 12 months after the passage of Severe Tropical Cyclone Larry (March 2006). The spatial configuration of sites allowed us to test if the effects of a cyclone and those from fragmentation interact. From all insect samples we extracted 12,568 beetles of 382 species from ten families. Beetle species composition was significantly different pre-and post-cyclone although the magnitude of faunal change was not large with 205 species, representing 96% of all individuals, present in both sampling events. Sites with the greatest changes to structure had the greatest changes in species composition. At the site level, increases in woody debris and wood-feeding beetle (Scolytinae) counts were significantly correlated but changes in the percent of ground vegetation were not mirrored by changes in the abundance of foliage-feeding beetles (Chrysomelidae). The overall direction of beetle assemblage change was consistent with increasing aridity, presumably caused by the loss of canopy cover. Sites with the greatest canopy loss had the strongest changes in the proportion of species previously identified in the pre-cyclone study as preferring arid or moist rainforest environments. The magnitude of fragmentation effects was virtually unaltered by the passage of Cyclone Larry. We postulate that in the short-term the effects of cyclonic disturbance and forest fragmentation both reduce the extent of moist, interior habitat.

Lianas as a food resource for herbivorous insects: a comparison with trees.

Woody climbers or, 'lianas', are one of the features that characterise rainforests. They contribute substantially to plant diversity and leaf biomass which makes them a potentially important food source for herbivores. Here, we focus on insect herbivores, folivores in particular, to show how disparities in the quantitative and qualitative availability of leaves between lianas and trees may differentially influence insect folivory and the herbivore communities themselves. We develop a conceptual model and show that lianas in general have lower structural and chemical defences, a greater nutritional profile and a preferable phenology in comparison with trees, which, contrary to our expectations, has led to assemblages of more-specialised insects. The impacts this has on higher trophic levels and broader ecological networks, however, are poorly known. We show through a study of four tropical floras from different biogeographic realms that lianas are likely to be a target for a wide range of insect herbivore taxa as they are a phylogenetically diverse group and increase diversity of higher taxa at local scales. This, in combination with their highly palatable leaves, may also make them a suitable temporary food source for insects during times when preferred host plants are scarce. This phenomenon has been observed in mammalian herbivores but awaits investigation in insects as does the effects this may have on survival and fitness. Apparent recent increases in liana abundances in some forests, likely due to climate change, makes understanding their role in supporting and maintaining biodiversity an increasingly important and necessary challenge. Since trees or saplings have usually been the subject of studies on insect herbivory, major knowledge gaps remain about the ways in which lianas contribute to, support and maintain the ecosystems in which they exist. We use our conceptual model to guide future research directions and express the necessity for caution when extrapolating explanations of herbivory derived from data on trees to growth forms with fundamentally different ecologies.

Edge effects and beta diversity in ground and canopy beetle communities of fragmented subtropical forest.

Clearing of dry forests globally creates edges between remnant forest and open anthropogenic habitats. We used flight intercept traps to evaluate how forest beetle communities are influenced by distance from such edges, together with vertical height, spatial location, and local vegetation structure, in an urbanising region (Brisbane, Australia). Species composition (but not total abundance or richness) differed greatly between ground and canopy. Species composition also varied strongly among sites at both ground and canopy levels, but almost all other significant effects occurred only at ground level, where: species richness declined from edge to interior; composition differed between positions near edges (<10 m) and interiors (> 50 m); high local canopy cover was associated with greater total abundance and richness and differing composition; and greater distances to the city centre were associated with increased total abundances and altered composition. Analyses of individual indicator species associated with this variation enabled further biological interpretations. A global literature synthesis showed that most spatially well-replicated studies of edge effects on ground-level beetles within forest fragments have likewise found that positions within tens of metres from edges with open anthropogenic habitats had increased species richness and different compositions from forest interior sites, with fewer effects on abundance. Accordingly, negative edge effects will not prevent relatively small compact fragments (if >10-20 ha) from supporting forest-like beetle communities, although indirect consequences of habitat degradation remain a threat. Retention of multiple spatially scattered forest areas will also be important in conserving forest-dependent beetles, given high levels of between-site diversity.

Conserving herbivorous and predatory insects in urban green spaces.

Insects are key components of urban ecological networks and are greatly impacted by anthropogenic activities. Yet, few studies have examined how insect functional groups respond to changes to urban vegetation associated with different management actions. We investigated the response of herbivorous and predatory heteropteran bugs to differences in vegetation structure and diversity in golf courses, gardens and parks. We assessed how the species richness of these groups varied amongst green space types, and the effect of vegetation volume and plant diversity on trophic- and species-specific occupancy. We found that golf courses sustain higher species richness of herbivores and predators than parks and gardens. At the trophic- and species-specific levels, herbivores and predators show strong positive responses to vegetation volume. The effect of plant diversity, however, is distinctly species-specific, with species showing both positive and negative responses. Our findings further suggest that high occupancy of bugs is obtained in green spaces with specific combinations of vegetation structure and diversity. The challenge for managers is to boost green space conservation value through actions promoting synergistic combinations of vegetation structure and diversity. Tackling this conservation challenge could provide enormous benefits for other elements of urban ecological networks and people that live in cities.