The fungi of Scottish Western oakwoods

Summary

Unfortunately the Atlantic oakwoods of Scotland have never been targeted by mycologists and information is very patchy. The best data lie in the lists of fungi from the Inner Hebridean oakwoods and it is on these records which go back to the end of the 19th century, and limited information from the mainland that a potential picture is offered. There appear to be no fungi specific to Western oakwoods and there are few differences between eastern and western oak communities in Scotland. Emphasis is placed on the need for more funding for those who can identify and recognise potential indicator species so that progress can be made.     

Atlantic oakwoods in Great Britain: Factors influencing their definition,distribution and occurrence

Summary

Atlantic oakwoods are the oceanic oak-dominated woodlands that fringe the western coast of Britain and Ireland and, due to their high humidity and rich and luxuriant lower plant assemblages, have been regarded as ‘temperate rainforest’. However, unlike the similarly iconic Caledonian pinewoods, Atlantic oakwoods have no formal designation or associated woodland classification type, which gives some uncertainty to the extent of the oakwoods habitat on its fringes. This paper examines the various elements central to the concept of an ‘Atlantic oakwood’ and discuses their occurrence in Great Britain.     

A forest habitat network for the Atlantic Oakwoods in Highland region,Scotland

Summary

The need to conserve biological diversity and ensure the future viability and integrity of Atlantic oakwoods in a fragmented landscape has led to strategies that facilitate a more holistic view of biodiversity conservation across extensive areas. The Scottish Forestry Strategy contains a major aspiration to develop forest habitat networks through the restoration and improvement of existing woodland and the expansion of new woodland. The Forest Research landscape ecology model BEETLE (Biological and Environmental Evaluation Tools for Landscape Ecology) uses a focal species approach to assess the functional connectivity of habitat within the wider landscape matrix. This model has been used to predict the current habitat network for Atlantic oakwood specialists in the Highland region of Scotland. The analysis outputs are presented with an approach to help practitioners and planners visualise the opportunities to target expansion, conversion or restoration of Atlantic oakwoods and their intrinsic biodiversity.     

Woodlands at the edge: A European perspective on the Atlantic Oakwood Plant Communities

Summary

The Atlantic Oakwoods can be seen as ‘at the edge’ in two ways. First, they lie at the extreme oceanic end of a series of Quercus robur and Q. petraea woodlands which stretches eastwards from the North Atlantic coast to the continental heartland of Europe. Second, they are at the northern limit of a range of oceanic oakwoods which extends southwards to Iberia where Q. pyrenaica is an important canopy companion. This paper outlines the main variations in trees, shrubs, herbs and cryptogams that can be seen along these two climatic gradients, so as to highlight what is truly distinctive about the Atlantic Oakwoods and how we might rightly value them.     

Argyll oakwoods: Use and ecological change, 1000 to 2000 AD — a palynological-historical investigation

Summary

This paper explores the relationship between ecological change and human use in Atlantic oakwoods during the last millennium. Information on vegetation change derived from high resolution pollen analytical studies can sometimes be satisfactorily linked with localised documentary evidence of historical woodland management. Key findings based on this approach using case studies of individual woods in Lorn are presented. Relatively diverse woods, probably containing old-growth, were transformed in the medieval period into disturbed open stands used for pasturage and local domestic wood supply. Around or prior to 1700 AD commercial management began and modes of exploitation changed so that, for a period until the late 19th or early 20th century, stands were cropped intensively. In the most recent 100 years or so, use of the woods has been relatively minor except as a grazing resource. The consequences of this history for the development of the current condition and character of Atlantic oakwood resources are discussed with reference to elements of woodland biodiversity such as arboreal species composition and Atlantic bryophyte assemblages.     

Lichens — the biodiversity value of western woodlands

Summary

The Atlantic broad-leaved woodlands of Britain are of international renown for their lichen floras. They are inhabited by 517 lichens, representing 28.3% of the total lichen flora and 73.2% of all British woodland lichens, and they are the main habitat for 165 species. Of these, 31 have a marked southern distribution and do not reach Scotland, whereas 26 species are found in Scotland, but not England or Wales. Their British Red-listed species are outnumbered by the 86 species for which Britain has International Responsibility.

Within the Atlantic broad-leaved woodlands, only 30 lichens show a preponderance for oak. With the exception of some ancient oakwoods in southern England, a high lichen biodiversity is rarely dependent on a dominance of oak in the woodland canopy, more usually it is the result of a long ecological continuity, often a varied tree and shrub composition, a varied canopy density, and good air quality. Consequently, the oak stands within former ‘industrial’ woodlands have a much lower lichen biodiversity compared with woodlands that have a history as ‘pasture woodland’ or, as with some ravine woodlands, have otherwise escaped intensive management.

The life-history of an oak tree is considered in relation to the niches it provides for lichen colonisation with time. Some management scenarios are provided with the enhancement of the lichen interest of former ‘industrial’ oakwoods as an objective.     

Overall biodiversity and the spatial patterns of atlantic oakwoods

Summary

Maintenance of biodiversity in these complex woodlands, that have been historically much altered, is based on actions promoting conservation of existing values but which may also prepare them for future changes, such as a decline in the dominance of oak in the canopy. Site condition monitoring of the 9000 ha of Atlantic oakwoods already designated indicates damage from excessive browsing, invading species and lack of deadwood. These conditions and potential hazards from climate change are being addressed by attempting to expand and interconnect existing woodlands to reduce fragmentation and increase possibilities for species to migrate.     

Regeneration in Atlantic oakwoods: has deer management had a beneficial effect?

Summary

The fragmented Atlantic oakwoods of western Europe are a habitat of conservation importance, and in the UK are threatened by browsing of young trees, invasion by exotic species and under-planting with conifers. Previous research showed that small tree saplings were abundant in Scottish oakwoods, but growth was limited by shading and frequent browsing. Within this study, we aimed to assess whether subsequent changes in deer management had resulted in increased sapling growth, and to determine whether planting of oak within lines of brash left after conifer clearance had offered long-term protection from browsing.

Pellet group counts at two sites in Glen Nant, Argyll, indicated that use by red deer, roe deer and sheep was lower than five years previously. Within one site, sapling survival was high despite continued browsing, and some saplings, particularly hazel, had gained substantial height increments. In contrast, at the second site, survival was low and the surviving saplings had managed no net growth. At a third site in Glen Etive, Argyll, partially protected by fencing, survival of oak saplings was high, and two thirds had increased in height. Planting oak saplings amongst lines of conifer brash did not offer long-term protection, but within a fenced exclosure saplings had grown substantially, despite vigorous competition from the field layer.

Deer management policies within these woods are beginning to show some success in promoting regeneration, but they will need to be continued for a long period to overcome the low growth rates of hardwood species.     

Biodiversity and ecology of the invertebrates of Atlantic Oakwoods

Summary

Woodland in general supports the richest invertebrate fauna of any habitat in Britain. The number of species depends on the structure of the wood and the age of the trees within it. The greatest biodiversity is associated with oaks although the difference between the communities found on the two oak species is not clearly understood, the Sessile Oak woods have been less well studied than their lowland equivalents, particularly so in Scotland. A number of minor habitats within the wood are important for many invertebrates and the presence of many of these may be due to past management. Every part of the oak tree, the leaves, flowers, acorns, bark and wood, supports invertebrates, herbivores, predators and parasites, and the fallen leaves and deadwood provides habitat for a complex of detritus feeders. Within the woodland the other trees, the fungi and the ground flora all have characteristic woodland invertebrates. Only a few studies, on beetles and spiders, have looked at which invertebrates are particularly associated with acidic oakwoods and there have been no specific studies of Atlantic oakwoods. It is known that there are some rare and local invertebrate species associated with western woods on ancient sites, including some recently discovered suggesting that the biodiversity of these sites and their management for invertebrates might be more important than currently thought.     

Fit is fat: winter body mass of Atlantic Puffins Fratercula arctica

Capsule: Atlantic Puffins Fratercula arctica from Scottish and Norwegian populations were significantly heavier in winter than when rearing chicks.

Aims: To compare body masses of Atlantic Puffins on their wintering grounds off the Faroe Islands with those of birds rearing chicks at colonies in Scotland and Norway.

Methods: We took standardized measurements of wing length and body mass of Atlantic Puffins during the summer chick-rearing period and on the wintering grounds near the Faroe Islands. These measurements were used to estimate seasonal changes in body mass for the two breeding populations. In three cases data were available for individuals weighed both at the colony and on the wintering grounds.

Results: On average, Atlantic Puffins breeding in Scotland and Norway increased their body mass by 20–30% between the chick-rearing period and winter. The very limited individual level data accorded well with the population level estimates.

Conclusions: Our results provide the first estimates of the order of magnitude in mass change between two key life history stages in this species. They indicate that gains in body mass between chick-rearing and winter are at least double the decline in mass previously recorded between incubation and chick-rearing. Given the Atlantic Puffin’s deteriorating conservation status, improved information on seasonal changes in body condition should help determine the underlying causes of die-offs in major wreck incidents such as those reported in recent years.     

Oak as a commercial crop in the eighteenth and nineteenth centuries

Summary

The main period of the exploitation of the Atlantic oakwoods lasted from around 1700 –1900, and was intense only from 1750–1850, though it had been preceded by a long period of anthropogenic interference of which medieval boatbuilding at least had the potential to alter the character of the woods. The Irish were pioneers when they began to search the coast from the Solway to the Great Glen for fresh supplies of oak bark for their tanning industry, and early in the 18th century this involved an attempt to buy large swathes of Argyll oakwoods, and also to create iron works at Glen Kinglas to utilise local charcoal. This failed, but by mid-century English ironmasters had followed them at Bunawe and Craleckan, works ambitious enough to draw fuel from most of the area within reach of the sea, and this led to greater care being taken of the woods involving the exclusion of stock, systematic rotational coppicing and apparently species selection in favour of oak. Management for tanbark by Scots to supply Scottish markets outpaced charcoaling before 1800, however, and affected areas like Loch Lomondside and Perthshire far from the sea, with similar consequences.

A crisis developed when the tanbark and charcoal markets collapsed, notably after 1840, but it took another generation of owners to adapt. Before the end of the century, coppice was largely abandoned, though in places where there were pyroligneous acid works or opportunities to sell spoke wood, the practice lingered on. Generally, however, stock were readmitted, and the woods converted to high forest or game coverts.     

The effects of drift and selection on latitudinal genetic variation in Scandinavian common toads (Bufo bufo) following postglacial recolonisation

Clinal variation is paramount for understanding the factors shaping genetic diversity in space and time. During the last glacial maximum, northern Europe was covered by glacial ice that rendered the region uninhabitable for most taxa. Different evolutionary processes during and after the recolonisation of this area from different glacial refugia have affected the genetic landscape of the present day European flora and fauna. In this study, we focus on the common toad (Bufo bufo) in Sweden and present evidence suggesting that these processes have resulted in two separate lineages of common toad, which colonised Sweden from two directions. Using ddRAD sequencing data for demographic modelling, structure analyses, and analysis of molecular variance (AMOVA), we provide evidence of a contact zone located between Uppland and Västerbotten in central Sweden. Genetic diversity was significantly higher in southern Sweden compared to the north, in accordance with a pattern of decreased genetic diversity with increasing distance from glacial refugia. Candidate genes under putative selection are identified through outlier detection and gene–environment association methods. We provide evidence of divergent selection related to stress response and developmental processes in these candidate genes. The colonisation of Sweden by two separate lineages may have implications for how future conservation efforts should be directed by identifying management units and putative local adaptations.Subject terms: Evolutionary genetics, Genomics, Genetic variation     

Ice-age survival of Atlantic cod: agreement between palaeoecology models and genetics

Scant scientific attention has been given to the abundance and distribution of marine biota in the face of the lower sea level, and steeper latitudinal gradient in climate, during the ice-age conditions that have dominated the past million years. Here we examine the glacial persistence of Atlantic cod (Gadus morhua) populations using two ecological-niche-models (ENM) and the first broad synthesis of multi-locus gene sequence data for this species. One ENM uses a maximum entropy approach (Maxent ); the other is a new ENM for Atlantic cod, using ecophysiological parameters based on observed reproductive events rather than adult distribution. Both the ENMs were tested for present-day conditions, then used to hindcast ranges at the last glacial maximum (LGM) ca 21kyr ago, employing climate model data. Although the LGM range of Atlantic cod was much smaller, and fragmented, both the ENMs agreed that populations should have been able to persist in suitable habitat on both sides of the Atlantic. The genetic results showed a degree of trans-Atlantic divergence consistent with genealogically continuous populations on both sides of the North Atlantic since long before the LGM, confirming the ENM results. In contrast, both the ENMs and the genetic data suggest that the Greenland G. morhua population post-dates the LGM.     

The migration of beech (Fagus sylvatica L.) up the Rhone: the Mediterranean history of a “mountain” species

New anthracological data on the beech (Fagus sylvatica L.), obtained through rescue excavations, allows the proposal of a new pattern of population by beech of the South of France since the Lateglacial. Due to its current ecology and distribution it is considered to be a mountain species in the Mediterranean area despite the presence of beech forests, usually described as glacial relicts, at low altitudes in the South of France and in the Rhone valley (Sainte-Baume, Valbonne and Grand Fays). Our results, combined with previous pollen and charcoal data, show both an early spread from glacial refuges and a late disappearance of the species. During the glacial period, the beech remained in the southernmost zones. From there, it first spread during the Preboreal towards outposts in the Rhone delta, and during the Atlantic along the rhodanian corridor. During the Subboreal, a retreat of beech towards the Pre-Alps and the Rhone delta can be traced. The Iron Age and the Classical period experienced the maximal concentration of beech along the Rhone river, while it became more rare in the North and in the South. During the Middle Ages, it first disappeared from these distal areas, and later also in the valley (around A.D. 1000), finally settling in its present mountainous distribution area (probably around the 14th century A.D.). Some crucial points of the proposed dynamics are further analysed: the paradoxical regression of beech in the valley during the Subboreal, the factors determining the expansion of beech from refuge areas during the Subatlantic and finally, the disappearance of the beech in the middle Rhone valley in rather recent times, a disappearance attributed to the strong human impact on the landscape.Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Distributional and demographic consequences of Pleistocene climate fluctuations for a marine demersal fish in the north-eastern Atlantic

Aim The Pleistocene glaciations were the most significant historical event during the evolutionary life span of most extant species. However, little is known about the consequences of these climate changes for the distribution and demography of marine animals of the north‐eastern Atlantic. The present study focuses on the phylogeographic and demographic patterns of the sand goby, Pomatoschistus minutus (Teleostei: Gobiidae), a small marine demersal fish. Location North‐eastern Atlantic, Mediterranean, Irish, North and Baltic seas. Methods Analysis was carried out by sequencing the mtDNA cytochrome b gene of sand gobies from 12 localities throughout the species’ range, and using this information in combination with published data of allozyme markers and mtDNA control region sequences. Several phylogenetic methods and a network analysis were used to explore the phylogeographic pattern. The historical demography of P. minutus was studied through a mismatch analysis and a Bayesian skyline plot. Results Reciprocal monophyly was found between a Mediterranean Sea (MS) clade and an Atlantic Ocean (AO) clade, both with a Middle Pleistocene origin. The AO Clade contains two evolutionary significant units (ESUs): the Iberian Peninsula (IB) Group and the North Atlantic (NA) Group. These two groups diverged during Middle Pleistocene glacial cycles. For the NA Group there is evidence for geographic sorting of the ancestral haplotypes with recent radiations in the Baltic Sea, Irish Sea, North Sea and Bay of Biscay. The demographic histories of the Mediterranean Clade and the two Atlantic ESUs were influenced mainly by expansions dated as occurring during the Middle Pleistocene glaciations and post‐Eem, respectively. Main conclusions The pre‐LGM (Last Glacial Maximum) subdivision signals were not erased for P. minutus during the LGM. Middle Pleistocene glaciations yielded isolated and differently evolving sets of populations. In contrast to the case for most other taxa, only the northern Atlantic group contributed to the post‐glacial recolonization. The historical demography of Mediterranean sand gobies was influenced mainly by Middle Pleistocene glaciations, in contrast to that of the Atlantic populations, which was shaped by Late Pleistocene expansions.     

Environmental confirmation of multiple ice age refugia for Pacific cod, Gadus macrocephalus

The concept of species surviving through quaternary climatic extremes by retreating to glacial refugia, and then evolving genetically during re-population movements of the following interglacial, has been in the literature for over 40 years. Recently, advances in genetic analysis have enabled this concept to be validated and theories regarding population expansions and contractions to be built. For the major Northern Hemisphere species of cod, Gadus morhua (Atlantic cod) and Gadus macrocephalus (Pacific cod), genetic analysis has suggested retreat to separate refugia on both sides of their respective ocean basins during the last glacial period. Ecosystem niche modelling has previously confirmed that environmental conditions during the last glacial were compatible with the existence of these separate refugia for Atlantic cod. Here it is shown that such modelling also confirms a reduced core glacial distribution for G. macrocephalus, but probable refugia on either side of the Pacific. Existing mitochondrial DNA analyses suggest two separate glacial populations in the northwest Pacific, which modelling confirms, with predicted separate marine refugia in the land-locked Sea of Japan basin and the Sea of Okhotsk. Existing mitochondrial DNA for the northeast Pacific populations is less conclusive regarding whether there were one or two separate refugia off this coast, and their location. Using environmental niche models this study shows the glacial NE Pacific environment could support two marine refugia, one centred in the Aleutians/Gulf of Alaska and the other off British Columbia. The intervening Cordilleran Ice Sheet, and the glacially sub-aerial and ice-free Queen Charlotte Islands shelf, is hypothesised to have constrained exchange between glacial stocks either side of the Islands. The postulated southern marine refugium is off-shore from an established terrestrial refugium, suggesting greater dependence of species and ecosystems during environmental change. An earth system approach to evolutionary change could enhance understanding of past and future ecosystems.     

Reconciling historical processes and population structure in the sooty tern Sterna fuscata

To test the influence of past vicariant events on population genetic structure of the sooty tern Sterna fuscata , we examined sequence variation in the mitochondrial control region of individuals from the Indo-Pacific and Atlantic Oceans. Our analyses indicate a rapid population expansion at a global scale during the last 100 000 years, consistent with global recolonisation during the interstade following the Pleistocene glacial maxima (125 000–175 000 years bp). We estimate islands of the Great Barrier Reef and Coral Sea were colonised no more than 16 000 years ago, most likely in association with the appearance of new breeding habitat following the final Pleistocene glacial retreat (19 000–22 000 years bp). Our results suggest that ice sheets linked to major glacial events not only impact genetic structuring in temperate seabirds, but that sea level changes in the tropics associated with these same events have also significantly impacted contemporary genetic structuring in tropical seabird species.     

Past and future climate-driven shifts in the distribution of a warm-adapted bird species,the European Roller Coracias garrulus

ABSTRACT

Capsule: The distribution range of the European Roller Coracias garrulus has undergone large changes over geological times, but although the species is warm-adapted, the human induced climate change is predicted to affect negatively the range of the currently large populations.

Aim: Information on species-specific vulnerability to climate change is crucial not only for designing interventions and setting conservation goals, but also to inform conservation decision-making. Our goal was to map climate suitability for the European Roller in the Western Palaearctic under current climate, and for past (last glacial maximum and mid-Holocene) and future (2050 and 2070) climate scenarios.

Methods: We used MaxEnt for species distribution modelling based on the reconstructed distribution map of the species.

Results: Our results suggest that during glacial periods Rollers persisted in small southern refugia, and then spread and colonized northern latitudes during the mid-Holocene. In the future, our models forecast a shift in climatically suitable range towards northern latitudes and an overall small range contraction (4.5–5.5%). Warmer temperatures will increase climate suitability in northern countries where the species is currently declining or became locally extinct. On the other hand, wide suitable areas under current climatic conditions are predicted to become unsuitable in the future (35–38% by 2050 and 2070, respectively), significantly impacting large populations such as those in Romania, Spain, Bulgaria and Hungary. French and Italian populations are identified to be future key populations for Roller conservation.

Conclusions: Our findings suggest that future climate changes will likely amplify the impacts of existing threats on the majority of large European Roller populations in Europe.     

Molecular markers provide evidence for a broad‐fronted recolonisation of the widespread calcareous grassland species Sanguisorba minor from southern and cryptic northern refugia

• Calcareous grasslands belong to the most species‐rich and endangered habitats in Europe. However, little is known about the origin of the species typically occurring in these grasslands. In this study we analysed the glacial and post‐glacial history of Sanguisorba minor, a typical plant species frequently occurring in calcareous grasslands.
• The study comprised 38 populations throughout the whole distribution range of the species across Europe. We used molecular markers (AFLP) and applied Bayesian cluster analysis as well as spatial principal components analysis (sPCA) to identify glacial refugia and post‐glacial migration routes to Central Europe.
• Our study revealed significant differences in the level of genetic variation and the occurrence of rare fragments within populations of S. minor and a distinct separation of eastern and western lineages. The analyses uncovered traditional southern but also cryptic northern refugia and point towards a broad fronted post‐glacial recolonisation.
• Based on these results we postulate that incomplete lineage sorting may have contributed to the detected pattern of genetic variation and that S. minor recolonised Central Europe post‐glacially from Iberia and northern glacial refugia in France, Belgium or Germany. Our results highlight the importance of refugial areas for the conservation of intraspecific variation in calcareous grassland species.

     

Phylogeography of Potentilla fruticosa (Rosaceae) in the Qinghai-Tibetan Plateau revisited: a reappraisal and new insights

Background: Most empirical studies to date have suggested that alpine plants in the central Qinghai-Tibetan Plateau (QTP) originated from the south-eastern QTP. However, previous phylogeographcial analysis of Potentilla fruticosa suggested that it diversified in the central QTP, which has also been assumed to have been a glacial refugium, and expanded to the north-eastern regions during the mid-Holocene period.

Aim: We reappraise this claim in the light of information acquired from sampling, haplotyping and analysing additional populations.

Methods: We sequenced the chloroplast trnT-L intergenic spacer of 346 individuals collected from 52 populations. We reconstructed phylogenetic relationships among haplotypes through Bayesian and Network analysis, and used several methods for demographic analysis.

Results: Some common and highly divergent haplotypes were distributed in the central, western and south-eastern QTP. Mismatch and other population genetic analyses applied to our data suggested that a distinct range expansion had occurred between approximately 30,000 and 325,000 years ago (kya).

Conclusions: Besides the possible postglacial expansion in the north-eastern QTP, our results further suggested that the range of the species radically expanded across the whole Plateau before the last glacial maximum (around 18 kya) and the south-eastern, western and central QTP regions had together provided important refugia during recent glacial stages. These findings, in contrast to the previous conclusions, highlight the importance of adequate sampling strategies in phylogeographical studies.