Quaternary beetles : a review and issues for Australian studies

Few Australian entomologists, ecologists, biogeographers or Quaternary researchers are familiar with the details of Quaternary beetle research. Since the 1950s the study of fossil beetles has developed to become an important discipline of the Quaternary sciences. Unfortunately, however, the significance of the discipline for ecological and evolutionary research has been slow to penetrate mainstream entomological, ecological, and evolutionary thought. This paper outlines the history, methods and results of Quaternary beetle studies, based primarily upon research from the well-studied Northern Hemisphere, and then examines issues relevant to Australian research. Analysis of Quaternary beetle assemblages from Australia can contribute to the reconstruction of past environments and climates, in particular quantitative estimation of past temperature regimes, and potentially, effective precipitation. Of more significance to entomology, however, is the potential to reconstruct climatically-induced changes in distribution, essential for understanding the Quaternary biogeographic history of Australia's insect fauna. Furthermore, it will be possible to examine evidence, or the lack thereof, for speciation during the Quaternary, in the context of Quaternary environmental change.

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.

Species overlap, speciation and the evolution of aggregation pheromones in bark beetles

Theoretical predictions suggest that species-specific signals used in the attraction of mates should evolve to reduce the risk of mismating and hybridization. These predictions lead to the hypothesis that the signals of spatially overlapping (i.e. sympatric or syntopic) species should differ more substantially than those of non-overlapping species. Earlier studies have tested this prediction for auditory and visual signals. Here we test the hypothesis using olfactory signals, specifically the aggregation pheromones of species from two genera of bark beetles, Dendroctonus and Ips. We found no direct evidence from within these genera regarding the fact that the chemical blends that make up these pheromones differ more substantially in species that overlap in their geographical ranges and/or host-tree use than in allopatric taxa. However, when comparing between genera, the pheromones of overlapping species appear to be more similar than non-overlapping species. We hypothesize that the species of host tree utilized by the beetles may have some influence on their pheromone blends. Additionally, within genera, species that overlap in host use tend to be more closely related than species that use different hosts. These results may provide indirect evidence for an effect of species overlap on the evolution of bark beetle pheromones.

Plantations, not farmlands, cause biotic homogenisation of ground-active beetles in South-Eastern Australia

Following landscape change, species invasions and extinctions may lead to biotic homogenisation, resulting in increased taxonomic and functional similarity between previously distinct biotas. Biotic homogenisation is more likely to occur in landscapes where the matrix contrasts strongly with native vegetation patches. To test this, we examined the distribution of ground-active beetles in a landscape of remnant Eucalyptus open woodland patches where large areas of lower contrast matrix (farmland) are being transformed to high-contrast pine plantations in south-eastern Australia. We sampled beetles from 30 sites including six replicates of five categories; (1) remnants adjacent to farmland, (2) remnants adjacent to plantation, (3) farmland, (4) plantation, and, (5) remnants between pine plantation and farmland. Community composition in the pine matrix was similar to native patches embedded in pine (ANOSIM, Global R=. 0.49, P<. 0.000), which we suggest is due to biotic homogenisation. Remnant patches with edges of both farmland and pine plantation did not represent an intermediate community composition between patches surrounded by either matrix type, but rather a unique habitat with unique species. Farmland supported the greatest number of individuals (. F=. 9.049, df. =. 25, P<. 0.000) and species (. F=. 5.875, df. =. 25, P=. 0.002), even compared to native remnant patches. Our results suggest that matrix transformations can reduce species richness and homogenise within-patch populations. This may increase the risk of species declines in fragmented landscapes where plantations are not only replacing native vegetation patches, but also other matrix types that may better support biodiversity. Our findings are particularly concerning given expanding plantation establishment worldwide.

Unveiling the diversification dynamics of Australasian predaceous diving beetles in the Cenozoic

During the Cenozoic, Australia experienced major climatic shifts that have had dramatic ecological consequences for the modern biota. Mesic tropical ecosystems were progressively restricted to the coasts and replaced by arid-adapted floral and faunal communities. Whilst the role of aridification has been investigated in a wide range of terrestrial lineages, the response of freshwater clades remains poorly investigated. To gain insights into the diversification processes underlying a freshwater radiation, we studied the evolutionary history of the Australasian predaceous diving beetles of the tribe Hydroporini (147 described species). We used an integrative approach including the latest methods in phylogenetics, divergence time estimation, ancestral character state reconstruction, and likelihood-based methods of diversification rate estimation. Phylogenies and dating analyses were reconstructed with molecular data from seven genes (mitochondrial and nuclear) for 117 species (plus 12 outgroups). Robust and well-resolved phylogenies indicate a late Oligocene origin of Australasian Hydroporini. Biogeographic analyses suggest an origin in the East Coast region of Australia, and a dynamic biogeographic scenario implying dispersal events. The group successfully colonized the tropical coastal regions carved by a rampant desertification, and also colonized groundwater ecosystems in Central Australia. Diversification rate analyses suggest that the ongoing aridification of Australia initiated in the Miocene contributed to a major wave of extinctions since the late Pliocene probably attributable to an increasing aridity, range contractions and seasonally disruptions resulting from Quaternary climatic changes. When comparing subterranean and epigean genera, our results show that contrasting mechanisms drove their diversification and therefore current diversity pattern. The Australasian Hydroporini radiation reflects a combination of processes that promoted both diversification, resulting from new ecological opportunities driven by initial aridification, and a subsequent loss of mesic adapted diversity due to increasing aridity.

Are only the strong surviving? Little influence of restoration on beetles (Coleoptera) in an agricultural landscape

Habitat restoration has become an important part of biodiversity conservation in the face of extensive habitat loss and fragmentation, especially in agricultural landscapes. Study of invertebrates such as beetles (Coleoptera) may be important to assess the effectiveness of restoration techniques in maintaining native fauna, because they provide a variety of trophic roles and ecosystem services. In this study we examined the conservation value for beetles of revegetation in linear strips and alongside remnant patches compared with remnant vegetation and cleared roadsides. We also assessed how habitat variables structured beetle community composition. Beetle species richness and abundance did not substantially differ between revegetated, remnant and cleared areas, and was not substantially influenced by vegetation type and structure. Herbivorous beetles and the family Curculionidae were more species rich in cleared linear strips. Beetle fauna in these agricultural landscapes may be a robust subset of the pre-clearing beetle community, possibly due to the widespread degradation of remnant areas and the ground layer habitats within them. One beetle species had slightly higher abundance in remnant linear strips, suggesting that remnant habitats may be important for some beetle species. Importantly, environmental variables strongly influenced beetle community composition, signifying that beetle communities are still responding to factors such as soil type and native vegetation, rather than variables mainly associated with land management. The restoration practices currently being undertaken in agricultural areas may not maintain beetle species that require specific habitat variables to survive. Ground-layer attributes need to be included in future revegetation projects, and translocation of specialist species of beetles may be required to restore communities.

Predation of a small passerine by the Purple-winged Roller Coracias temminckii, an endemic species of Sulawesi

The usual diet of the Purple-winged Roller, a species endemic to Sulawesi, is grasshoppers, locusts, beetles and small lizards. This note describes an observation of this species preying on an adult Eurasian Tree Sparrow Passer montanus. The presence of birds in the diet of other roller species is discussed.

Effects of humidity on the response of the bark beetle Ips grandicollis (Eichhoff) (Coleoptera: Curculionidae: Scolytinae) to synthetic aggregation pheromone

Environmental factors may affect chemical communication between individuals by limiting their ability to detect and respond to these signals. One such factor, high humidity, has been shown to interfere with the normal response of some invertebrate species to their attractant pheromones. The effect of humidity on the response of the five-spined bark beetle, Ips grandicollis, to a synthetic form of the aggregation pheromone component ipsenol, was tested in an experimental chamber in the laboratory. The response was measured as both the number of beetles to reach the pheromone source and the time taken, and was tested under high (>80% relative humidity) and low (30–40% relative humidity) conditions of humidity. There was no significant difference in response of beetles between the two treatments although there was a reduction in response in the high-humidity treatment when relative humidity levels were in excess of 90%. These findings suggest that atmospheric humidity does not influence bark beetles response to synthetic pheromone, except perhaps in unlikely conditions of excessive humidity.

Positive and negative effects of phoretic mites on the reproductive output of an invasive bark beetle

When multiple species coexist upon a single host, their combined effect on the host can be unpredictable. We explored the effect of phoretic mites on the reproductive output of the five-spined bark beetle, Ips grandicollis. Using correlative approaches and experimental manipulation of mite numbers we examined how mite load affected the number, size and condition of bark beetle offspring produced. We found that mites have both negative and positive consequences on different aspects of bark beetle reproduction. Females from which mites were removed were more fecund and produced larger offspring than females with mites, implying a cost of mite loads. However, when mites were present on females, those bearing the highest mite loads produced offspring that were larger and in better condition, indicating a beneficial effect of mites. These data suggest that phoretic interactions between mites and bark beetles differ over the course of the host’s lifespan, with either the mites interacting in different ways with different life stages of the host (parasitic on adult, mutualistic with larvae), and/or the beetles being host to different mite assemblages over their lifetime.

Harem size and oviposition behaviour in a polygynous bark beetle

1. Harem polygyny can have fitness benefits and costs on females. In bark beetles of the genus Ips the latter may include within-harem competition between larvae. However, earlier competition between females for male care and mating opportunities may also influence oviposition behaviour. There has been relatively little investigation into the relationship between harem size and initial egg output. The present study investigated this relationship in the bark beetle Ips grandicollis.
2. The measure of egg output used was the number of eggs in the gallery with the most eggs in each harem. Mean ( ± SE) harem size of 242 observed harems was 3.25 ± 0.10. A curvilinear relationship was found between egg output and harem size, with females in smaller harems (one to four females) laying more eggs with increased harem size. However, females in larger harems (five to seven females) laid fewer eggs as harem size increased. The optimal harem size (in terms of number of eggs laid) was close to four females.
3. We found no evidence from a behavioural assay that females could preferentially choose unmated males over mated males with harems of two females. Additionally, the distribution of harem sizes suggests that females distribute themselves among males randomly.
4. The results suggest that harem size has effects on female reproduction that extend beyond larval competition and influence patterns of oviposition. The mechanism that determines why egg laying is greatest at intermediate levels is unknown. There is no evidence that smaller harems belong to lower quality males, but females may adjust egglaying behaviour in large harems as a result of reduced male attendance or anticipated larval competition.

The mode of evolution of aggregation pheromones in Drosophila species

Aggregation pheromones are used by fruit flies of the genus Drosophila to assemble on breeding substrates, where they feed, mate and oviposit communally. These pheromones consist of species-specific blends of chemicals. Here, using a phylogenetic framework, we examine how differences among species in these pheromone blends have evolved. Theoretical predictions, genetic evidence, and previous empirical analysis of bark beetle species, suggest that aggregation pheromones do not evolve gradually, but via major, saltational shifts in chemical composition. Using pheromone data for 28 species of Drosophila we show that, unlike with bark beetles, the distribution of chemical components among species is highly congruent with their phylogeny, with closely related species being more similar in their pheromone blends than are distantly related species. This pattern is also strong within the melanogaster species group, but less so within the virilis species group. Our analysis strongly suggests that the aggregation pheromones of Drosophila exhibit a gradual, not saltational, mode of evolution. We propose that these findings reflect the function of the pheromones in the ecology of Drosophila, which does not hinge on species specificity of aggregation pheromones as signals.

Reassembling a lost lowland carabid beetle assemblage (Coleoptera) from Kauai, Hawaiian Islands

A late Holocene but prehistoric carabid beetle fauna from the lowland Makauwahi Cave, Kauai, is characterised. Seven extinct species - Blackburnia burneyi, B. cryptipes, B. godzilla, B. menehune, B. mothra, B. ovata and B. rugosa, spp. nov. (tribe Platynini) - represent the first Hawaiian insect species to be newly described from subfossil specimens. Four extant Blackburnia spp. - B. aterrima (Sharp), B. bryophila Liebherr, B. pavida (Sharp), and B. posticata (Sharp) - and three extant species of tribe Bembidiini - Bembidion ignicola Blackburn, B. pacificum Sharp and Tachys oahuensis Blackburn - are also represented. All subfossil fragments are disarticulated, with physical dimensions and cladistic analysis used to associate the major somites - head, prothorax and elytra - for description of the new species. The seven new Makauwahi Cave species support recognition of a lowland area of endemism adjoining Haupu, a low-stature 700m elevation ridgeline in southern Kauai. Four of the extinct Blackburnia are adelphotaxa to extant species currently found at higher elevations in Kauai. Addition of these lowland specialists to the phylogenetic hypothesis undercuts applicability of the taxon cycle for interpreting evolutionary history of these taxa. Two of the extinct species are Kauai representatives in clades that subsequently colonised younger Hawaiian Islands, enhancing support for the progressive biogeographic colonisation of the archipelago by this lineage. And three of the extinct Blackburnia species comprised larger beetles than those of any extant Kauai Blackburnia, consistent with the evolution of island gigantism in the lowland habitats of Kauai.

Breeding habitat selection in an obligate beach bird: a test of the food resource hypothesis

The food resource hypothesis of breeding habitat selection in beach-nesting birds suggests that birds breed at sites with more prey to meet the increased energetic requirements associated with breeding. We compare prey resources using pitfall traps and core samples at breeding sites and absence sites of the eastern population of hooded plover, Thinornis rubricollis rubricollis, which, in this part of its range, is a threatened obligate beach bird. Breeding sites had higher abundances, equivalent species richness, and different assemblages of invertebrate prey compared with absence sites. Assemblages at breeding sites were characterised by more isopods, and fewer beetles of the family Phycosecidae. Breeding habitat selection by plovers appears to be associated with selection for sites with more food, and any process that degrades food resources at a site (e.g. kelp harvesting or marine pollution events) may reduce the likelihood of occupancy of that site by breeding birds.