Sub-fossil Insects and ecosystem dynamics in wetlands; implications for biodiversity and conservation.

We review the uses of fossil insects, particularly Coleoptera (beetles) and Chironomidae (non-biting midges) from ancient deposits to inform the study of wetland ecosystems and their ecological and restoration processes. In particular, we focus on two contrasting ecosystems, drawing upon research undertaken by us on British raised mire peats and shallow lake systems, one an essentially terrestrial ecosystem, the other aquatic, but in which wetland insects play an important and integral part. The study of raised mire peats suggests that faunal stability is a characteristic of these wetland systems, over what appear to be extensive periods of time (up to several millennia), whilst studies of shallow lake ecosystems over recent timescales indicates that faunal instability appears to be more common, usually driven by increasing eutrophication. Drawing upon a series of fossil Coleoptera records spanning several thousand years from Hatfield Moors, south Yorkshire, we reconstruct in some detail the mire’s ontogeny and fluctuations in site hydrology and vegetation cover, illustrating the intimate association between substrate, topography and peat development. A comparison between fossil and modern beetle populations indicates that the faunal characteristics of this mire and its adjacent neighbour, Thorne Moors, become established during the early phases of peat development, including its rare endemics, and that the faunal biodiversity on the sites today is dictated by complex site histories. The over-riding characteristic of these faunas is of stability over several thousand years, which has important implications for the restoration of degraded sites, especially those where refugial areas are limited. In contrast, analyses of fossil Chironomidae from shallow lakes allow researchers to track changes in limnological status and while attempts have been made to reconstruct changes in nutrient levels quantitatively, the chironomids respond indirectly to such changes, typically mediated through complex ecosystem dynamics such as changes in fish and/or macrophyte communities. These changes are illustrated via historic chironomid stratigraphies and diversity indices from a range of shallow lakes located across Britain: Slapton Ley, Frensham Great Pond, Fleet Pond, Kyre Pool and Barnes Loch. These sites have shown varying degrees of eutrophication over recent timescales which tends to be associated with a decline in chironomid diversity. While complex functional processes exist within these ecosystems, our evidence suggests that one of the key drivers in the loss of shallow lake chironomid diversity appears to be the loss of aquatic macrophytes. Overall, while chironomids do show a clear response to altered nutrient regimes, multi-proxy reconstructions are recommended for a clear interpretation of past change. We conclude that if we are to have a better understanding of biota at the ecosystem level we need to know more of the complex interactions between different insect groups as well as with other animal and plant communities. A palaeoecological approach is thus crucial in order to assess the role of insect groups in ecosystem processes, both in the recent past and over long time scales, and is essential for wetland managers and conservation organisations involved in long term management and restoration of wetland systems.

Forest fires and insects: palaeoentomological research from a sub-fossil burnt forest.

The role of fire within Pinus-mire ecosystems is explored by focusing on a palaeoentomological investigation of the extensive burnt fossil forest preserved within the basal deposits of the raised mires of Thorne and Hatfield Moors, Humberhead Levels, eastern England. Remains of charred tree macrofossils (roots, stumps and trunks) are widely distributed across both sites, mainly comprising Pinus and Betula. Evidence from this research and elsewhere suggest fires were a common event on Pinus mires, and may indicate that such episodes played an important role in the development of raised bogs. A fire-loving (pyrophilous) insect fauna appears to have been attracted to the burnt areas and the decline and extirpation in Britain of a number of pyrophilous species (e.g. Stagetus borealis Isrealsson) suggests the former importance of this type of habitat within British Pinus-mire systems. The lack of consideration given to the role of natural fires within the British landscape is questioned and the interpretation of charcoal within mire deposits as a possible anthropogenic indicator is highlighted as an area that would benefit some reconsideration.

Melanism and disease resistance in insects

There is growing evidence that insects in high-density populations invest relatively more in pathogen resistance than those in low-density populations (i.e. density-dependent prophylaxis). Such increases in resistance are often accompanied by cuticular melanism, which is characteristic of the high-density form of many phase polyphenic insects. Both melanism and pathogen resistance involve the prophenoloxidase enzyme system. In this paper the link between resistance, melanism and phenoloxidase activity is examined in Spodoptera lanae. In S. exempta, cuticular melanism was positively correlated with phenoloxidase activity in the cuticle, haemolymph and midgut. Melanic S. exempta larvae were found to melanize a greater proportion of eggs of the ectoparasitoid Euplectrus laphygmae than non-melanic larvae, and melanic S. littoralis were more resistant to the entomopathogenic fungus Beauveria bassiana (in S. exempta the association between melanism and fungal resistance was non-signficant). These results strengthen the link between melanism and disease resistance and implicate the involvement of phenoloxidase.

Quantifying forage specialisation in polyphagic insects: the polylectic and rare solitary bee, Colletes floralis (Hymenoptera: Colletidae)

Many insect species vary in their degree of foraging specialisation, with many bee species considered polyphagic (polylectic). Wild, non-managed bee species vary in their conservation status, and species-specific biological traits such as foraging specialisation may play an important role in determining variance in population declines. Current agri-environment schemes (AESs) prescribe the introduction of flower seed mixes for agricultural systems to aid the conservation of wild bees. However, the extent to which flower combinations adequately meet bee foraging requirements is poorly known. We quantitatively assessed pollen use and selectivity using two statistical approaches: Bailey's Intervals and Compositional Analysis, in an examplar species, a purportedly polylectic and rare bee, Colletes floralis, across 7 sites through detailed analysis of bee scopal pollen loads and flower abundance. Both approaches provided good congruence, but Compositional Analysis was more robust to small sample sizes. We advocate its use for the quantitative determination of foraging behaviour and dietary preference. Although C. floralis is polylectic, it showed a clear dietary preference for plants within the family Apiaceae. Where Apiaceae was uncommon, the species exploited alternative resources. Other plant families, such as the Apiaceae, could be included, or have their proportion increased in AES seed mixes, to aid the management of C. floralis and potentially other wild solitary bees of conservation concern. © 2011 The Authors. Insect Conservation and Diversity © 2011 The Royal Entomological Society.

How and why do insects migrate?

Countless numbers of insects migrate within and between continents every year, and yet we know very little about the ultimate reasons and proximate mechanisms that would explain these mass movements. Here we suggest that perhaps the most important reason for insects to migrate is to hedge their reproductive bets. By spreading their breeding efforts in space and time, insects distribute their offspring over a range of environmental conditions. We show how the study of individual long-distance movements of insects may contribute to a better understanding of migration. In the future, advances in tracking methods may enable the global surveillance of large insects such as desert locusts.

Dynamics of macronutrient self-medication and illness-induced anorexia in virally infected insects

Some animals change their feeding behaviour when infected with parasites, seeking out substances that enhance their ability to overcome infection. This 'self-medication' is typically considered to involve the consumption of toxins, minerals or secondary compounds. However, recent studies have shown that macronutrients can influence the immune response and that pathogen-challenged individuals can self-medicate by choosing a diet rich in protein and low in carbohydrates. Infected individuals might also reduce food intake when infected (i.e. illness-induced anorexia). Here, we examine macronutrient self-medication and illness-induced anorexia in caterpillars of the African armyworm (Spodoptera exempta) by asking how individuals change their feeding decisions over the time course of infection with a baculovirus. We measured self-medication behaviour across several full-sib families to evaluate the plasticity of diet choice and underlying genetic variation. Larvae restricted to diets high in protein (P) and low in carbohydrate (C) were more likely to survive a virus challenge than those restricted to diets with a low P : C ratio. When allowed free choice, virus-challenged individuals chose a higher protein diet than controls. Individuals challenged with either a lethal or sublethal dose of virus increased the P : C ratio of their chosen diets. This was mostly due to a sharp decline in carbohydrate intake, rather than an increased intake of protein, reducing overall food intake, consistent with an illness-induced anorexic response. Over time the P : C ratio of the diet decreased until it matched that of controls. Our study provides the clearest evidence yet for dietary self-medication using macronutrients and shows that the temporal dynamics of feeding behaviour depends on the severity and stage of the infection. The strikingly similar behaviour shown by different families suggests that self-medication is phenotypically plastic and not a consequence of genetically based differences in diet choice between families. © 2013 British Ecological Society.

Climatic signals in the life histories of insects:the distribution and abundance of heather psyllids (Strophingia spp.) in the UK

1. The population density and age structure of two species of heather psyllid Strophingia ericae and Strophingia cinereae, feeding on Calluna vulgaris and Erica cinerea, respectively, were sampled using standardized methods at locations throughout Britain. Locations were chosen to represent the full latitudinal and altitudinal range of the host plants.

2. The paper explains how spatial variation in thermal environment, insect life-history characteristics and physiology, and plant distribution, interact to provide the mechanisms that determine the range and abundance of Strophingia spp.

3. Strophingia ericae and S. cinereae, despite the similarity in the spatial distribution patterns of their host plants within Britain, display strongly contrasting geographical ranges and corresponding life-history strategies. Strophingia ericae is found on its host plant throughout Britain but S. cinereae is restricted to low elevation sites south of the Mersey-Humber line and occupies only part of the latitudinal and altitudinal range of its host plant. There is no evidence to suggest that S. ericae has reached its potential altitudinal or latitudinal limit in the UK, even though its host plant appears to reach its altitudinal limit.

4. There was little difference in the ability of the two Strophingia spp. to survive shortterm exposure to temperatures as low as - 15 degrees C and low winter temperatures probably do not limit distribution in S. cinereae.

5. Population density of S. ericae was not related to altitude but showed a weak correlation with latitude. The spread of larval instars present at a site, measured as an index of instar homogeneity, was significantly correlated with a range of temperature related variables, of which May mean temperature and length of growing season above 3 degrees C (calculated using the Lennon and Turner climatic model) were the most significant. Factor analysis did not improve the level of correlation significantly above those obtained for single climatic variables. The data confirmed that S. ericae has a I year life cycle at the lowest elevations and a 2 year life cycle at the higher elevations. However, there was no evidence, as previously suggested, for an abrupt change from a one to a 2 year life cycle in S. ericae with increasing altitudes or latitudes.

6. By contrast with S. ericae, S. cinereae had an obligatory 1 year life cycle, its population decreased with altitude and the index of instar homogeneity showed little correlation with single temperature variables. Moreover, it occupied only part of the range of its host plant and its spatial distribution in the UK could be predicted with 96% accuracy using selected variables in discriminant analysis.

7. The life histories of the congeneric heather psyllids reflect adaptations that allow them to exploit host plants with different distributions in climatic and thereby geographical space. Strophingia ericae has the flexible life history that enables it to exploit C. vulgaris throughout its European boreal temperate range. Strophingia cinereae has a less flexible life history and is adapted for living on an oceanic temperate host. While the geographic ranges of the two Strophingia spp. overlap within the UK, the psyllids appear to respond differently to variation in their thermal environment.

“Islands” in Holocene forests: implications for forest openness, landscape clearance and “culture-steppe” species

Human activity has undoubtedly had a major impact on Holocene forested ecosystems, with the concurrent expansion of plants and animals associated with cleared landscapes and pasture, also known as 'culture-steppe'. However, this anthropogenic perspective may have underestimated the contribution of autogenic disturbance (e.g. wind-throw, fire), or a mixture of autogenic and anthropogenic processes, within early Holocene forests. Entomologists have long argued that the north European primary forest was probably similar in structure to pasture woodland. This idea has received support from the conservation biologist Frans Vera, who has recently strongly argued that the role of large herbivores in maintaining open forests in the primeval landscapes of Europe has been seriously underestimated. This paper reviews this debate from a fossil invertebrate perspective and looks at several early Holocene insect assemblages. Although wood taxa are indeed important during this period, species typical of open areas and grassland and dung beetles, usually associated with the dung of grazing animals, are persistent presences in many early woodland faunas. We also suggest that fire and other natural disturbance agents appear to have played an important ecological role in some of these forests, maintaining open areas and creating open vegetation islands within these systems. More work, however, is required to ascertain the role of grazing animals, but we conclude that fossil insects have a significant contribution to make to this debate. This evidence has fundamental implications in terms of how the palaeoecological record is interpreted, particularly by environmental archaeologists and palaeoecologists who may be more interested in identifying human-environment interactions rather than the ecological processes which may be preserved within palaeoecological records.

Pinus and Prostomis: a dendrochronological and palaeoentomological study of a study of a mid-Holocene woodland in Eastern England

Tree-ring analysis of sub-fossil Pinus sylvestris L. and Quercus sp. and their associated sub-fossil insect assemblages from tree rot holes have been used to study a prehistoric forest buried in the basal peats at Tyrham Hall Quarry, Hatfield Moors SSSI, in the Humberhead Levels, eastern England. The site provided a rare opportunity to examine the date, composition, age structure and entomological biodiversity of a mid-Holocene Pinus-dominated forest. The combined approaches of dendrochronology and palaeoentomology have enabled a detailed picture of the forest to be reconstructed, within a precise time frame. The Pinus chronology has been precisely dated to 2921- 2445 BC against the English Quercus master curve and represents the first English Pinus chronology to be dendrochronologically dated. A suite of important xylophilous (wood-loving) beetles that are today very rare and four species that no longer live within the British Isles were also recovered, their disappearance associated with the decline in woodland habitats as well as possible climate change. The sub-fossil insects indicate that the characteristic species of the site's modern-day fauna were already in place 4000 years ago. These findings have important implications in terms of maintaining long-term invertebrate biodiversity of mire sites.

Is big best? Queen size, usurpation and nest closure in a primitively eusocial sweat bee (Lasioglossum malachurum).

For primitively eusocial insects in which a single foundress establishes a nest at the start of the colony cycle, the solitary provisioning phase before first worker emergence represents a risky period when other, nestless foundresses may attempt to usurp the nest. In the primitively eusocial sweat bee Lasioglossum malachurum (Hymenoptera, Halictidae), spring foundresses compete for nests which are dug into hard soil. Nest-searching foundresses (‘floaters’) frequently inspected nests during this solitary phase and thereby exerted a usurpation pressure on resident queens. Usurpation has been hypothesised to increase across the solitary provisioning phase and favour closure of nests at an aggregation, marking the termination of the solitary provisioning phase by foundresses, before worker emergence. However, our experimental and observational data suggest that usurpation pressure may remain constant or even decrease across the solitary provisioning phase and therefore cannot explain nest closure before first worker emergence. Levels of aggression during encounters between residents and floaters were surprisingly low (9% of encounters across 2 years), and the outcome of confrontations was in favour of residents (resident maintains residency in 94% of encounters across 2 years). Residents were significantly larger than floaters. However, the relationship between queen size and offspring production, though positive, was not statistically significant. Size therefore seems to confer a considerable advantage to a queen during the solitary provisioning phase in terms of nest residency, but its importance in terms of worker production appears marginal. Factors other than intraspecific usurpation need to be invoked to explain the break in provisioning activity of a foundress before first worker emergence.

Temporal and spatial variation in otter Lutra lutra diet in Northern Ireland

The European otter (Lutra lutra L.) has a highly specialised diet that is composed predominantly of fish. The current study investigates the percentage composition of food items in otter spraints collected in six river catchments in Northern Ireland in 1980 and again from the same locations in 2003. Spraints contained significantly more salmonids than any other prey item. The composition of spraints differed among catchments. More salmonids and three-spined stickleback (Gasterosteus aculeatus L.) were found in spraints from the Glens of Antrim, while spraints from the Lagan catchment had significantly more eel (Anguilla anguilla L.). There were significantly more spraints containing non-fish food items in 2003 compared with 1980. These non-fish items were insects, amphipods, birds, rats (Rattus norvegicus Berkenhout) and lagomorphs. Increased consumption of non-fish items was apparent in all but one of the river catchments. The mean diversity of spraint composition was significantly greater in 2003 than in 1980. Therefore, our findings indicate that otters have diversified their diet since 1980 and now eat more non-fish prey.

Pollinators of an endemic and endangered species, Mammillaria gaumeri (Cactaceae), in its natural habitat (coastal dune) and in a botanical garden.

Mammillaria gaumeri (Britton & Ross) Orcutt (Cactaceae), an endemic plant of the Yucatan Peninsula, is included by the Mexican government in the list of species that require special protection. Its natural habitat is now restricted to fragmented areas and protection programs involve botanical gardens in growing individuals rescued from disturbed areas. Little information is available on the reproductive characteristics of this species and nothing is known of its pollinators. We investigated the visitors of M. gaumeri flowers, collecting and observing bee species in its natural habitat (i.e., coastal dune) and in a botanical garden, where coastal dune vegetation had been created. Observations were made on plants whose density was artificially increased by grouping flowering individuals. At each site, we: 1) collected insects visiting the flowers; 2) recorded number of visits; and 3) video-recorded bee movements on the flowers. As expected, the number of bee species and visitation frequency were higher at the botanical garden than at the coastal dune. After landing on a flower, bees either inspected the anthers or dived among them. These behaviors, carried out by all observed species, seemed related to the state of the anthers (full or empty of pollen) and stigma lobes (opened or closed). Specifically, visits lasted longer when anthers were full of pollen and stigma lobes were opened. The same bee species recorded on the dune were also recorded at the botanical garden, suggesting that the artificial dune at the botanical garden offered suitable conditions for the natural pollinators of this endangered cactus.