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.

“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.

Can we characterise ‘openness’ in the Holocene palaeoenvironmental record? Modern analogue studies of insect faunas and pollen spectra from Dunham Massey deer park and Epping Forest, England

This paper examines the degree to which tree-associated Coleoptera (beetles) and pollen could be used to predict the degree of ‘openness’ in woodland. The results from two modern insect and pollen analogue studies from ponds at Dunham Massey, Cheshire and Epping Forest, Greater London are presented. We explore the reliability of modern pollen rain and sub-fossil beetle assemblages to represent varying degrees of canopy cover for up to 1000m from a sampling site. Modern woodland canopy structure around the study sites has been assessed using GIS-based mapping at increasing radial distances as an independent check on the modern insect and pollen data sets. These preliminary results suggest that it is possible to use tree-associated Coleoptera to assess the degree of local vegetation openness. Additionally, it appears that insect remains may indicate the relative intensity of land use by grazing animals. Our results also suggest most insects are collected from within a 100m to 200m radius of the sampling site. The pollen results suggest that local vegetation and density of woodland in the immediate area of the sampling site can have a strong role in determining the pollen signal.

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.

The Holocene British and Irish ancient woodland fossil beetle fauna: implications for forest history, biodiversity and faunal colonisation

This paper presents a new review of our knowledge of the ancient forest beetle fauna from Holocene archaeological and palaeoecological sites in Great Britain and Ireland. It examines the colonisation, dispersal and decline of beetle species, highlighting the scale and nature of human activities in the shaping of the landscape of the British Isles. In particular, the paper discusses effects upon the insect fauna, and examines in detail the fossil record from the Humberhead Levels, eastern England. It discusses the local extirpation of up to 40 species in Britain and 15 species in Ireland. An evaluation of the timing of extirpations is made, suggesting that many species in Britain disappear from the fossil record between c. 3000 cal BC and 1000 cal BC (c. 5000-3000 cal BP), although some taxa may well have survived until considerably later. In Ireland, there are two distinct trends, with a group of species which seem to be absent after c. 2000 cal BC (c. 4000 cal BP) and a further group which survives until at least as late as the medieval period. The final clearance of the Irish landscape over the last few hundred years was so dramatic, however, that some species which are not especially unusual in a British context were decimated. Reasons behind the extirpation of taxa are examined in detail, and include a combination of forest clearance and human activities, isolation of populations, lack of temporal continuity of habitats, edaphic and competition factors affecting distribution of host trees (particularly pine), lack of forest fires and a decline in open forest systems. The role of climate change in extirpations is also evaluated. Consideration is given to the significance of these specialised ancient forest inhabitants in Ireland in the absence of an early Holocene land-bridge which suggests that colonisation was aided by other mechanisms, such as human activities and wood-rafting. Finally, the paper discusses the Continental origins of the British and Irish fauna and its hosts and the role played by European glacial refugia.

Crossing forest thresholds: inertia and collapse in a Holocene sequence from south-central Spain

A Holocene palaeoecological sequence from Villaverde, south-central Spain, is presented. The pollen stratigraphy is used to infer past vegetation changes within a catchment area that represents the boundary between semi-arid, plateau and mountain vegetation. From c. 9700–7530 cal. yr BP, Pinus is dominant, probably as a result of a combination of a relatively dry climate and natural fire disturbance. From c. 7530–5900 cal. yr BP, moderate invasion by Quercus appears to be a migrational response following increased moisture and temperature, but in part shaped by competitive adjustments. From c. 5900–5000 cal. yr BP, the pine forests are replaced by deciduous-Quercus forests with an important contribution from Corylus, Betula, Fraxinus and Alnus. Mediterranean-type forests spread from c. 5000 to 1920 cal. yr BP coincident with expansions of Artemisia, Juniperus and other xerophytes. From c. 1920–1160 cal. yr BP, Pinus becomes dominant after a disturbance- mediated invasion of the oak forests. Human impact upon the regional landscape was negligible during the Neolithic, and limited in the Bronze and Iron Ages. Local deforestation and the expansion of agro-pastoral activities occur after c. 1600 cal. yr BP.

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.

Response of surface horizons in an oak forest to prescribed burning

The reduction of forest floor ground cover and litter layers by prescribed fires may alter the morphology (field and micro) and physical properties of surface horizons. This study determined long-term (35 yr) changes in surface horizon bulk density, organic matter concentration and content, and morphology in response to periodic (5 yr) and annual (1 yr) prescribed fires. Soils were fine-silty, siliceous, thermic Glossic Fragiuldults, supporting mixed oak vegetation in middle Tennessee. Upper mineral soils (0- to 2-cm and 0- to 7.6-cm depths) were sampled and detailed field descriptions made. Periodic and control plots had a thin layer of Oi, Oe, and Oa horizons 5 yr after the 1993 burn, whereas on annual burn plots a 1- to 2-cm charred layer was present. Significant reductions in organic matter concentration and mean thickness of the A horizon were found from burning (A horizons thicknesses were 6.4, 4.6, and 2.9 cm in control, periodic, and annual plots, respectively). Periodic burns did not significantly alter the organic matter and bulk density of the upper 7.6 cm of mineral soil; however, annual burns did result in significantly higher bulk densities (1.01, 1.07, and 1.29 Mg m-3 in control, periodic, and annual plots, respectively) and lower organic matter concentrations and contents. Microscopic investigations confirmed that compaction was increased from annual burning. Thin sections also revealed that the granular structure of the A horizons in control and periodic plots resulted from bioterbation of macro and mesofauna, fungi, and roots. Long-term annual burning greatly affected surface soil properties, whereas periodic burning on a 5-yr cycle had only limited effects.