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.

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.

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.

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.

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.

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.

Logging cuts the functional importance of invertebrates in tropical rainforest.

Invertebrates are dominant species in primary tropical rainforests, where their abundance and diversity contributes to the functioning and resilience of these globally important ecosystems. However, more than one-third of tropical forests have been logged, with dramatic impacts on rainforest biodiversity that may disrupt key ecosystem processes. We find that the contribution of invertebrates to three ecosystem processes operating at three trophic levels (litter decomposition, seed predation and removal, and invertebrate predation) is reduced by up to one-half following logging. These changes are associated with decreased abundance of key functional groups of termites, ants, beetles and earthworms, and an increase in the abundance of small mammals, amphibians and insectivorous birds in logged relative to primary forest. Our results suggest that ecosystem processes themselves have considerable resilience to logging, but the consistent decline of invertebrate functional importance is indicative of a human-induced shift in how these ecological processes operate in tropical rainforests.

Response to comments on "Can we name Earth's species before they go extinct?".

Mora et al. disputed that most species will be discovered before they go extinct, but not our main recommendations to accelerate species' discoveries. We show that our conclusions would be unaltered by discoveries of more microscopic species and reinforce our estimates of species description and extinction rates, that taxonomic effort has never been greater, and that there are 2 to 8 million species on Earth.

Insects on flowers: The unexpectedly high biodiversity of flower-visiting beetles in a tropical rainforest canopy.

Insect biodiversity peaks in tropical rainforest environments where a large but as yet unknown proportion of species are found in the canopy. While there has been a proliferation of insect biodiversity research undertaken in the rainforest canopy, most studies focus solely on insects that inhabit the foliage. In a recent paper, we examined the distribution of canopy insects across five microhabitats (mature leaves, new leaves, flowers, fruit and suspended dead wood) in an Australian tropical rainforest, showing that the density (per dry weight gram of microhabitat) of insects on flowers were ten to ten thousand times higher than on the leaves. Flowers also supported a much higher number of species than expected based on their contribution to total forest biomass. Elsewhere we show that most of these beetle species were specialized to flowers with little overlap in species composition between different canopy microhabitats. Here we expand our discussion of the implications of our results with respect to specialization and the generation of insect biodiversity in the rainforest canopy. Lastly, we identify future directions for research into the biodiversity and specialization of flower-visitors in complex tropical rainforests.

Can we name Earth's species before they go extinct?

Some people despair that most species will go extinct before they are discovered. However, such worries result from overestimates of how many species may exist, beliefs that the expertise to describe species is decreasing, and alarmist estimates of extinction rates. We argue that the number of species on Earth today is 5 ± 3 million, of which 1.5 million are named. New databases show that there are more taxonomists describing species than ever before, and their number is increasing faster than the rate of species description. Conservation efforts and species survival in secondary habitats are at least delaying extinctions. Extinction rates are, however, poorly quantified, ranging from 0.01 to 1% (at most 5%) per decade. We propose practical actions to improve taxonomic productivity and associated understanding and conservation of biodiversity.

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.

The overlooked biodiversity of flower-visiting invertebrates.

Estimates suggest that perhaps 40% of all invertebrate species are found in tropical rainforest canopies. Extrapolations of total diversity and food web analyses have been based almost exclusively on species inhabiting the foliage, under the assumption that foliage samples are representative of the entire canopy. We examined the validity of this assumption by comparing the density of invertebrates and the species richness of beetles across three canopy microhabitats (mature leaves, new leaves and flowers) on a one hectare plot in an Australian tropical rainforest. Specifically, we tested two hypotheses: 1) canopy invertebrate density and species richness are directly proportional to the amount of resource available; and 2) canopy microhabitats represent discrete resources that are utilised by their own specialised invertebrate communities. We show that flowers in the canopy support invertebrate densities that are ten to ten thousand times greater than on the nearby foliage when expressed on a per-unit resource biomass basis. Furthermore, species-level analyses of the beetle fauna revealed that flowers support a unique and remarkably rich fauna compared to foliage, with very little species overlap between microhabitats. We reject the hypothesis that the insect fauna on mature foliage is representative of the greater canopy community even though mature foliage comprises a very large proportion of canopy plant biomass. Although the significance of the evolutionary relationship between flowers and insects is well known with respect to plant reproduction, less is known about the importance of flowers as resources for tropical insects. Consequently, we suggest that this constitutes a more important piece of the 'diversity jigsaw puzzle' than has been previously recognised and could alter our understanding of the evolution of plant-herbivore interactions and food web dynamics, and provide a better foundation for accurately estimating global species richness.

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 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.

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.

Beetle assemblages from an Australian tropical rainforest show that the canopy and the ground strata contribute equally to biodiversity.

There remains great uncertainty about how much tropical forest canopies contribute to global species richness estimates and the relative specialization of insect species to vertical zones. To investigate these issues, we conducted a four-year sampling program in lowland tropical rainforest in North Queensland, Australia. Beetles were sampled using a trap that combines Malaise and flight interception trap (FIT) functions. Pairs of this trap, one on the ground and a second suspended 15-20 m above in the canopy were located at five sites, spaced 50 m or more apart. These traps produced 29986 beetles of 1473 species and 77 families. There were similar numbers of individuals (canopy 14473; ground 15513) and species (canopy 1158; ground 895) in each stratum, but significantly more rare species in the canopy (canopy 509; ground 283). Seventy two percent of the species (excluding rare species) were found in both strata. Using IndVal, we found 24 and 27% of the abundant species (n>or=20 individuals) to be specialized to the canopy and the ground strata, respectively, and equivalent analyses at the family level showed figures of 30 and 22%, respectively. These results show that the canopy and the ground strata both provide important contributions to rainforest biodiversity.