Nigel E. Stork

Insects in fragmented forests: a functional approach

Insects are highly susceptible to the adverse effects of forest fragmentation. It is now beyond any doubt that fragmentation-induced changes in abundance and species richness occur in many insect groups. However, the study of insects in fragmented forests is still in its infancy and lacks real direction. Simple empirical studies are not answering the questions we most want to answer about fragmented systems. Are we in the midst of a mass-extinction crisis? What is the functional significance of the immense insect biodiversity? Does biodiversity loss affect ecosystem functioning? A more focused, functional approach to the study of forest fragmentation is required to move beyond the description of pattern and to determine how changes in insect communities affect ecosystem processes in fragmented forests.

BEETLE SPECIES RESPONSES TO TROPICAL FOREST FRAGMENTATION

The effects of forest fragmentation on beetle species composition were investigated in an experimentally fragmented tropical forest landscape in Central Amazonia. Leaf-litter beetles were sampled at seven distances from the forest edge (0–420 m) along forest edge-to-interior transects in two 100-ha forest fragments and two continuous forest edges, and at an identical series of distances along two deep continuous forest transects. Additional samples were taken at the centers of two 10-ha forest fragments and two 1-ha fragments. This sampling regime allowed discrimination between edge and fragment area effects. Beetle species composition changed significantly and independently with both decreasing distance from forest edge and decreasing fragment area. Edge effects on species composition were mediated by six important environmental variables: air temperature, canopy height, percent ground cover of twigs, litter biomass, litter moisture content, and an air temperature × distance from edge interaction effect, due to the different temperature profiles of edges with differing edge vegetation density. Population densities of 15 of the 32 most abundant beetle species tested (47%) were significantly affected by forest fragmentation. Species responses were classified empirically into four major categories: (A) edge sensitive, area insensitive; (B) area sensitive, edge insensitive; (C) edge and area sensitive; and (D) edge and area insensitive. Within these categories, trends in density were either positive (deep-forest species), or negative (disturbed-area species), with species showing the full spectrum of responses to fragmentation. The vast majority of species were adversely affected. Estimated species loss rates from forest fragments were: 49.8% of common species from 1-ha fragments, 29.8% from 10-ha fragments, and 13.8% from 100-ha fragments. Declining density was a significant precursor of species loss from forest fragments, but other species that did not show significant population density responses to fragmentation were also absent from some fragments, presumably by chance. The probability of species loss from forest fragments was not correlated with body size or trophic group for the 32 common species, although for the entire beetle assemblage (993 species) proportions of species in different trophic groups changed significantly with fragmentation. Rarity and population variability (in undisturbed forest) were significant predictors of susceptibility to fragmentation. Surprisingly, though, common species were significantly more likely to become locally extinct in small fragments than rarer species. This lends empirical support to models of multispecies coexistence under disturbance that suggest competitively dominant but poorly dispersing species are the first to become extinct due to habitat destruction. Thus, rarer species are predicted to be better dispersers and better at persisting.

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.

Can We Name Earth's Species Before They Go Extinct?

Completing the Catalog Despite the widely held belief that the number of taxonomists is decreasing, there is evidence that increasing numbers of authors are describing species new to science. In parallel, several statistically sophisticated attempts have been made to better quantify the number of species that may exist on Earth, including the oceans. Estimates of recent extinction rates have also been re-examined to question whether we are in, or heading toward, an anthropogenic mass extinction event. Costello et al. (p. 413) review these findings, provide hope that science will describe most species within this century, and suggest how this complete description can be facilitated. 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.

Species number, species abundance and body length relationships of arboreal beetles in Bornean lowland rain forest trees

ABSTRACT. 1 The relationships between number of species, abundance per species, and body length are examined for 859 species of beetles in samples of arthropods collected from ten Bornean lowland forest trees by insecticide fogging. Similar relationships are examined for different feeding guilds of these beetles, and for those beetles from different species of trees. 2 The data are used to construct four interrelated graphs, namely species: abundance, species: body length, population abundance: body length and total number of individuals: body length distributions. 3 In contrast to a number of previous studies, no consistent linear relationship between population density and body length was found for the Bornean beetles and it is suggested that, as in birds, the added dispersal ability of flight reduces critical population densities necessary for persistence in small species. Previous relationships between body weight and population abundance may also be artefacts of the way in which data were gathered. 4 Despite large samples, we failed to locate the mode in plots of the number of species in each abundance category (species: abundance distribution). 5 Species: body length and total number of individuals: body length plots were similar to those found in previous studies, although using data for Coleoptera alone may have produced a steeper decline in the total number of individuals as body size increases than is apparent in samples of all arthropods. 6 We present the first three‐dimensional graph relating numbers of species, body lengths and population abundances. The surface of this three‐dimensional relationship is relatively simple.

Invertebrates as determinants and indicators of soil quality

Invertebrates are an integral part of soils and are important in determining the suitability of soils for the sustainable production of healthy crops or trees. We discuss the importance of the soil invertebrate fauna in relation to terrestrial habitats and global biodiversity as we understand it. We describe the role of the main invertebrate groups in soils, including earthworms, termites, springtails, and nematodes, and how they determine soil quality. Practical problems in dealing with the invertebrate fauna include sampling, taxonomy and availability of biological information on species. Various measures are available that use invertebrates to assess soil quality, each with its advantages and disadvantages. They include abundance, biomass, density, species richness, trophic/guild structure, food web structure, keystone species and ecosystem engineers. We propose the three most useful and practical of these as suitable to be combined with other biological (microbial) and non-biological (hydrological, physical, chemical) criteria into a single index of soil quality that might be used on a regional, if not international basis.

The composition of the arthropod fauna of Bornean lowland rain forest trees

Data on the taxonoinic composition of the arboreal arthropod fauna of 10 Bornean lowland raini forest trees are presented, based on samples obtainied using insecticide fogging. Combined samples from all trees comprised 23,874 individuals of at least 3000 species. The mean number of species on each tree was 616.7 with one tree sample containing more thani 1007 specics. The relative rank of the major orders of arthropods in terms of both species anid individuals was remarkably constanit across the trees. Hymenoptera, Coleoptera, Diptera and Hemiptera were the orders with the most species and individuals. Formnicidae, though inot particularly species rich, was the most abundanit family in terms of individuals and the commonest specics in six of the 10 trees was an ant. Refogging of one of the trees 10 days after initial sampling showed that the arthropod fauna had not completely recovered. Wlhat many arthropods are doinig in the canopy is discussed.

Guild structure of arthropods from Bornean rain forest trees

ABSTRACT. 1. 23275 arthropods collected by insecticide fogging from ten Bornean lowland rain forest trees were sorted to approximately 3000 species and assigned to guilds using two sets of criteria.

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.

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 29 986 beetles of 1473 species and 77 families. There were similar numbers of individuals (canopy 14 473; ground 15 513) 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≥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.