Biodiversity loss from deforestation may be partly offset by the expansion of secondary forests and plantation forestry in the tropics. However, our current knowledge of the value of these habitats for biodiversity conservation is limited to very few taxa, and many studies are severely confounded by methodological shortcomings. We examined the conservation value of tropical primary, secondary, and plantation forests for 15 taxonomic groups using a robust and replicated sample design that minimized edge effects. Different taxa varied markedly in their response to patterns of land use in terms of species richness and the percentage of species restricted to primary forest (varying from 5% to 57%), yet almost all between-forest comparisons showed marked differences in community structure and composition. Cross-taxon congruence in response patterns was very weak when evaluated using abundance or species richness data, but much stronger when using metrics based upon community similarity. Our results show that, whereas the biodiversity indicator group concept may hold some validity for several taxa that are frequently sampled (such as birds and fruit-feeding butterflies), it fails for those exhibiting highly idiosyncratic responses to tropical land-use change (including highly vagile species groups such as bats and orchid bees), highlighting the problems associated with quantifying the biodiversity value of anthropogenic habitats. Finally, although we show that areas of native regeneration and exotic tree plantations can provide complementary conservation services, we also provide clear empirical evidence demonstrating the irreplaceable value of primary forests.

https://www.pnas.org/content/pnas/104/47/18555.full.pdf

Quantifying the biodiversity value of tropical primary, secondary, and plantation forests.

The term beta diversity has been used to refer to a wide variety of phenomena. Although all of these encompass some kind of compositional heterogeneity between places, many are not related to each other in any predictable way. The present two-part review aims to put the different phenomena that have been called beta diversity into a common conceptual framework, and to explain what each of them measures. In this first part, the focus is on defining a beta component of diversity. This involves deciding what diversity is and how the observed total or gamma diversity (g) is partitioned into alpha (a) and beta (b) components. Several different definitions of ‘‘beta diversity’’ that result from these decisions have been used in the ecological literature. True beta diversity is obtained when the total effective number of species in a dataset (true gamma diversity g) is multiplicatively partitioned into the effective number of species per compositionally distinct

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A diversity of beta diversities: straightening up a concept gone awry. Part 1. Defining beta diversity as a function of alpha and gamma diversity

One of the primary goals of research on bioindicators is to identifyspecies or other taxonomic units that would reliably indicate disturbances inthe environment, and reflect the responses of other species or the overallbiodiversity. However, there is no perfect bioindicator and selecting the mostsuitable one depends to a great extent on the goal of the survey. In this paperwe examine the suitability of carabids as bioindicators. Carabids are frequentlyused to indicate habitat alteration. They have been used in grasslands andboreal forests where species number and/or abundances have been noted to changealong a habitat disturbance gradient. A common trend is that large, poorlydispersing specialist species decrease with increased disturbance while smallgeneralist species with good dispersal ability increase. Some species are notaffected by moderate disturbance. There is, however, not enough research todetermine how suitable carabids are for biodiversity studies, or how well theyrepresent the response of other species. We conclude that carabids are usefulbioindicators, but as crucial understanding of their relationship with otherspecies is incomplete, they should be used with caution.

Ground beetles (Coleoptera: Carabidae) as bioindicators

Land use and related pressures have reduced local terrestrial biodiversity, but it is unclear how the magnitude of change relates to the recently proposed planetary boundary (“safe limit”). We estimate that land use and related pressures have already reduced local biodiversity intactness—the average proportion of natural biodiversity remaining in local ecosystems—beyond its recently proposed planetary boundary across 58.1% of the world’s land surface, where 71.4% of the human population live. Biodiversity intactness within most biomes (especially grassland biomes), most biodiversity hotspots, and even some wilderness areas is inferred to be beyond the boundary. Such widespread transgression of safe limits suggests that biodiversity loss, if unchecked, will undermine efforts toward long-term sustainable development.

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Has land use pushed terrestrial biodiversity beyond the planetary boundary? A global assessment

Nuptial feeding encompasses any form of nutrient transfer from the male to the female during or directly after courtship and/or copulation. In insects, nuptial gifts may take the form of food captured or collected by the male, parts, or even the whole of the male's body, or glandular products of the male such as salivary secretions, external glandular secretions, the spermatophore and substances in the ejaculate. Over the past decade, there has been considerable debate over the current function of nuptial feeding in insects. This debate has centred on the issue of whether nuptial gifts function as paternal investment (i.e. function to increase the fitness and/or number of the gift-giving male's own offspring) or as mating effort (i.e. function to attract females, facilitate coupling, and/or to maximize ejaculate transfer), although the two hypotheses are not mutually exclusive. In the present article, evidence for the potential of nuptial gifts to function as either paternal investment, mating effort, or both is reviewed for each form of nuptial feeding in each insect taxon for which sufficient data are available. Empirical evidence suggests that many diverse forms of nuptial feeding in different insect taxa function, at least in part, as mating effort. For example, nuptial prey and salivary masses in the Mecoptera, regurgitated food in Drosophila (Diptera), hind-wing feeding in Cyphoderris (Orthoptera) and the secretion of the male's cephalic gland in Neopyrochroa (Coleoptera) and Zorotypus (Zoraptera) appear to function to entice females to copulate and/or to facilitate coupling. Nuptial prey and salivary masses in the Mecoptera also appear to function to maximize ejaculate transfer (which is also a form of mating effort), as do nuptial prey in Empis (Diptera), external glandular secretions in Oecanthus and Allonemobius (Orthoptera) and the spermatophylax in gryllids and tettigoniids (Orthoptera). Large spermatophores in, for example, the Lepidoptera and Coleoptera, also appear to be maintained by selection on the male to maximize ejaculate transfer and thereby counter the effects of sperm competition. In contrast to the large amount of evidence in support of the mating effort hypothesis, there is a relative lack of good evidence to support the paternal investment hypothesis. Certain studies have demonstrated an increase in the weight and/or number of eggs laid as a result of the receipt of larger gifts, or a greater number of gifts, in tettigoniids, gryllids, acridids, mantids, bruchid beetles, drosophilids and lepidopterans. However, virtually all of these studies (with the possible exception of studies of the spermatophylax in tettigoniids) have failed to control adequately for hormonal substances in the ejaculate that are known to affect female reproductive output. Furthermore, in at least four tettigoniids (but not in the case of two species), three lepidopterans, a drosophilid and probably also bruchid beetles and bittacids, evidence suggests that the male has a low probability of fertilising the eggs that stand to benefit from his nuptial gift nutrients. Therefore, the hypothesis that paternal investment might account for the function of nuptial gifts in general is not supported.

The function of nuptial feeding in insects: a review of empirical studies

Sustainable use of tropical forest systems requires continuous monitoring of biological diversity and ecosystem functions. This can be efficiently done with ‘early warning‘ (short-cycle) indicator groups of non-economical insects, whose population levels and resources are readily measured. Twenty-one groups of insects are evaluated as focal indicator taxa for rapid assessment of changes in Neotropical forest systems. Composite environmental indices for heterogeneity, richness, and natural disturbance are correlated positively with butterfly diversity in 56 Neotropical sites studied over many years. Various components of alpha, beta and gamma-diversity show typical responses to increased disturbance and different land-use regimes. Diversity often increases with disturbance near or below natural levels, but some sensitive species and genes are eliminated at very low levels of interference. Agricultural and silvicultural mosaics with over 30% conversion, including selective logging of three or more large trees per hectare, show shifts in species composition with irreversible loss of many components of the butterfly community, indicating non-sustainable land and resource use and reduction of future options. Monitoring of several insect indicator groups by local residents in a species-rich Brazilian Amazon extractive reserve has helped suggest guidelines for cologically, economically, and socially sustainable zoning and use regimes.

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Diversity, disturbance, and sustainable use of Neotropical forests: insects as indicators for conservation monitoring

The Atlantic Forest region (wide sense) includes very complex tropical environments, increasingly threatened by extensive anthropogenic conversion (>90%). Ecologically specialized, short-generation insects (butterflies) are evaluated here as indicators for monitoring community richness, landscape integrity, and sustainable resource use in the region. The > 2100 butterfly species in the Atlantic Forest region have been censused in many sites over 35 years, giving comparable daily, weekly, monthly, and long-term site lists. The 21 most thoroughly studied sites include 218–914 species, of which half can be censused in a week or less. The butterfly communities are divided into six relatively distinct faunal regions, centered in the northeast, the central coastal tablelands, the southeast coastal plain, the mountains plus interior of the southeastern states, the central plateau, and the southern states. Species richness shows the highest values in coastal mountains from 15 to 23°S. Local butterfly com...

Atlantic Forest Butterflies: Indicators for Landscape Conservation1

Ithomiine butterflies, the original distasteful models in neotropical mimicry complexes, show warning coloration and are usually rejected by both vertebrate and invertebrate predators1–4. Based on co-evolutionary theory1,5–7 and the Danainae–cardenolide example7, this protection has been ascribed to toxic compounds sequestered by larval Ithomiinae from their host-plants, usually poisonous Solanaceae8–10. However, although the giant tropical orb spider Nephila clavipes cuts out field-caught Ithomiinae from its web4,11 it readily eats freshly emerged adults. Solanaceae toxins are not found stored in adult Ithomiinae6,11 and when applied to the palatable1 butterfly Biblis hyperia do not induce rejection behaviour in Nephila (Table 1). I show here that the protection of ithomiines against this abundant predator is due to dehydropyrrolizidine alkaloid monoesters and their N-oxides, absent from Solanaceae but sequestered by these butterflies as adults from flowers (mostly Eupatorieae) and decomposing foliage (mostly Boraginaceae)11–13. These compounds are also important in the reproduction of the Ithomiinae11,14.

Adult-obtained pyrrolizidine alkaloids defend ithomiine butterflies against a spider predator

A generic-level phylogeny for the butterfly family Nymphalidae was produced by cladistic analysis of 234 characters from all life stages. The 95 species in the matrix (selected from the 213 studied) represent all important recognized lineages within this family. The analysis showed the taxa grouping into six main lineages. The basal branch is the Libytheinae, with the Danainae and Ithomiinae on the next branch. The remaining lineages are grouped into two main branches: the Heliconiinae-Nymphalinae, primarily flower-visitors (but including the fruit-attracted Coeini); and the Limenitidinae (sensu strictu), Biblidinae, and the satyroid lineage (Apaturinae, Charaxinae, Biinae, Calinaginae, Morphinae, Brassolinae, and Satyrinae), primarily fruit-attracted. Data partitions showed that the two data sets (immatures and adults) are very different, and a partitioned Bremer support analysis showed that the adult characters are the main source of conflict in the nodes of the combined analysis tree. This phylogeny includes the widest taxon coverage of any morphological study on Nymphalid butterflies to date, and supports the monophyly and relationships of most presently recognized subgroups, providing strong evidence for the presently accepted phylogenetic scheme.

Phylogeny Of The Nymphalidae (lepidoptera).

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Keith S. Brown

Papers

Diversity, disturbance, and sustainable use of Neotropical forests: insects as indicators for conservation monitoring

Atlantic Forest Butterflies: Indicators for Landscape Conservation1

Adult-obtained pyrrolizidine alkaloids defend ithomiine butterflies against a spider predator

Phylogeny Of The Nymphalidae (lepidoptera).

Phylogeny of the Nymphalidae (Lepidoptera).