The mode of pheromone evolution: evidence from bark beetles

Sex and aggregation pheromones consist of species-specific blends of chemicals. The way in which different species' blends have evolved has been the subject of some debate. Theoretical predictions suggest that differences between species have arisen not through the accruing of small changes, but through major shifts in chemical composition. Using data on the aggregation pheromones of 34 species of bark beetle from two genera, Dendroctonus and Ips, we investigated how the distributions of the chemical components of their pheromone blends mirror their phylogenetic relationships. We tested whether there were consistent patterns that could be used to help elucidate the mode of pheromone evolution. Although there were obvious differences in pheromone blends between the two genera, the differences between species within each genus followed a less clear phylogenetic pattern. In both genera, closely related species are just as different as more distantly related species. Within Dendroctonus, particularly, most chemical components were distributed randomly across the phylogeny. Indeed, for some chemicals, closely related species may actually be more different than would be expected from a random distribution of chemical components. This argues strongly against the idea of minor shifts in pheromone evolution. Instead, we suggest that, within certain phylogenetic constraints, pheromone evolution in bark beetles is characterized by large saltational shifts, resulting in sibling species being substantially phenotypically (i.e. pheromonally) different from one another, thus agreeing with theoretical predictions.     

The evolutionary significance of wing dimorphism in carabid beetles (Coleoptera: Carabidae)

A review of data on the background of wing dimorphism in carabid beetles (Coleoptera: Carabidae) and especially of the closely relatedCalathus cinctus andC. melanocephalus is given. In bothCalathus species wing dimorphism is inherited in a simple Mendelian fashion with the brachypterous condition dominant, but inC. melanocephalus the expression of the long winged genotype is under environmental control as well. The development of long winged phenotypes in the latter species is favoured by relatively favourable environmental conditions, such as high temperatures and a high food-supply. The higher fecundity of the larger and heavier long winged females of both species may compensate for losses of long winged phenotypes by flight activities. The evolutionary significance of both types of inheritance is discussed in relation to dispersal. The ‘fixed type’ as found inC. cinctus is considered an opportunistic short term ‘between sites strategy’, whereas the ‘dynamic type’ ofC. melanocephalus represents a flexible long term ‘within sites strategy’.     

The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests

As compared to natural forests, managed boreal forests are younger, more homogeneous in terms of tree age and species composition, and consist of smaller fragments. Here we examine the effects of such characteristics caused by forestry on carabid beetles (Coleoptera, Carabidae) in the boreal region. The main results are the following. (1) Fragmentation of forests and the size of a fragment appear not to be crucial for the survival of the majority of forest carabids, as they tend to be distributed over various successional stages, but species requiring old-growth habitats suffer. (2) For carabids there appear to be no or very few edge specialist species, and forest-open land edges appear to be effective barriers for species associated with forest or open habitat. However, generalist species easily cross the edge, and edges of forest fragments may be invaded by species from the surrounding open habitat. (3) Habitat change following clear-cutting dramatically changes the composition of carabid assemblages: species restricted to mature forests disappear and open-habitat species invade, while habitat generalists survive at least in the short term. Carabid diversity can probably best be maintained if forest management mimics natural processes, maintains natural structures and includes the natural composition of vegetation and other structural elements (such as dead wood) within the stands, provided that these forest features can be maintained and recreated through forest management practices. At a larger scale, the whole spectrum of forest types and ages (especially old-growth forests), and different successional processes (especially fire) should be maintained. These require the development and use of innovative logging methods, and the planning, implementation, and assessment of landscape-scale ecological management strategies.     

Phylogeny of carabid beetles as inferred from 18S ribosomal DNA (Coleoptera: Carabidae)

The phylogeny of carabid tribes is examined with sequences of 18S ribosomal DNA from eighty-four carabids representing forty-seven tribes, and fifteen outgroup taxa. Parsimony, distance and maximum likelihood methods are used to infer the phylogeny. Although many clades established with morphological evidence are present in all analyses, many of the basal relationships in carabids vary from analysis to analysis. These deeper relationships are also sensitive to variation in the sequence alignment under different alignment conditions. There is moderate evidence against the monophyly of Migadopini + Amarotypini, Scaritini + Clivinini, Bembidiini and Brachinini. Psydrini are not monophyletic, and consist of three distinct lineages (Psydrus, Laccocenus and a group of austral psydrines, from the Southern Hemisphere consisting of all the subtribes excluding Psydrina). The austral psydrines are related to Harpalinae plus Brachinini. The placements of many lineages, including Gehringia, Apotomus, Omophron, Psydrus and Cymbionotum, are unclear from these data. One unexpected placement, suggested with moderate support, is Loricera as the sister group to Amarotypus. Trechitae plus Patrobini form a monophyletic group. Brachinini probably form the sister group to Harpalinae, with the latter containing Pseudomorpha, Morion and Cnemalobus. The most surprising, well supported result is the placement of four lineages (Cicindelinae, Rhysodinae, Paussinae and Scaritini) as near relatives of Harpalinae + Brachinini. Because these four lineages all have divergent 18S rDNA, and thus have long basal branches, parametric bootstrapping was conducted to determine if their association and placement could be the result of long branch attraction. Simulations on model trees indicate that, although their observed association might be due to long branch attraction, there was no evidence that their placement near Harpalinae could be so explained. These simulations also suggest that 18S rDNA might not be sufficient to infer basal carabid relationships.     

Movements of carabid beetles (Coleoptera: Carabidae) inhabiting cereal fields: a field tracing study

Summary The individual movements of four carabid species were investigated by tracing beetles in the field using a portable radar system (harmonic radar). The field-inhabiting a Pterostichus melanarius Illiger, P. niger Schaller and Harpalus rufipes De Geer were traced in cereal fields. The movements of P. melanarius Mnd P. niger were also tracked in a wood and then compared with the movements of the forest-species Carabus nemoralis Müller. P. melanarius and P. niger displayed directed movements in cereal fields, although P. niger moved faster than P. melanarius. The movement pattern of H. rufipes in the field, and C. nemoralis and P. malanarius in the wood can be described as a c correlated random walk. In cereal fields, P. melanarius burrowed actively into the soil, and P. niger searched out crevices. In contrast, none of these two carabids burrowed in the forest soil, whereas C. nemoralis did. No statistically significant difference was observed between the movements of hungry and satiated beetles, although starved beetles tended to disperse shorter distances per move and turned back more often (especially P. niger). The significance of the various movement patterns exhibited by the species studied for their reproduction and survival on arable land is discussed.     

The occurrence and distribution of carabid beetles (Carabidae) on islands in the Baltic Sea: a review

The distribution and occurrence of carabid beetles have been studied extensively in a number of archipelagoes in the Baltic Sea, a ca. 377,000 km², fairly shallow and young sea in northern Europe. This work has revealed some surprising results related to colonisation success and maintenance of populations. Dispersal from the mainland and between islands appears to be relatively easy because inter-island distances are small (tens or hundreds of metres) and the salinity of the Baltic is low, which allows survival during drift. This is reflected in the higher than expected proportion of brachypterous species on small islands. A lack of energetically expensive wings, flight muscles and flight fuel may give brachypterous species an advantage - in terms of resource allocation for survival and fecundity - over macropterous species in the colonisation of an island. Once an island is reached, successful colonisation mainly depends on the presence of suitable habitat. The most abundantly collected carabids on these islands are habitat generalists, but some degree of habitat association exists. Yet, the composition of carabid assemblages differs substantially among islands, possibly a result of chance effects. There is evidence that species on smaller islands occur in a wider variety of vegetation types than species on larger islands. Both habitat diversity and area per se are implicated in the positive species-area relationships found, but the small island effect has been identified as being significant in most of the archipelagoes investigated. Pterostichus melanarius, a eurytopic, wing-polymorphic species abundantly collected on the mainland is, unexpectedly, absent from small islands. Possible reasons for this include; (1) interspecific predation and competition for food by the closely related but slightly larger P. niger, which is dominant on small islands, (2) the apparent inability of P. melanarius to reproduce on small islands, (3) its reduced dispersal power as compared to P. niger and (4) the possible absence of some essential resources from small islands. In conclusion, the conservation value of highly threatened cultural landscapes on Baltic Sea islands, such as species rich wooded meadows, is discussed as well as the possible negative effects of climate change on carabid beetles.     

Microorganisms in the gut of beetles: evidence from molecular cloning

We have regularly cultured yeasts from the gut of certain beetles in our ongoing research. In this study cloned PCR products amplified from the gut contents of certain mushroom-feeding and wood-ingesting beetles in four families (Erotylidae, Tenebrionidae, Ciidae, and Passalidae) were sequenced and compared with culture results. Cultural techniques detected some yeasts present in the gut of the beetles, including a Pichia stipitis-like yeast associated with wood-ingesting passalid beetles. Clone sequences similar to several ascomycete yeasts and Malassezia restricta, a fastidious basidiomycetous yeast requiring special growth media, however, were not detected by culturing. Unexpectedly, phylogenetic analysis of additional clone sequences discovered from passalid beetles showed similarity to members of the Parabasalia, protists known from other wood-ingesting insects, termites, and wood roaches. Examination of all gut regions of living passalids, however, failed to reveal parabasalids, and it is possible that they were parasites in the gut tissue present in low numbers.     

The effects of carabid beetles (Coleoptera: Carabidae) on the arthropod fauna of wheat fields in Chile

The role of carabid beetles in reducing populations of phytophagous insects has been an elusive subject. A field experiment was established on a commercial wheat crop (cv. Otto) with an area of 4.5 ha in Valdivia, Chile, during the spring and summer of 1996-1997. The field had been under a prairie system for two years, before wheat sowing (fertilization and a pesticide had been applied during crop development). Samples were taken at approximately monthly intervals. Carabid beetles were sampled with a grid of pitfall traps and other insects were sampled with a vacuum insect net and soil cores. The genera of the carabids found are of neotropical origin. Exclusion by polythene barriers, together with removal of carabid beetles using traps, was an effective technique for controlling carabid populations in a commercial wheat crop. A reduction in the number of carabid beetles was associated with an increase in the number of springtails and arachnids, and a decrease of agromyzid adults. Phytophagous insects, such as homopterans and lepidopterous larvae, were not affected by carabid exclusion and removal. The action of carabid beetles on the arthropod fauna can be extremely complex, due to its predatory activity at multitrophic levels.     

Gap felling as a forest harvesting method in boreal forests: responses of carabid beetles (Coleoptera, Carabidae)

In order to study the ecological effects of logging combined with mechanical soil preparation, we sampled carabid beetles with pitfall trapping in nine spruce-dominated stands in central Finland in 1995–1998. Three of the stands were left intact as controls. Three 1600-m² openings per stand were logged in the winter 1995–1996 into six stands. In three of these stands, light soil preparation was applied. Logging affected the species assemblages, but soil preparation per se had no clear effects. Open-habitat species increased in abundance in the openings one year after logging, but catches of generalist species in the different treatments did not differ from each other. A forest species, Calathus micropterus , was least abundant in the prepared openings. The amounts of logging residue, exposed mineral soil and aggregated humus, as well as the abundance of red wood ants, significantly explained variation in carabid assemblages.     

The contribution of small individuals to density-body size relationships: examination of energetic equivalence in reef fishes

A key relationship in ecology is that between density and body size, with the emphasis placed on energetic rules constraining the abundance of larger organisms below that of smaller organisms. Most studies have focused upon the density-body size relationship at the species level. However, energy is gathered at an individual level. We therefore examined this relationship in a coral reef fish assemblage, focusing on individuals. Using a comprehensive data set, with over 14,000 observations we found that the relationship between local density and adult body size differs from the linear relationship predicted by the energetic equivalence rule. However, excluding the smallest size classes, the relationship between body size and individual abundance for intermediate to larger fish did not depart from the predicted –0.75. Unlike plants and intermediate to large reef fishes, the smallest fishes appear to have constraints that may reflect different patterns of resource acquisition.     

A simple rule for the costs of vigilance: empirical evidence from a social forager

It is commonly assumed that anti-predator vigilance by foraging animals is costly because it interrupts food searching and handling time, leading to a reduction in feeding rate. When food handling does not require visual attention, however, a forager may handle food while simultaneously searching for the next food item or scanning for predators. We present a simple model of this process, showing that when the length of such compatible handling time Hc is long relative to search time S, specifically Hc/S > 1, it is possible to perform vigilance without a reduction in feeding rate. We test three predictions of this model regarding the relationships between feeding rate, vigilance and the Hc/S ratio, with data collected from a wild population of social foragers (samango monkeys, Cercopithecus mitis erythrarchus). These analyses consistently support our model, including our key prediction: as Hc/S increases, the negative relationship between feeding rate and the proportion of time spent scanning becomes progressively shallower. This pattern is more strongly driven by changes in median scan duration than scan frequency. Our study thus provides a simple rule that describes the extent to which vigilance can be expected to incur a feeding rate cost.     

The responses of polyphagous predators to prey spatial heterogeneity: aggregation by carabid and staphylinid beetles to their cereal aphid prey

Abstract. 1. Discrete patches of aphids were artificially created in winter wheat fields in 1982 and in 1983 using field cages.
2. Aggregation by polyphagous predators at these patches following cage removal was measured using pitfall traps in patch and control areas.
3. Several species of polyphagous beetle (Carabidae and Staphylinidae) aggregated in these patches while other species did not.
4. The consequences of aggregation by polyphagous predators to a single prey species are discussed.     

Quality control by leaf-cutting ants: evidence from communities of endophytic fungi in foraged and rejected vegetation

Leaf-cutting ants of the genera Acromyrmex and Atta forage vegetation for incorporation into their mutualistic fungal gardens. However, the presence of certain endophytic fungi in this predominantly leaf-based material could affect the fungal garden and thus the choice of material by the ants. The present study was conducted to document the endophytic fungal communities occurring in the vegetation being transported by workers of Atta laevigata into their nests and to compare this community structure with that of the vegetative material subsequently rejected from the nests. We found considerable diversity in the fungi isolated. Acremonium, Cylindrocladium, Drechslera, Epicoccum, Fusarium, Trichoderma, Ulocladium and two unidentified morphospecies were significantly more common in rejected compared with foraged material, and some of these genera include mycoparasites, which could represent a threat to the fungal gardens. Conversely, Colletotrichum, Pestalotiopsis, Phomopsis, Xylaria and an unidentified morphospecies were more common in carried compared with rejected material. The possibility that ants have a ‘quality-control’ mechanism based on the presence of antagonistic fungal endophytes is discussed, as is the potential use of these fungi as biocontrol agents against Attini pests.     

The scaling of abundance in consumers and their resources: implications for the energy equivalence rule

The negative scaling of plant and animal abundance with body mass is one of the most fundamental relationships in ecology. However, theoretical approaches to explain this phenomenon make the unrealistic assumption that species share a homogeneous resource. Here we present a simple model linking mass and metabolism with density that includes the effects of consumer size on resource characteristics (particle size, density, and distribution). We predict patterns consistent with the energy equivalence rule (EER) under some scenarios. However, deviations from EER occur as a result of variation in resource distribution and productivity (e.g., due to the clumping of prey or variation in food particle size selection). We also predict that abundance scaling exponents change with the dimensionality of the foraging habitat. Our model predictions explain several inconsistencies in the observed scaling of vertebrate abundance among ecological and taxonomic groups and provide a broad framework for understanding variation in abundance.     

Disturbance regimes in a wetland remnant: implications for trait-displacements and shifts in the assemblage structure of carabid beetles (Coleoptera: Carabidae)

Studies on disturbance regimes involving carabid beetles have mainly focused on forest habitats. We therefore decided to analyze the effects of disturbance on carabid communities in a wetland remnant (Lake Chiusi, central Italy). Results highlighted the presence of a disturbance gradient affecting the species richness and trait-displacement of carabid communities. Carabids were sampled with pitfall traps from March to October 2008 at nine randomly selected sample stations; a set of landscape attributes were also collected. Principal Component Analysis (PCA) and generalized linear mixed models (GLMMs) were used to link the distribution of carabid life-history traits and species richness with the most informative combination of landscape attributes. The first PCA axis (PC1) showed significant correlation with “distance to the lake shoreline” and “perimeter-area ratio”, highlighting the presence of a disturbance-axis. The second and third axes accounted only for a trivial portion of the total variance. GLMMs revealed a progressive decrease in the number of hygrophilous species from the core of the wetland to its outer areas. Similar trends were observed for species richness and for predator species with good dispersal ability and larval period in summer. Our results highlight the importance of taking into account community-wide functional implications in landscape ecology studies.     

The systematic component of phylogenetic error as a function of taxonomic sampling under parsimony

The effect of taxonomic sampling on phylogenetic accuracy under parsimony is examined by simulating nucleotide sequence evolution. Random error is minimized by using very large numbers of simulated characters. This allows estimation of the consistency behavior of parsimony, even for trees with up to 100 taxa. Data were simulated on 8 distinct 100-taxon model trees and analyzed as stratified subsets containing either 25 or 50 taxa, in addition to the full 100-taxon data set. Overall accuracy decreased in a majority of cases when taxa were added. However, the magnitude of change in the cases in which accuracy increased was larger than the magnitude of change in the cases in which accuracy decreased, so, on average, overall accuracy increased as more taxa were included. A stratified sampling scheme was used to assess accuracy for an initial subsample of 25 taxa. The 25-taxon analyses were compared to 50- and 100-taxon analyses that were pruned to include only the original 25 taxa. On average, accuracy for the 25 taxa was improved by taxon addition, but there was considerable variation in the degree of improvement among the model trees and across different rates of substitution.