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G. Russell Coope: Papers honouring his life and career
Quaternary International 341 (2014) 1e5
Contents lists available at ScienceDirect
Quaternary International journal homepage: www.elsevier.com/locate/quaint
Guest Editorial
G. Russell Coope: Papers honouring his life and career a b s t r a c t Keywords: Quaternary Pleistocene Holocene Insect fossils Beetles Midges
The passing of Russell Coope in 2011 brought an end to a vigorous, dynamic research career that launched the field of Quaternary entomology. This issue of Quaternary International is composed mostly of papers given in his honour in June, 2012 at Royal Holloway University of London. It comprises 21 papers that cover a wide range of topics. The reconstruction of British Pleistocene environments was arguably Coope's most important contribution to science. Three papers containing previously unpublished Middle and Late Pleistocene beetle faunas and their interpretations are included here. A discussion paper on the origins of the insect faunas of North Atlantic islands echoes another of Coope's research interests, as do two studies of late glacial climates of northwest Europe. A suite of several papers discussing the environmental archaeology of sites ranging in age from the Bronze Age to the early 20th century honour Coope's pioneering work in this field. Pleistocene research from sites in North America and Japan complete the volume, followed by descriptions of two large-scale insect fossil databases. © 2014 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
2. British Pleistocene studies
Russell Coope was the pioneer in the field of Quaternary entomology, almost single-handedly bringing the field from complete obscurity to the forefront of Quaternary science, beginning in the late 1950s. His passing in 2011 brought an end to a vigorous, dynamic research career that launched the field. He also trained a younger generation of paleoentomologists who went on to work in Britain, Continental Europe and North America. His former students and other colleagues met in June, 2012 to celebrate Russell's life and contributions to science in a symposium at Royal Holloway University of London. A number of outstanding papers were presented in this symposium, many of which are presented in this special issue of Quaternary International. One of Russell Coope's defining characteristics was his wideranging interest in all aspects of natural history, not just Quaternary insects. But even within his own field, he was instrumental in developing research in both Pleistocene and Holocene insect fossils from natural and anthropogenic deposits. He helped launch this field in North America during his participation in a scientific meeting in Ohio in the 1960s, and he worked on sites throughout northwest Europe, in the process training students from France, Denmark and Sweden. It is only fitting, therefore, that the contents of this special issue are wide ranging, both geographically and topically. The authors offer these articles in honour of his memory. Coope remained active in the field of Quaternary entomology right up to the end of his life, so some of his work remained unpublished during his lifetime. Some aspects of this work are included here.
We present some of Coope's contributions to multidisciplinary research in British Pleistocene environmental reconstructions in a set of two papers led by Harry Langford. One of these concerns Middle to Late Pleistocene environments from cold/cool stage deposits at Whittlesey, Cambridgeshire, England. Organic deposits exposed along the River Nene yielded a sequence of fossil assemblages. The study examined pollen, plant macrofossils, molluscs, beetles, ostracods, foraminifera and vertebrates. The youngest of these fossil assemblages dates to MIS 3. The other three assemblages pre-date the last interglacial (MIS 5e), with the uppermost of these three apparently representing the MIS 7e6 transition. The three assemblages older than the last interglacial all contained beetle and ostracod faunas indicating mean July temperatures (TMAX) that were 1e3 C warmer than those of the LGM, but mean January temperatures (TMIN) were 8 C colder than those estimated for the LGM. Langford et al. also reported on an MIS 7 interglacial deposit from the River Nene. This is the first fully interglacial fluvial fossil assemblage described so far for this catchment. Pollen, mollusc and insect fossil remains were studied from the deposit. The palaeobotanical and insect evidence indicate the presence of extensive deciduous woodland at the site, while the ostracod evidence suggests that saline habitats were also present locally, suggesting that sea level at the time was not much different to today. Temperature estimates based on insect and ostracod data indicate that mean July temperatures were 16e21 C, while mean January temperatures were 1 to 3 C. The modern TMAX and TMIN values fall within these boundaries.
http://dx.doi.org/10.1016/j.quaint.2014.06.056 1040-6182/© 2014 Elsevier Ltd and INQUA. All rights reserved.
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Guest Editorial / Quaternary International 341 (2014) 1e5
Coope also analysed fossil insect assemblages of early Middle Pleistocene age from a site at Sidestrand in northeast Norfolk, England (Larkin et al.). Here a beetle fauna was identified that represents one of the most extensive cold-stage assemblages from this interval, with 59 taxa indicative of cold, continental climate. TMAX estimates for this fauna were 10e13 C, and TMIN estimates were 17 to 10 C. These temperature regimes are found today near northern treeline in the Arctic. The beetle evidence from this site contradicts the evidence derived from other climate proxies, such as the global ice volume record, which indicate less intense cold stages during the early Middle Pleistocene. However, the age of the Sidestrand cold-adapted fauna remains uncertain. An underlying Sidestrand Hall Member of the Cromer Forest-Bed Formation is thought to represent MIS 13, but this cold-stage fossil bed is separated from that stratigraphic horizon by shallow marine deposits representing at least two marine regression phases and a cold marine regression. 3. North Atlantic studies One of the most interesting puzzles for biogeographers is the question of how and when the North Atlantic islands were colonized by flora and fauna. This is a problem that fascinated Russell Coope, who wrote an article about it in 1986. Panagiotakopulu offers a hypothesis that insect colonization of the North Atlantic region was all postglacial, based on the lack of endemic species that might otherwise have survived on the Faroe Islands, Iceland and Greenland during the last glaciation. The bulk of the fossil data points towards postglacial immigration onto islands left devoid of insect life by the effects of the last glaciation. The evidence suggests that the majority of successful immigration efforts occurred through insects being carried west across the North Atlantic on ice rafts created by the disintegration of the Scandinavian ice sheet. Some of these massive blocks of ice may well have been covered by layers of soil and organic debris, providing temporary habitats for insects that either were carried away from the land on these blocks, or which landed on them when they were not too far from the shore, in attempted aerial dispersals from the west coastal regions of Norway. 4. British late glacial studies Study of the sequence of environmental changes during the late glacial (or ‘lateglacial’ in most British publications) interval was one of the most important themes in Russell Coope's research. He published 18 papers on this topic. Although the British late glacial interval is probably one of the best-studied in the world, it still sparks very lively debates. Elias and Matthews contributed to this topic by examining the thermal history of the British late glacial using the Mutual Climatic Range (MCR) method. They performed MCR analyses and used Bayesian approaches to recalibrate the radiocarbon chronologies for four important previously published beetle faunal sequences: Gransmoor, St. Bees, Llanilid, and Glannynau, but instead of generating MCR estimates for entire faunal assemblages, they split the assemblages into terrestrial and aquatic groups, and undertook separate MCR analyses of the two groups. Their working hypothesis was that the MCR estimates of TMAX derived from the aquatic species might be colder than those derived from terrestrial species, if late glacial pond waters were cooled by meltwater from ice sheets or snowbeds. Only a few instances of difference between the two sets of TMAX estimates were found, and all of these fell within the transition into the late glacial interstadial interval, during the transition from interstadial conditions into the Younger Dryas cooling, or during the Younger Dryas itself. The authors attributed the differences to the influence of meltwater from
snowbanks surrounding the study sites, and suggest that future late glacial beetle studies from northwest Europe should adopt the protocol of generating separate aquatic and terrestrial MCR estimates, for comparative purposes. The late glacial interval and the early Holocene were also the focus of an article by Brooks and Langdon, who examined fossil midge (Diptera: Chironomidae) records from northwest Europe in order to estimate summer air temperatures. They examined a large suite of faunal assemblages from 22 late glacial sites and 34 early Holocene sites, drawing isotherm maps across the study region. These maps show a strong east-west gradient during the late glacial interval, presumably due to the influence of thermohaline circulation in the regions bordering the North Atlantic. They also found a strong north-south gradient for the late glacial, likely due the persistent cooling effects of the Scandinavian ice sheet. Warming intervals, both during the late glacial interstadial and in postglacial times, were first seen in the south, then progressed northwards. Interestingly, the midge-based paleotemperature estimates for Northwest Europe were consistently cooler than those derived from beetle assemblages. This trend is consistent with the findings of Elias and Matthews that meltwater input in ponds and lakes during intervals of late lying snow may have cooled the water bodies in which both aquatic beetle and midge larvae lived. 5. Holocene studies Mansell et al. used insect fossils, pollen and plant macrofossil data to reconstruct Holocene floodplain palaeoecology of the Humberhead Levels region of South Yorkshire, examining the implications for regional wetland development along the River Torne, and connections with nearby wetlands. Between 10,200 cal BP and 2300 cal BP, the floodplain experienced multiple phases of ecological change. Early in the sequence, a cut-off channel of the river began to infill with peat, while the surrounding floodplain remained relatively dry with pine forests growing nearby. After about 7000 cal BP, the local woodland had diversified and expanded to mixed deciduous tree cover. By about 6000 cal BP, widespread floodplain paludification had occurred, mainly controlled by relative sea-level rise. Floodplain expansion fostered alder fen woodland. The local Torne floodplain experienced varying levels of wetness accompanied by changes in local vegetation. Floodplain alluviation, linked to rising water tables, and probably agricultural activities within the catchment, began about 4300 cal BP, significantly earlier than previously thought for this region. When the findings of the floodplain investigations were compared with previously published insect faunal assemblages from nearby Hatfield Moors (cf. Whitehouse, 2004), Mansell et al. found that whilst some early aspects of wetland response occurred within similar timeframes between the two ecosystems, implying linked responses to regionally driven wetland development, likely linked to sea level changes, that once ombrotrophy had set in the nearby raised mire ecosystem, it effectively was de-coupled from regional water table changes and became responsive to local precipitation changes. 6. Environmental archaeology Three environmental archaeological studies employing British insect fossil evidence are presented in this issue. The first is a study of stable carbon and nitrogen isotopes from fossil beetle chitin, based on specimens from Roman-age sites in northern England by King. The results of the isotope analyses found the isotopic signatures from insect chitin to be comparable to previous studies based on stable isotopes from contemporaneous bone collagen samples in the study region. The marriage of stable isotope
Guest Editorial / Quaternary International 341 (2014) 1e5
analysis to archaeoentomology during this preliminary investigation provided new data for the study of animal husbandry and agricultural practices during the Roman Period. In particular, the insect isotope data shed light on the availability and use of agricultural crops as well as the diet and grazing environments of domesticated animals. The second British archaeological study, by Smith et al. concerned the study of dung beetles from the palaeoecological and archaeological record, in an attempt to reconstruct herd concentration and the identity of herbivores. There were two parts to the paper. The first describes the results of experimental studies based on modern sub-fossil dung beetle faunas retrieved from a range of small ponds at sites in Dunham Massey, Cheshire, and Epping Forest, London. These studies suggest that it may be possible to use the proportions of dung beetles and associated species to differentiate between high and low density grazing in the fossil record. The second part examines the insect faunas recovered from the modern dung of cattle, horses, deer and pigs, chosen to replicate, as closely as possible, Pleistocene taxa. The results suggest that it may indeed be possible to differentiate between the dung of a range of different animals, based on the numbers and kinds of dung beetles found in a fossil assemblage. The third of the British archaeological studies is by King, and concerned the environmental context for disease in Medieval York. The evidence indicates that the level of squalor declined following the Norman conquest in the 11th century, but York's waterways remained fairly foul from the 11th to 14th centuries, and this contamination carried over into residential areas of the town. During the 14th century, efforts were made to clean up human and animal waste through their removal from the town and dumping in designated areas on the outskirts. The fossil evidence shows that this move towards better hygiene brought marked results, as the number of remains of human intestinal parasites recovered from ancient latrines dropped after the 14th century. A study of Medieval environmental archaeology was carried out by Reilly on the basis of insect fossil assemblages from several sites across Dublin, Ireland. This work was also inspired by Russell Coope, who published the first such study on insect remains from Christchurch Place, Dublin, in 1981. Reilly's study focuses on the Fishamble Street site. This study used insect fossils to tease apart occupational subdivisions within houses. For instance, the superabundance of the blind beetle Aglenus brunneus in side aisles of these houses supports the archaeologist's contention that the style of construction of Viking-age houses in Dublin followed the previous Irish pattern of having raised benches. These benches were built up with the kind of brushwood, wood chips and other dry plant material that would have provided an ideal habitat for A. brunneus. The fossil insect faunas also showed the influence of the Viking European trading network on Dublin. Ponel and Andrieu-Ponel present the results of their study of insect fossils and pollen from an archaeological site in Marseille, France, dating from the 14th to 17th century AD. Both lines of evidence document the cultivation of olives and figs at the site, along with cereals. Beetle species associated with wood soaked with seawater suggest the nearby presence of a shipyard, and beetles and plants associated with salty soil suggest proximity to saltpan environments. The comparative study is instructive in that there were instances (most commonly) in which both lines of evidence agree, and others in which phytophagous insects indicated the presence of specific plants that were not recorded in the pollen spectra. Also, the insect data provided abundant, original information on both human activities and anthropogenic environments developed at the site. There are few comparable studies focusing on urban environments in this region of the Mediterranean, and as such this study highlights the importance and value
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of opportunities to investigate urban archaeological areas outside the current geographic focus of northern Europe, typical of many studies. Environmental archaeology is also the focus of a paper by Lemdahl et al., who studied Lateglacial insect assemblages from the Palaeolithic site of Slotseng in southwest Denmark. The Slotseng site represents Palaeolithic settlements dated about 15,000 to 14,000 cal BP. They recovered insect remains from a kettle hole adjacent to the archaeological site, part of a multidisciplinary project that also involved pollen, macroscopic plant remains, and vertebrate remains. In this article, the results from the insect analysis are presented. The diverse beetle assemblages indicate an open heath environment at the site, which agrees well with the reconstructions based on pollen and plant macrofossil analyses. The presence of dung beetles indicates that reindeer herds grazed in the vicinity of the site. A number of finds of the carrion beetle Thanatophilus dispar suggest that fish may have been a complementary food resource for the hunters at Slotseng. MCR reconstructions indicate arctic/subarctic climate conditions during the periods GS-2a (Pre-Bølling) and GI-1d (Older Dryas) with TMAX ca. 9e13 C and TMIN ca. -3 to 20 C. During the interstadials GI-1e (Bølling) and GI-1c (Allerød 1) TMAX values were ca. 14e16 C, but TMIN values remained similar to those during the colder periods. Another study from a late Holocene archaeological setting in south-central Sweden is presented by Hellqvist. Here, a spring deposit yielded insect remains in peat. The spring was apparently used by people as early as the Bronze Age, and continued to be used until the late Iron Age, between AD 600 and 800. The earliest beetle assemblages indicate open landscapes with grazing land, adjacent to forests. This environment is similar to the modern one at the site. The decline in beetle species diversity through time reflects the regional dominance of agriculture, and the loss of natural habitats. The environmental archaeology of late 19th and early 20th century turf buildings in Iceland is the focus of a study by Forbes and Milek. These were mainly outbuildings on Icelandic farms, used for a variety of purposes ranging from human habitation to storage areas and animal stalls. This was not a traditional archaeological study, because in some cases the farmers still living on these farms had memories of how the outbuildings had been used by previous generations. Subtle variations in the insect faunas associated with the various outbuilding uses were not sufficiently robust to be trustworthy, so the authors concluded that it would be difficult to identify a room's function solely on the basis of the entomological evidence. This kind of experimental archaeology is helping to set important boundaries on the reliable interpretation of insect fossil assemblages from various archaeological settings. 7. North American research Two North American studies are included in this issue. One, by Kuzmina et al., discusses Middle Pleistocene (MIS 7) to Holocene fossil insect assemblages from the Old Crow basin of the northern Yukon, Canada. Oddly enough, this site has a unique Russell Coope connection, as it is the only fossil insect site in arctic Canada he ever visited. The oldest organic deposits discussed here lie below the Old Crow tephra, dated at 124 ± 10 ka. The whole sequence of insect assemblages from Old Crow indicates that regional environments were dominated by dry tundra and steppe-tundra throughout the late Pleistocene. Forest insect species were relatively rare, even during the last interglaciation (MIS 5e), when a relatively wet and warm tundra environment dominated. The interglacial faunas include thermophilous species suggesting warmer than modern climate. In contrast to this, the Early
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Guest Editorial / Quaternary International 341 (2014) 1e5
Holocene faunas contained an abundance of forest insects. On the whole, the insect data suggest that the contrast between cold and warm stages in the Old Crow region was less pronounced than in other regions of Yukon and Alaska during the Pleistocene, possibly due to the persistent influence of large regional lakes. Ashworth and Nelson studied fossil beetles from the Olympia beds, Discovery Park, Seattle, Washington. Coope visited this site with Nelson in 1980, when the latter was a student at the University of Washington. This paper reports on two fossil insect assemblages with ages of 17,959 and 19,786 cal BP. The assemblages yielded abundant, diverse faunas. The habitats represented by fossils include sparsely vegetated sand and gravel bars, open muddy areas, and vegetation-shaded banks. Outside of the floodplain, the habitat was a grassland with isolated stands of coniferous trees indicated by scolytids (bark beetles). Large mammals probably used the river as a watering hole, based on the abundance of dung-feeding beetles. Most of the beetle species have modern geographic ranges which include northern Washington State, but not the temperate rainforest fauna characteristic of the Pacific Northwest. The older of the two assemblages yielded a TMAX estimate of 17 C, similar to Seattle today, however, TMIN estimates suggest winters colder than modern. The younger assemblage yielded a TMAX estimate 1e2 C colder than the older assemblage, possibly associated with the advance of the Puget Lobe. Both pollen and fossil beetle evidence indicate that the climate was dry, with only half of today's precipitation. While the palynologists have considered the MIS 2 vegetation as tundra, cold steppe, or subarctic parkland, there are no obligate tundra beetles in the assemblages, and the authors consider that the local vegetation was prairie. 8. Japanese research Two Japanese studies are presented in this issue. Hayashi and Sota studied Quaternary Donaciine beetles (Coleoptera, Chrysomelidae) from Japan, reconstructing the history of colonization and divergence patterns on the basis of fossil and molecular data. The oldest fossils of the extant species are of two endemic species, Plateumaris akiensis and Plateumaris constricticollis from the Pliocene. P. akiensis is currently confined to a single locality, whereas P. constricticollis occurs over a wide geographic range and shows marked morphological and genetic divergence. Early Pleistocene fossil Donaciines include two endemic species (Donacia japana and Donacia ozensis), and the non-endemic Donacia vulgaris. During the Middle Pleistocene Plateumaris sericea and Donacia splendens made their first appearance in Japan. Mitochondrial DNA evidence indicates that the modern lineages of the Japanese specimens date back to the Middle-Late Pleistocene in most extant species, suggesting the occurrence of colonization and/or bottleneck events during this period. P. sericea likely colonized the islands of Hokkaido and Honshu/Kyushu separately during the Middle Pleistocene, following the divergence of two different lineages during the Early Pleistocene. The other Japanese paper by Shiyake reports on a first attempt to apply the Mutual Climatic Range method of palaeoclimate reconstruction to fossil beetle assemblages in Japan. Japan enjoys a wealth of modern climate data because of its dense web of meteorological stations. Further, the modern distributions of beetles are well known in Japan. Many of the cold adapted species which were in the Japanese lowlands during glacial periods still survive in high mountain refuges. The author has devised a new method of constructing species climate envelopes, drawing convex polygons to make the method more accurate and objective. The paper presents a test of this method, using four beetle species from a stratum in the Nojiri-ko site, Nagano Prefecture that dates approximately to MIS 4
(100e53 ka). The test shows a decline of temperatures during the Late Pleistocene, with TMAX 4.5 C cooler than modern, and TMIN very similar to modern values in the study region. These results agree with previous estimates based on modern range overlaps and on reconstructions based on vegetation. 9. Database compilations Buckland contributes a paper describing the development and uses of the BUGS electronic database of fossil insect data. The database currently contains data for 1124 sites in Europe, the Middle East and North Africa. Russell Coope analysed over 154 of these assemblages. BugsCEP, The Bugs Coleopteran Ecology Package, is the latest version of the database (http://www.bugscep.com). This version of the programme includes analytical tools for quantitative habitat reconstruction and visualisation, correlation matrices, and MCR climate reconstruction. It also provides a system for entering, storing and managing palaeoentomological data. Work is underway to create an online version of the programme, implemented through the Strategic Environmental Archaeology Database (SEAD) project (http://www.sead.se). Along similar lines to the BUGS database, Kuzmina reviewed the QUINSIB database, a set of computerized insect fossil data, mainly from sites in north eastern Siberia. The QUINSIB database is far smaller than the BUGS CEP database, but it still contains over 13,400 fossils records from more than 600 fossil assemblages. It also includes modern distribution and habitat data for more than 400 species found in these assemblages (mainly beetles). As with BUGS, the principal aim of QUINSIB is to make these data available to interested students and researchers. In future, the authors hope to improve the utility of the database by adding distribution maps, references, and more ecological data. 10. Concluding thoughts The science of Quaternary Entomology that was essentially launched by Russell Coope in the late 1950s has grown by leaps and bounds in the intervening years. Coope's great enthusiasm and intellectual curiosity have inspired multiple generations of researchers, and it is highly doubtful that this branch of science would have progressed as it has, without his tremendous influence. All of us involved in the development of this special issue of Quaternary International wish to honour his memory through this publication.
Acknowledgements We would like to thank all the authors for their contributions to this issue, as well as the reviewers who dealt with the manuscripts. The managing editor, Norm Catto, did a wonderful job of steering us through some choppy editorial waters, and he provided useful guidance throughout the development of the issue.
References Coope, G.R., 1981. Report on the Coleoptera from an eleventh-century house at Christ Church Place, Dublin. In: Bekker-Nielson, H., Foote, P., Olsen, O. (Eds.), Proceedings of the Eighth Viking Congress (1977). Odense University Press, Odense, Denmark, pp. 51e56. Coope, G.R., 1986. The invasion and colonization of the North Atlantic islands: a palaeoecological solution to a biogeographic problem. Philosophical Transactions of the Royal Society of London, Series B 314, 619e635. Whitehouse, N.J., 2004. Mire ontogeny, environmental and climate change inferred from fossil beetle successions from Hatfield Moors, eastern England. The Holocene 14, 79e93.
Guest Editorial / Quaternary International 341 (2014) 1e5
Scott A. Elias* Geography Department, Royal Holloway University of London, Egham Hill, Egham, Surrey TW20 0EX, United Kingdom Nicki J. Whitehouse School of Geography, Earth and Environmental Sciences, Plymouth University, Drake Circus, Plymouth, PL4 8AA England, UK E-mail address: [email protected].
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* Corresponding author. E-mail addresses: [email protected], [email protected] (S.A. Elias).
Available online 25 July 2014
Contents lists available at ScienceDirect
Quaternary International journal homepage: www.elsevier.com/locate/quaint
Guest Editorial
G. Russell Coope: Papers honouring his life and career a b s t r a c t Keywords: Quaternary Pleistocene Holocene Insect fossils Beetles Midges
The passing of Russell Coope in 2011 brought an end to a vigorous, dynamic research career that launched the field of Quaternary entomology. This issue of Quaternary International is composed mostly of papers given in his honour in June, 2012 at Royal Holloway University of London. It comprises 21 papers that cover a wide range of topics. The reconstruction of British Pleistocene environments was arguably Coope's most important contribution to science. Three papers containing previously unpublished Middle and Late Pleistocene beetle faunas and their interpretations are included here. A discussion paper on the origins of the insect faunas of North Atlantic islands echoes another of Coope's research interests, as do two studies of late glacial climates of northwest Europe. A suite of several papers discussing the environmental archaeology of sites ranging in age from the Bronze Age to the early 20th century honour Coope's pioneering work in this field. Pleistocene research from sites in North America and Japan complete the volume, followed by descriptions of two large-scale insect fossil databases. © 2014 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
2. British Pleistocene studies
Russell Coope was the pioneer in the field of Quaternary entomology, almost single-handedly bringing the field from complete obscurity to the forefront of Quaternary science, beginning in the late 1950s. His passing in 2011 brought an end to a vigorous, dynamic research career that launched the field. He also trained a younger generation of paleoentomologists who went on to work in Britain, Continental Europe and North America. His former students and other colleagues met in June, 2012 to celebrate Russell's life and contributions to science in a symposium at Royal Holloway University of London. A number of outstanding papers were presented in this symposium, many of which are presented in this special issue of Quaternary International. One of Russell Coope's defining characteristics was his wideranging interest in all aspects of natural history, not just Quaternary insects. But even within his own field, he was instrumental in developing research in both Pleistocene and Holocene insect fossils from natural and anthropogenic deposits. He helped launch this field in North America during his participation in a scientific meeting in Ohio in the 1960s, and he worked on sites throughout northwest Europe, in the process training students from France, Denmark and Sweden. It is only fitting, therefore, that the contents of this special issue are wide ranging, both geographically and topically. The authors offer these articles in honour of his memory. Coope remained active in the field of Quaternary entomology right up to the end of his life, so some of his work remained unpublished during his lifetime. Some aspects of this work are included here.
We present some of Coope's contributions to multidisciplinary research in British Pleistocene environmental reconstructions in a set of two papers led by Harry Langford. One of these concerns Middle to Late Pleistocene environments from cold/cool stage deposits at Whittlesey, Cambridgeshire, England. Organic deposits exposed along the River Nene yielded a sequence of fossil assemblages. The study examined pollen, plant macrofossils, molluscs, beetles, ostracods, foraminifera and vertebrates. The youngest of these fossil assemblages dates to MIS 3. The other three assemblages pre-date the last interglacial (MIS 5e), with the uppermost of these three apparently representing the MIS 7e6 transition. The three assemblages older than the last interglacial all contained beetle and ostracod faunas indicating mean July temperatures (TMAX) that were 1e3 C warmer than those of the LGM, but mean January temperatures (TMIN) were 8 C colder than those estimated for the LGM. Langford et al. also reported on an MIS 7 interglacial deposit from the River Nene. This is the first fully interglacial fluvial fossil assemblage described so far for this catchment. Pollen, mollusc and insect fossil remains were studied from the deposit. The palaeobotanical and insect evidence indicate the presence of extensive deciduous woodland at the site, while the ostracod evidence suggests that saline habitats were also present locally, suggesting that sea level at the time was not much different to today. Temperature estimates based on insect and ostracod data indicate that mean July temperatures were 16e21 C, while mean January temperatures were 1 to 3 C. The modern TMAX and TMIN values fall within these boundaries.
http://dx.doi.org/10.1016/j.quaint.2014.06.056 1040-6182/© 2014 Elsevier Ltd and INQUA. All rights reserved.
2
Guest Editorial / Quaternary International 341 (2014) 1e5
Coope also analysed fossil insect assemblages of early Middle Pleistocene age from a site at Sidestrand in northeast Norfolk, England (Larkin et al.). Here a beetle fauna was identified that represents one of the most extensive cold-stage assemblages from this interval, with 59 taxa indicative of cold, continental climate. TMAX estimates for this fauna were 10e13 C, and TMIN estimates were 17 to 10 C. These temperature regimes are found today near northern treeline in the Arctic. The beetle evidence from this site contradicts the evidence derived from other climate proxies, such as the global ice volume record, which indicate less intense cold stages during the early Middle Pleistocene. However, the age of the Sidestrand cold-adapted fauna remains uncertain. An underlying Sidestrand Hall Member of the Cromer Forest-Bed Formation is thought to represent MIS 13, but this cold-stage fossil bed is separated from that stratigraphic horizon by shallow marine deposits representing at least two marine regression phases and a cold marine regression. 3. North Atlantic studies One of the most interesting puzzles for biogeographers is the question of how and when the North Atlantic islands were colonized by flora and fauna. This is a problem that fascinated Russell Coope, who wrote an article about it in 1986. Panagiotakopulu offers a hypothesis that insect colonization of the North Atlantic region was all postglacial, based on the lack of endemic species that might otherwise have survived on the Faroe Islands, Iceland and Greenland during the last glaciation. The bulk of the fossil data points towards postglacial immigration onto islands left devoid of insect life by the effects of the last glaciation. The evidence suggests that the majority of successful immigration efforts occurred through insects being carried west across the North Atlantic on ice rafts created by the disintegration of the Scandinavian ice sheet. Some of these massive blocks of ice may well have been covered by layers of soil and organic debris, providing temporary habitats for insects that either were carried away from the land on these blocks, or which landed on them when they were not too far from the shore, in attempted aerial dispersals from the west coastal regions of Norway. 4. British late glacial studies Study of the sequence of environmental changes during the late glacial (or ‘lateglacial’ in most British publications) interval was one of the most important themes in Russell Coope's research. He published 18 papers on this topic. Although the British late glacial interval is probably one of the best-studied in the world, it still sparks very lively debates. Elias and Matthews contributed to this topic by examining the thermal history of the British late glacial using the Mutual Climatic Range (MCR) method. They performed MCR analyses and used Bayesian approaches to recalibrate the radiocarbon chronologies for four important previously published beetle faunal sequences: Gransmoor, St. Bees, Llanilid, and Glannynau, but instead of generating MCR estimates for entire faunal assemblages, they split the assemblages into terrestrial and aquatic groups, and undertook separate MCR analyses of the two groups. Their working hypothesis was that the MCR estimates of TMAX derived from the aquatic species might be colder than those derived from terrestrial species, if late glacial pond waters were cooled by meltwater from ice sheets or snowbeds. Only a few instances of difference between the two sets of TMAX estimates were found, and all of these fell within the transition into the late glacial interstadial interval, during the transition from interstadial conditions into the Younger Dryas cooling, or during the Younger Dryas itself. The authors attributed the differences to the influence of meltwater from
snowbanks surrounding the study sites, and suggest that future late glacial beetle studies from northwest Europe should adopt the protocol of generating separate aquatic and terrestrial MCR estimates, for comparative purposes. The late glacial interval and the early Holocene were also the focus of an article by Brooks and Langdon, who examined fossil midge (Diptera: Chironomidae) records from northwest Europe in order to estimate summer air temperatures. They examined a large suite of faunal assemblages from 22 late glacial sites and 34 early Holocene sites, drawing isotherm maps across the study region. These maps show a strong east-west gradient during the late glacial interval, presumably due to the influence of thermohaline circulation in the regions bordering the North Atlantic. They also found a strong north-south gradient for the late glacial, likely due the persistent cooling effects of the Scandinavian ice sheet. Warming intervals, both during the late glacial interstadial and in postglacial times, were first seen in the south, then progressed northwards. Interestingly, the midge-based paleotemperature estimates for Northwest Europe were consistently cooler than those derived from beetle assemblages. This trend is consistent with the findings of Elias and Matthews that meltwater input in ponds and lakes during intervals of late lying snow may have cooled the water bodies in which both aquatic beetle and midge larvae lived. 5. Holocene studies Mansell et al. used insect fossils, pollen and plant macrofossil data to reconstruct Holocene floodplain palaeoecology of the Humberhead Levels region of South Yorkshire, examining the implications for regional wetland development along the River Torne, and connections with nearby wetlands. Between 10,200 cal BP and 2300 cal BP, the floodplain experienced multiple phases of ecological change. Early in the sequence, a cut-off channel of the river began to infill with peat, while the surrounding floodplain remained relatively dry with pine forests growing nearby. After about 7000 cal BP, the local woodland had diversified and expanded to mixed deciduous tree cover. By about 6000 cal BP, widespread floodplain paludification had occurred, mainly controlled by relative sea-level rise. Floodplain expansion fostered alder fen woodland. The local Torne floodplain experienced varying levels of wetness accompanied by changes in local vegetation. Floodplain alluviation, linked to rising water tables, and probably agricultural activities within the catchment, began about 4300 cal BP, significantly earlier than previously thought for this region. When the findings of the floodplain investigations were compared with previously published insect faunal assemblages from nearby Hatfield Moors (cf. Whitehouse, 2004), Mansell et al. found that whilst some early aspects of wetland response occurred within similar timeframes between the two ecosystems, implying linked responses to regionally driven wetland development, likely linked to sea level changes, that once ombrotrophy had set in the nearby raised mire ecosystem, it effectively was de-coupled from regional water table changes and became responsive to local precipitation changes. 6. Environmental archaeology Three environmental archaeological studies employing British insect fossil evidence are presented in this issue. The first is a study of stable carbon and nitrogen isotopes from fossil beetle chitin, based on specimens from Roman-age sites in northern England by King. The results of the isotope analyses found the isotopic signatures from insect chitin to be comparable to previous studies based on stable isotopes from contemporaneous bone collagen samples in the study region. The marriage of stable isotope
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analysis to archaeoentomology during this preliminary investigation provided new data for the study of animal husbandry and agricultural practices during the Roman Period. In particular, the insect isotope data shed light on the availability and use of agricultural crops as well as the diet and grazing environments of domesticated animals. The second British archaeological study, by Smith et al. concerned the study of dung beetles from the palaeoecological and archaeological record, in an attempt to reconstruct herd concentration and the identity of herbivores. There were two parts to the paper. The first describes the results of experimental studies based on modern sub-fossil dung beetle faunas retrieved from a range of small ponds at sites in Dunham Massey, Cheshire, and Epping Forest, London. These studies suggest that it may be possible to use the proportions of dung beetles and associated species to differentiate between high and low density grazing in the fossil record. The second part examines the insect faunas recovered from the modern dung of cattle, horses, deer and pigs, chosen to replicate, as closely as possible, Pleistocene taxa. The results suggest that it may indeed be possible to differentiate between the dung of a range of different animals, based on the numbers and kinds of dung beetles found in a fossil assemblage. The third of the British archaeological studies is by King, and concerned the environmental context for disease in Medieval York. The evidence indicates that the level of squalor declined following the Norman conquest in the 11th century, but York's waterways remained fairly foul from the 11th to 14th centuries, and this contamination carried over into residential areas of the town. During the 14th century, efforts were made to clean up human and animal waste through their removal from the town and dumping in designated areas on the outskirts. The fossil evidence shows that this move towards better hygiene brought marked results, as the number of remains of human intestinal parasites recovered from ancient latrines dropped after the 14th century. A study of Medieval environmental archaeology was carried out by Reilly on the basis of insect fossil assemblages from several sites across Dublin, Ireland. This work was also inspired by Russell Coope, who published the first such study on insect remains from Christchurch Place, Dublin, in 1981. Reilly's study focuses on the Fishamble Street site. This study used insect fossils to tease apart occupational subdivisions within houses. For instance, the superabundance of the blind beetle Aglenus brunneus in side aisles of these houses supports the archaeologist's contention that the style of construction of Viking-age houses in Dublin followed the previous Irish pattern of having raised benches. These benches were built up with the kind of brushwood, wood chips and other dry plant material that would have provided an ideal habitat for A. brunneus. The fossil insect faunas also showed the influence of the Viking European trading network on Dublin. Ponel and Andrieu-Ponel present the results of their study of insect fossils and pollen from an archaeological site in Marseille, France, dating from the 14th to 17th century AD. Both lines of evidence document the cultivation of olives and figs at the site, along with cereals. Beetle species associated with wood soaked with seawater suggest the nearby presence of a shipyard, and beetles and plants associated with salty soil suggest proximity to saltpan environments. The comparative study is instructive in that there were instances (most commonly) in which both lines of evidence agree, and others in which phytophagous insects indicated the presence of specific plants that were not recorded in the pollen spectra. Also, the insect data provided abundant, original information on both human activities and anthropogenic environments developed at the site. There are few comparable studies focusing on urban environments in this region of the Mediterranean, and as such this study highlights the importance and value
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of opportunities to investigate urban archaeological areas outside the current geographic focus of northern Europe, typical of many studies. Environmental archaeology is also the focus of a paper by Lemdahl et al., who studied Lateglacial insect assemblages from the Palaeolithic site of Slotseng in southwest Denmark. The Slotseng site represents Palaeolithic settlements dated about 15,000 to 14,000 cal BP. They recovered insect remains from a kettle hole adjacent to the archaeological site, part of a multidisciplinary project that also involved pollen, macroscopic plant remains, and vertebrate remains. In this article, the results from the insect analysis are presented. The diverse beetle assemblages indicate an open heath environment at the site, which agrees well with the reconstructions based on pollen and plant macrofossil analyses. The presence of dung beetles indicates that reindeer herds grazed in the vicinity of the site. A number of finds of the carrion beetle Thanatophilus dispar suggest that fish may have been a complementary food resource for the hunters at Slotseng. MCR reconstructions indicate arctic/subarctic climate conditions during the periods GS-2a (Pre-Bølling) and GI-1d (Older Dryas) with TMAX ca. 9e13 C and TMIN ca. -3 to 20 C. During the interstadials GI-1e (Bølling) and GI-1c (Allerød 1) TMAX values were ca. 14e16 C, but TMIN values remained similar to those during the colder periods. Another study from a late Holocene archaeological setting in south-central Sweden is presented by Hellqvist. Here, a spring deposit yielded insect remains in peat. The spring was apparently used by people as early as the Bronze Age, and continued to be used until the late Iron Age, between AD 600 and 800. The earliest beetle assemblages indicate open landscapes with grazing land, adjacent to forests. This environment is similar to the modern one at the site. The decline in beetle species diversity through time reflects the regional dominance of agriculture, and the loss of natural habitats. The environmental archaeology of late 19th and early 20th century turf buildings in Iceland is the focus of a study by Forbes and Milek. These were mainly outbuildings on Icelandic farms, used for a variety of purposes ranging from human habitation to storage areas and animal stalls. This was not a traditional archaeological study, because in some cases the farmers still living on these farms had memories of how the outbuildings had been used by previous generations. Subtle variations in the insect faunas associated with the various outbuilding uses were not sufficiently robust to be trustworthy, so the authors concluded that it would be difficult to identify a room's function solely on the basis of the entomological evidence. This kind of experimental archaeology is helping to set important boundaries on the reliable interpretation of insect fossil assemblages from various archaeological settings. 7. North American research Two North American studies are included in this issue. One, by Kuzmina et al., discusses Middle Pleistocene (MIS 7) to Holocene fossil insect assemblages from the Old Crow basin of the northern Yukon, Canada. Oddly enough, this site has a unique Russell Coope connection, as it is the only fossil insect site in arctic Canada he ever visited. The oldest organic deposits discussed here lie below the Old Crow tephra, dated at 124 ± 10 ka. The whole sequence of insect assemblages from Old Crow indicates that regional environments were dominated by dry tundra and steppe-tundra throughout the late Pleistocene. Forest insect species were relatively rare, even during the last interglaciation (MIS 5e), when a relatively wet and warm tundra environment dominated. The interglacial faunas include thermophilous species suggesting warmer than modern climate. In contrast to this, the Early
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Guest Editorial / Quaternary International 341 (2014) 1e5
Holocene faunas contained an abundance of forest insects. On the whole, the insect data suggest that the contrast between cold and warm stages in the Old Crow region was less pronounced than in other regions of Yukon and Alaska during the Pleistocene, possibly due to the persistent influence of large regional lakes. Ashworth and Nelson studied fossil beetles from the Olympia beds, Discovery Park, Seattle, Washington. Coope visited this site with Nelson in 1980, when the latter was a student at the University of Washington. This paper reports on two fossil insect assemblages with ages of 17,959 and 19,786 cal BP. The assemblages yielded abundant, diverse faunas. The habitats represented by fossils include sparsely vegetated sand and gravel bars, open muddy areas, and vegetation-shaded banks. Outside of the floodplain, the habitat was a grassland with isolated stands of coniferous trees indicated by scolytids (bark beetles). Large mammals probably used the river as a watering hole, based on the abundance of dung-feeding beetles. Most of the beetle species have modern geographic ranges which include northern Washington State, but not the temperate rainforest fauna characteristic of the Pacific Northwest. The older of the two assemblages yielded a TMAX estimate of 17 C, similar to Seattle today, however, TMIN estimates suggest winters colder than modern. The younger assemblage yielded a TMAX estimate 1e2 C colder than the older assemblage, possibly associated with the advance of the Puget Lobe. Both pollen and fossil beetle evidence indicate that the climate was dry, with only half of today's precipitation. While the palynologists have considered the MIS 2 vegetation as tundra, cold steppe, or subarctic parkland, there are no obligate tundra beetles in the assemblages, and the authors consider that the local vegetation was prairie. 8. Japanese research Two Japanese studies are presented in this issue. Hayashi and Sota studied Quaternary Donaciine beetles (Coleoptera, Chrysomelidae) from Japan, reconstructing the history of colonization and divergence patterns on the basis of fossil and molecular data. The oldest fossils of the extant species are of two endemic species, Plateumaris akiensis and Plateumaris constricticollis from the Pliocene. P. akiensis is currently confined to a single locality, whereas P. constricticollis occurs over a wide geographic range and shows marked morphological and genetic divergence. Early Pleistocene fossil Donaciines include two endemic species (Donacia japana and Donacia ozensis), and the non-endemic Donacia vulgaris. During the Middle Pleistocene Plateumaris sericea and Donacia splendens made their first appearance in Japan. Mitochondrial DNA evidence indicates that the modern lineages of the Japanese specimens date back to the Middle-Late Pleistocene in most extant species, suggesting the occurrence of colonization and/or bottleneck events during this period. P. sericea likely colonized the islands of Hokkaido and Honshu/Kyushu separately during the Middle Pleistocene, following the divergence of two different lineages during the Early Pleistocene. The other Japanese paper by Shiyake reports on a first attempt to apply the Mutual Climatic Range method of palaeoclimate reconstruction to fossil beetle assemblages in Japan. Japan enjoys a wealth of modern climate data because of its dense web of meteorological stations. Further, the modern distributions of beetles are well known in Japan. Many of the cold adapted species which were in the Japanese lowlands during glacial periods still survive in high mountain refuges. The author has devised a new method of constructing species climate envelopes, drawing convex polygons to make the method more accurate and objective. The paper presents a test of this method, using four beetle species from a stratum in the Nojiri-ko site, Nagano Prefecture that dates approximately to MIS 4
(100e53 ka). The test shows a decline of temperatures during the Late Pleistocene, with TMAX 4.5 C cooler than modern, and TMIN very similar to modern values in the study region. These results agree with previous estimates based on modern range overlaps and on reconstructions based on vegetation. 9. Database compilations Buckland contributes a paper describing the development and uses of the BUGS electronic database of fossil insect data. The database currently contains data for 1124 sites in Europe, the Middle East and North Africa. Russell Coope analysed over 154 of these assemblages. BugsCEP, The Bugs Coleopteran Ecology Package, is the latest version of the database (http://www.bugscep.com). This version of the programme includes analytical tools for quantitative habitat reconstruction and visualisation, correlation matrices, and MCR climate reconstruction. It also provides a system for entering, storing and managing palaeoentomological data. Work is underway to create an online version of the programme, implemented through the Strategic Environmental Archaeology Database (SEAD) project (http://www.sead.se). Along similar lines to the BUGS database, Kuzmina reviewed the QUINSIB database, a set of computerized insect fossil data, mainly from sites in north eastern Siberia. The QUINSIB database is far smaller than the BUGS CEP database, but it still contains over 13,400 fossils records from more than 600 fossil assemblages. It also includes modern distribution and habitat data for more than 400 species found in these assemblages (mainly beetles). As with BUGS, the principal aim of QUINSIB is to make these data available to interested students and researchers. In future, the authors hope to improve the utility of the database by adding distribution maps, references, and more ecological data. 10. Concluding thoughts The science of Quaternary Entomology that was essentially launched by Russell Coope in the late 1950s has grown by leaps and bounds in the intervening years. Coope's great enthusiasm and intellectual curiosity have inspired multiple generations of researchers, and it is highly doubtful that this branch of science would have progressed as it has, without his tremendous influence. All of us involved in the development of this special issue of Quaternary International wish to honour his memory through this publication.
Acknowledgements We would like to thank all the authors for their contributions to this issue, as well as the reviewers who dealt with the manuscripts. The managing editor, Norm Catto, did a wonderful job of steering us through some choppy editorial waters, and he provided useful guidance throughout the development of the issue.
References Coope, G.R., 1981. Report on the Coleoptera from an eleventh-century house at Christ Church Place, Dublin. In: Bekker-Nielson, H., Foote, P., Olsen, O. (Eds.), Proceedings of the Eighth Viking Congress (1977). Odense University Press, Odense, Denmark, pp. 51e56. Coope, G.R., 1986. The invasion and colonization of the North Atlantic islands: a palaeoecological solution to a biogeographic problem. Philosophical Transactions of the Royal Society of London, Series B 314, 619e635. Whitehouse, N.J., 2004. Mire ontogeny, environmental and climate change inferred from fossil beetle successions from Hatfield Moors, eastern England. The Holocene 14, 79e93.
Guest Editorial / Quaternary International 341 (2014) 1e5
Scott A. Elias* Geography Department, Royal Holloway University of London, Egham Hill, Egham, Surrey TW20 0EX, United Kingdom Nicki J. Whitehouse School of Geography, Earth and Environmental Sciences, Plymouth University, Drake Circus, Plymouth, PL4 8AA England, UK E-mail address: [email protected].
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* Corresponding author. E-mail addresses: [email protected], [email protected] (S.A. Elias).
Available online 25 July 2014