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Marine Biodiversity and Climate Change
Assessing and Predicting the Influence of Climatic
Change Using Intertidal Rocky Shore Biota
Final Report for United Kingdom Funders
N. Mieszkowska*, R. Leaper*, P. Moore*, M. A. Kendall†, M. T. Burrows#, D. Lear*, E. Poloczanska#, K.
Hiscock*, P.S. Moschella*, R.C. Thompson¶, R.J. Herbert, D. Laffoley§, J. Baxter‡, A. J. Southward* & S. J.
Hawkins*
The Marine Biological Association of the U.K.*
Plymouth Marine Laboratory†
The Scottish Association for Marine Sciences#
University of Plymouth¶
Medina Valley Centre, Isle of Wight & Bournemouth University
English Nature§
Scottish Natural Heritage‡
December 2005
MarClim Research undertaken by: MarClim UK was funded by:
Marine Biological Association Occasional Publications No. 20
2
ISSN 0260-2784
MarClim was a multi-partner British and Irish project led by the Marine Biological Association of the U.K. in collaboration with the Plymouth
Marine Laboratory, The Scottish Association for Marine Science, The University of Plymouth, University College Cork and a broader network of
collaborators. It was funded by the following organisations: Countryside Council for Wales, The Department for Environment Food and Rural
Affairs, English Nature, Environment Agency, Joint Nature Conservation Committee, Scottish Executive, Scottish Natural Heritage, The Crown
Estates, States of Jersey and WWF, under the umbrella of the UK Climate Impacts Programme (UKCIP). This report summarises the findings
of the work undertaken by the British team. A separate report summarising the work conducted by University College Cork (MarClim Ireland) is
available at www.mba.ac.uk/marclim. Additional peer and non-peer reviewed publications associated with this project can be obtained from the
website and a list of both as well as selected presentations is contained in Appendix 1. MarClim Ireland was funded through a post-doctoral
fellowship (PDOC/01/006) October 2002-2005 with the support of the Marine Institute and the Marine RTDI Measure, Productive Sector
Operational Programme, National Development Plan 2000–2006.
Contents
1. Executive Summary 2
2. Aims and Activities Undertaken by the MarClim Project 5
3. A Changing Climate 8
4. Responses of Marine Biodiversity to Climate Change 10
5. The Basis of the MarClim Project: Historical Data 13
6. The Basis of the MarClim Project: Resurvey Data 16
7. Main Findings: Changes in the Geographic Distribution of
Intertidal Species in Response to Climate Change and the
Biological Mechanisms Underlying these Changes
19
8. Prediction of Future Changes: Modelling Species Populations 28
9. Prediction of Future Changes: Modelling Species
Distributions
32
10. Conclusions and Project Legacy 35
11. The Implications of Climate Change for Marine Stewardship
Policy
38
12. Literature Cited 40
Appendix 1 ACFOR Abundance Categories 43
Appendix 2. MarClim Project Outputs 2001-2005 44
Appendix 3. MarClim Project Deliverables 50
2
1. Executive Summary
In the last 60 years climate change has altered the distribution
and abundance of many seashore species. Below is a
summary of the findings of this project.
The MarClim project was a four year multi-partner funded project created to investigate the effects of climatic
warming on marine biodiversity. In particular the project aimed to use intertidal species, whose abundances
had been shown to fluctuate with changes in climatic conditions, as indicator species of likely responses of
species not only on rocky shores, but also those found offshore. The project used historic time series data,
from in some cases the 1950s onwards, and contemporary data collected as part of the MarClim project
(2001-2005), to provide evidence of changes in the abundance, range and population structure of intertidal
species and relate these changes to recent rapid climatic warming. In particular quantitative counts of
barnacles, limpets and trochids were made as well as semi-quantitative surveys of up to 56 intertidal taxa.
Historic and contemporary data informed experiments to understand the mechanisms behind these changes
and models to predict future species ranges and abundances.
Main Findings
Range extensions have occurred at the northern limits of the geographical distributions of typically
southern, warm water species Osilinus lineatus (toothed topshell), Gibbula umbilicalis (flat topshell),
Chthamalus montagui (Montagu’s stellate barnacle), Chthamalus stellatus (Poli’s stellate barnacle)
and Balanus perforatus (acorn barnacle) since the mid-1980s in Wales, Northern Ireland and
Scotland, including greater penetration around the north of Scotland into the colder North Sea
(Section 7).
Eastward range extensions of the southern species Osilinus lineatus (toothed topshell), Gibbula
umbilicalis (flat topshell), Patella ulyssiponensis (china limpet), Patella depressa (black-footed
limpet), Melarhaphe neritoides (small periwinkle), Actinia fragacea (strawberry anemone) and
Balanus perforatus (acorn barnacle) have also occurred since the mid-1980s in the English Channel
beyond previous biogeographic boundaries (Section 7).
The northern species Alaria esculenta (dabberlocks) and Tectura testudinalis (common tortoiseshell
limpet) have shown small retractions in their southern distributional limits and declines in abundance
at populations close to these range edges, but the rate of recession is not as fast as the rate of
advancement in southern species (Section 7).
Synchronous increases in abundance have been recorded in populations of southern topshells
throughout Britain and northern France since the mid-1980s. These increases are an order of
magnitude greater than the inter-annual variation detected, increasing the confidence that these are
3
observations of decadal-scale change rather than the result of anomalous years, providing support
that these increases in abundance are climate-related (Section 7).
Annual reproductive cycles of the southern/lusitanian trochids are commencing earlier in response to
milder winters and warmer springs, coupled with increased survival of newly settled recruits exposed
to milder, shorter winters on the shore (Section 7).
The annual reproductive cycles of the southern/lusitanian limpet Patella depressa are starting earlier
and lasting longer in south-west Britain. In contrast, less than 20% of the population of the
northern/boreal limpet, Patella vulgata, are reaching gonad development stages at which spawning
can occur on some shores in south-west Britain (Section 7).
Fluctuations of the northern barnacle Semibalanus balanoides and the southern Chthamalus spp.
have been related to climate change, using historical data collected by Southward and advanced
statistical methods. These show that there is a direct negative effect of warm springs on survival of
Semibalanus balanoides which via release from competition has an indirect positive effect on
Chthamalus. These data have been used for hindcast and forecast modelling using UKCIP climate
scenarios. In particular these models have been able to incorporate characteristics such as species
mortality, larval supply and competitive interactions to create more biologically realistic predictions of
species responses to climate change (Section 8).
Models using the extensive broadscale resurvey data have been created for all MarClim indicator
species to predict changes in their abundance and distribution in response to wave action and sea
surface temperature regimes forecast by UKCIP (www.mba.ac.uk/marclim) (Section 9).
Conclusions
The MarClim project has provided strong evidence that recent rapid climate change has resulted in
changes in the abundance, population structure and biogeographic ranges of a number of intertidal
indicator species (Hawkins et al. 2003, Herbert et al. 2003, Kendall et al. 2004, Mieszkowska 2005,
Moore 2005, Southward et al. 2005, Mieszkowska et al. 2006, Mieszkowska et al. in prep), mirroring
changes offshore (Beaugrand et al. 2002, Genner et al. 2004, Richardson and Schoeman 2004, Sims et
al. 2004, Coombs et al. 2005, Perry et al. 2005, Southward et al. 2005). Experiments have shown that
many of the changes in southern/lusitanian species have occurred as a result of increased reproductive
output and juvenile survival in response to increased warming. In the case of the northern/boreal limpet,
P. vulgata, it would appear that decreases in its abundance may be linked to a decrease in this species
reproductive output, particularly on shores in south-west Britain. Evidence suggests that species range
expansion in response to climatic warming is occurring quicker in marine systems (plankton, fish, as well
as intertidal species) than terrestrial systems.
4
Project Legacy
The MarClim project has fulfilled its aims and in doing so collated extensive long-term time series data
on the abundance and distribution of intertidal indicator species around the British Isles. During the
project 20 peer-reviewed manuscripts have been published or have been submitted for publication to
date (Appendix 2). Many more manuscripts are expected to be published as a result of the data collated
and collected as part of the MarClim project. MarClim also disseminated its work by giving over 40
presentations to scientists, policy makers and the general public (Appendix 2).
The MarClim project has collated, undertaken extensive quality assurance measures and created a
database of historical quantitative counts for barnacles, limpets and trochids as well as semi-quantitative
broadscale data from 4400 sampling occasions at over 1000 locations. During the MarClim project 800
site visits were made to over 470 locations in the UK. An electronic database has been created, to
incorporate both the historic and contemporary surveys, to enable long-term availability of the data. All
data collated and collected by the MarClim project will be made available via the National Biodiversity
Network gateway or by contacting the Marine Environmental Change Network (MECN) co-ordinator
(www.mba.ac.uk/MECN).
Eighty sites for which there are long-term records have been designated to form a future monitoring
network as part of a wider British marine monitoring programme. The Countryside Council for Wales
(CCW) have incorporated MarClim monitoring sites and protocols into their annual marine intertidal
survey programme. Future monitoring of rocky shore climate indicator species will continue under the
auspices of the Marine Environmental Change Network (MECN).
5
2. Aims and activities undertaken by the MarClim project
At the outset of the project the main aims of MarClim were:
To use existing historical information and collect new data on intertidal indicator species from the last
50-100 years to develop and test hypotheses on the impact of climatic change on marine biodiversity
in Britain and Ireland.
To forecast future marine community changes on the basis of the Met Office’s Hadley Centre climate
change models and the United Kingdom Climate Impacts Partnership’s climate change scenarios. The
broad range of species known or thought likely to be temperature sensitive were covered.
To establish low-cost, fit-for-purpose, methodologies and networks to provide subsequent regular
updates and track how climate influences the marine biodiversity of Britain and Ireland.
To provide general contextual time series data to support reporting on the success or otherwise of
marine aspects of Biodiversity Action Plans, European initiatives including the Habitats, Birds and
Water Framework Directives, and management and monitoring of marine activities and resources,
including fisheries and Special Areas of Conservation.
To evaluate whether the climate indicator species used in this work have a wider contribution to make
as part of the sustainability indicators that are needed to underpin the UK sustainable development
strategy.
To disseminate the results widely, and accordingly elucidate the known impact that climate has had on
marine biodiversity over the last 100 years, and may have in the future.
To provide a basis for the development of a proposal for European Commission funding to establish a
pan-European network with related aims.
To assess and report on the likely consequences of the predicted changes in response to climate for
society, for commercial and non-commercial users of the marine environment and the policies and
frameworks that conserve, manage and protect marine biodiversity. To assess whether any more
serious impacts can be ameliorated or mitigated.
MarClim was organised in four phases:
Phase I (April 2001 – August 2001):
The project specification was refined and end-user liaison established via the MarClim Advisory
6
Group and discussions with other relevant programmes (MONARCH, Tyndall Centre for Climate
Change Research).
Electronic archiving of MBA and other historical datasets, particularly for fish was started and led to
various subsequent publications (Sims et al. 2001, Hawkins et al. 2002, Hawkins et al. 2003, Genner
et al. 2004, Sims et al. 2004, Coombs et al. 2005).
Conference presentations were given and workshops attended to introduce the MarClim project to
the research and policy end-user communities.
Phase II (September 2001 – March 2002)
Relevant intertidal datasets were identified and access granted from their stewards, leading to a
central archive held by MarClim (Crisp & Southward broadscale and time-series data, Lewis &
Kendall trochid data, Shetland data, Coastal Surveillance Unit intertidal data and Herbert south coast
of England data).
These datasets were archived and appropriate databases designed.
Fieldwork methods were trialled and a set of standard protocols for resurveys produced
(www.mba.ac.uk/marclim).
Research staff were recruited to the project. All staff were trained and survey techniques cross-
calibrated with Kendall, Hawkins and Southward, who had collected much of the historical data using
the field survey protocols.
Initial analyses of historical data were made.
Phase III (April 2002 – March 2004)
Historical data were analysed to inform re-survey (Herbert et al. 2003, Mieszkowska et al. 2006).
Quantitative surveys of barnacles, limpets and trochids and semi-quantitative broadscale surveys of
up to 56 indicator species were made at over 470 locations in Britain for which historical data were
available.
Annual quantitative barnacle counts of northern warm-water and southern cold-water species started
in the 1950s were continued at over 20 sites in south-west England.
Time-series of quantitative trochid data were restarted at over 30 sites in England, Wales, Scotland
and northern France.
Annual quantitative counts of limpets started in 1980 were continued at 20 sites in south and south-
west Britain.
Initiation of modelling distributions in relation to temperature and local wave action, and population
dynamics of key indicator species (co-supported by NERC small grant to SAMS).
Further publications were submitted (Herbert et al. 2003, Hiscock et al. 2004, Kendall et al. 2004,
Mieszkowska et al. 2006)
7
Phase IV (April 2004 – including extension to October 2005)
Field surveys were concluded and the monitoring network continued in south-west Britain.
An electronic archive of historical and resurvey data was completed and re-survey data entered into
Marine Recorder.
Long-term climate change monitoring methodologies and a network were designed for future
implementation.
Models forecasting likely changes in populations of key species and distributions of all MarClim
species under scenarios of climate change for the 21st century were completed and trialled.
Two Ph.Ds associated with the MarClim project were completed (Mieszkowska 2005, Moore 2005)
MarClim was subsumed with the Marine Environmental Change Network (MECN) in autumn 2005.
Summary report was written and several publications completed and submitted (Simkanin et al.
2005, Southward et al. 2005, Mieszkowska et al. in review, Moore et al. in review, Poloczanska et al.
in review)
A major policy paper was completed by the Advisory Group (Laffoley et al. 2005)
Outputs of MarClim were delivered to wider European audiences via invited participation in the
European Science Foundation Marine Board Working Group on Climate Change and the European
Platform on Biodiversity Strategy Research meeting hosted by Defra and the Scottish Executive as
part of the U.K. Presidency of the E.U.
8
3. A changing climate
There is now compelling evidence to suggest that temperature
increases over both land and sea in the northern hemisphere
during the 20th century have been greater than during any other
century in the last 1000 years.
It is now widely accepted that the planet is currently experiencing a period of rapid climate change, primarily
driven by human activities (Oreskes 2004). The global average surface temperature has increased by 0.7C
during the 20th century (Hulme et al. 2002). Globally, nine out of the ten warmest years on record were
recorded in the decade 1990-2000 and 2003 was the warmest year since instrumental records began in 1860
(Climatic Research Unit news release 16th December 2004). Sea surface temperatures (SST) around the
British coastline have increased, with some areas exceeding the global average rise. Data for SST show that
in the western English Channel there has been a 1C rise in SST since 1990, greater than any other change
recorded over the past 100 years (Hawkins et al. 2003). Similar changes have been recorded in the eastern
English Channel (Woehrling et al. 2005).
It is difficult to predict how the climate will alter over the coming decades because of uncertainties over the
rate of future greenhouse gas emissions and the response of climate to these emissions. Irrespective of the
timescale and magnitude of response, some anthropogenically driven climate change is now inevitable.
Global climate models (GCMs) predict an acceleration of the current warming trend during the first half of the
21st century as a response to continued anthropogenic emissions; thus the earth is expected to become
warmer than at any period during the past 40 million years (Houghton et al. 2001). The UK Climate Impacts
Programme (UKCIP) in conjunction with the Tyndall Centre for Climate Change Research has produced
scenarios of climate change for the UK based on climate modelling work undertaken by the Hadley Centre for
Climate Protection and Research at the Meteorological Office. This modelling is based on four different sets
of assumptions – or “storylines” – about the key drivers of greenhouse gas emissions and other climate-
altering pollutants. The UKCIP02 report (Hulme et al. 2002) presents four corresponding scenarios of future
climate change for the UK based on: Low, Medium-Low, Medium-High and High-Emissions, for three thirty-
year time slices (centred on the 2020s, 2050s and 2080s; Fig.1). The baseline convention for these scenarios
is the 30-year period 1961-1990 and scenarios are presented relative to this period (Hulme et al. 2002). The
scenarios provide a set of four alternative climate futures for the UK based on our understanding of the
science of climate change. These include the following:
An increase in annual average SST is expected under all four emissions scenarios over the next 100
years.
Southern North Sea and English Channel SST may increase by between 1.5°C under the Low Emissions
scenario and 3°C under the High Emissions scenario.
Irish Sea SST will increase between 0.5°C (Low Emissions) and 2.5°C (High Emissions).
North-east Atlantic SST will rise between 0.5°C (Low Emissions) and 1.5°C (High Emissions).
Increases in daily mean wind speeds of between 2% (Low Emissions) to 8% (High Emissions) in the
southern North Sea and English Channel during winter and spring are expected by the 2080s.
9
Summer and autumn wind speeds are expected to decrease by up to 10% off western Britain by 2080s
under the High Emissions scenario.
Fig. 1. Changes in average annual sea surface temperature (°C) by the 2020s, 2050s and 2080s, with respect to the model-simulated
1961-90 average. UKCIP02 Climate Change Scenarios (funded by DEFRA, produced by Tyndall and Hadley Centres for UKCIP).
10
4. Responses of Marine Biodiversity to Climate Change
The MarClim project has assessed the influence of climatic change on
rocky shore biota over the last 60 years and forecasts species
distributions into the future.
There is increasing concern regarding the impacts of climate change for the conservation and management of
marine biodiversity (Fields et al. 1993). As climate warming continues a general pole-ward shift in species
ranges is expected as species respond to the alteration of suitable ‘climate space’ they can inhabit (Graham
and Grimm 1990, Fields et al. 1993, Southward et al. 1995, Parmesan 1996, Sagarin et al. 1999). Species
are likely to respond to rapid temperature increases at different rates due to differences in their metabolism,
physiological processes and behaviour (Inouye et al. 2000, Réale et al. 2003, Sims et al. 2004), which will
influence adult growth and survival, reproductive output, phenology and recruitment success (Lewis 1996,
Walther et al. 2002, Herbert et al. 2003, Sims et al. 2004). Contractions and expansions of geographic range
edges will lead to species both being lost from and introduced to assemblages. Such changes will in turn
influence the outcomes of species interactions such as competition, facilitation and predation, ultimately
altering the structure of communities and marine ecosystem processes (Davis et al. 1998, Bertness et al.
1999, Case et al. 2005, Helmuth et al. 2005, Parmesan et al. 2005). Climate change could also cause
widespread ‘local extinction’ of species which could lead to global extinctions in those species that are unable
to adapt or respond to fluctuations in their physical environment (Grabherr et al. 1994, Thomas et al. 2004).
Thus the effects of climatic variability on the distributions of plants and animals must be measured in order to
understand and ultimately forecast changes in marine ecosystems.
As the most readily accessible marine habitat in Britain and Ireland, rocky shores have been the focus of
recording species distributions (Stephenson and Stephenson 1972, Lewis 1986, Little and Kitching 1996,
Raffaelli and Hawkins 1999), starting with the Victorian passion for the seashore. More formal charting of
geographic distributions started in the first half of the 20th century (Moore 1936, Moore and Kitching 1939)
with particularly valuable surveys being made in the 1950s (Crisp 1950, Southward 1950, Southward 1951,
Southward and Crisp 1952, Southward and Crisp 1954a, Southward and Crisp 1956, Crisp and Southward
1958, Crisp 1964b). Most species are sessile or sedentary and can be surveyed non-destructively.
Consequently, some of the best long-term data sets have been acquired for rocky shores – many in Britain
and Ireland (Crisp and Chipperfield 1948, Crisp 1950, Southward and Crisp 1952, Southward and Crisp
1954a, 1954b, Southward and Crisp 1956, Crisp and Southward 1958, Southward 1963, Crisp 1964,
Southward 1967, 1991, Southward et al. 1995). Furthermore, the biology of species and ecological
interactions are well known from both laboratory and field manipulative experiments, aiding the interpretation
of past change and future forecasts. Britain and Ireland straddle a biogeographic boundary between cold
‘northern’ boreal waters and warmer ‘southern’ lusitanian waters (Forbes 1858, Lewis 1964; Fig. 2). As a
consequence many intertidal species are either at the northern or southern edge of their biogeographic
ranges and will be particularly susceptible to changes in the climate (Southward and Crisp 1954a, Southward
1963, Southward et al. 1995, Lewis 1996). Thus the rocky intertidal of Britain and Ireland provides an ideal
system for studying the effects of climate in terms of alterations of geographic distribution of species and the
11
mechanisms driving these changes (Southward and Crisp 1954a, Southward and Crisp 1956, Southward
1967, 1991, Southward et al. 1995, Genner et al. 2004a).
Fig. 2. The biogeographical characteristics of the coast of the British Isles, including the range limits of some species. Redrawn from
Forbes 1858 and recently published by Hiscock et al. 2004 in Aquatic Conservation. Anglesey is absent from the map as it was in the
original publication. Acaema testudinalis is now Tectura testudinalis (a limpet); Cytherea chione is now Callista chione (a bivalve
mollusc); Echinus lividus is now Paracentrotus lividus (purple sea urchin); Fusus norvegicus is now Volutopsis norwegicus (a snail);
Haliotis is Haliotis tuberculata (the ormer); Rhynconella psittacea is now Hemithiris psttacea (a snail); Trichotropis borealis (a snail)
retains the same name; Echinus neglectus is now Strongylocentrotus droebachiensis (a sea urchin).
The MarClim project has investigated the following predictions on the current and future responses of
intertidal rocky shore fauna and flora to changes in environmental temperature regimes in Britain:
(1) Northern species will retreat northwards and their abundance will decline; such changes are likely be
driven by a reduction in reproductive output and/or decreased juvenile or adult survival during hotter
summer periods.
(2) Southern species will expand their range northwards and their abundance will increase. The
mechanisms underlying these responses are likely to be an increase in reproductive output and/or
juvenile survival during warmer summer periods and milder winters. The extent to which range
12
extensions can occur will also be dependent on length of larval life stages and presence of rocky
shores or artificial substrate beyond existing range edges.
(3) Biological interactions including competition, facilitation and predation will modulate the responses of
southern and northern species with implications for community structure and ecosystem functioning.
(4) Changes will be greater than in the last warm period prior to the cold winter of 1962/63.
These predictions have been tested using long-term data series on the abundance and distribution of rocky
intertidal species in Britain and Ireland detailed in Section 5, coupled with broadscale resurveys of selected
species (Table 1) at key sites between 2001 and 2005 (Fig. 5a). The period for which ecological information
exists spans the relatively warm 1950s, the severe winter of 1962-63 and a cool period from the 1970s to the
mid 1980s (Fig. 3). Experiments investigating the effects of warming on species reproduction and recruitment
of the lusitanian topshells Osilinus (Monodonta) lineatus and Gibbula umbilicalis have been undertaken to
investigate the mechanisms influencing these species increases in range and abundance (Mieszkowska
2005). In parallel, models were constructed on interactions between northern and southern indicator species,
focussing on barnacles and validated by comparisons with long-term time series (Poloczanska et al. in review;
Section 8). Models were also constructed to predict past and current distributions of indicator species and
validated against MarClim archived and contemporary data. These models were constructed using UKCIP
climate scenarios (Hulme et al. 2002) to assess the future distribution of key intertidal species, and the results
are detailed in Section 9.
50 - 51o N
11
11.5
12
12.5
13
13.5
1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000
Sea surface temperature ( oC)
Fig 3. Annual mean sea surface temperature trends from square 50º to 51ºN, 4º to 5ºW (Hadley Centre for Climate Prediction and
Research). Bold line is a five year running mean.
,
4 - 5º W
13
5. The Basis of the MarClim Project: Historic Data
A long history of records from the 1950s to the present day
exists for rocky intertidal species distributions.
Broadscale Surveys
Following in the footsteps of the pioneering long-term observations made on both sides of the English
Channel (Fischer-Piette 1933, 1936), the subsequent broadscale biogeographic surveys of rocky shores in
Britain and Ireland were initiated in the late 1940s and early 1950s. These surveys began with detailed
descriptions on the distribution of barnacles (Crisp and Chipperfield 1948, Crisp 1950, Southward 1950,
Southward 1951, Southward and Crisp 1952, Southward and Crisp 1954a), but were rapidly extended to
include many common intertidal organisms, so that the most important of the various possible abiotic and
biotic factors controlling distributions would be revealed (Southward and Crisp 1954b, Crisp and Southward
1958).
The broadscale surveys of French, British and Irish coasts (Fig. 4a for sites in the U.K.), at first focused on the
English Channel (Crisp and Southward 1958, Crisp and Fischer-Piette 1959) and around Ireland (Southward
and Crisp 1954b). The original aims of the surveys were threefold: (1) to ascertain, as precisely as possible in
the field, the environmental requirements of the species chosen; (2) to relate the distribution of each species
directly to the physical environment and by looking for a common basis in the various distributions find the
major factors responsible, and (3) to provide a reference against which any future changes in distribution
might be shown.
During these surveys Denis Crisp and Alan Southward developed a system of categorical abundance
estimation for key species that remains in wide use today (Southward and Crisp 1954b). The ACFOR
(Abundant, Common, Frequent, Occasional and Rare) abundance scales permitted very rapid assessment of
many tens of species in relatively short visits to survey sites. Categories ranged from ‘Not found’ through
‘Rare’ to ‘Abundant’: each class having clearly defined species-specific abundance categories on a semi-
logarithmic scale (Appendix 1). A number of survey sites in south and south-west England were re-surveyed
on a regular basis using the ACFOR abundance scale by Southward until 1987. The surveys were also
extended to other areas of Britain; in particular to sites in Scotland and Wales. Crisp and Southward
continued to visit selected sites intermittently until 1987 (Crisp et al. 1981), and by 1996 only a very small
subset of the original sites were still being surveyed (Southward 1991). It was not until 1997 that more regular
surveys were re-started in south-west Britain by Hawkins, primarily to continue the barnacle time series
detailed below.
Quantitative Barnacle Time Series
Detailed surveys of the distribution of barnacles were made in Scotland, Wales and south-west England (Fig.
4b; Southward 1950, Southward 1951, Southward and Crisp 1952, Southward and Crisp 1954a, Southward
and Crisp 1956). A joint survey between Denis Crisp and Alan Southward was then developed along two
lines. (1) Transect surveys at four sites close to Plymouth previously worked by Hilary Moore in 1934. (2)
14
More-or-less annual (non-transect) surveys at sites in the south-west, with less regular surveys conducted in
the rest of Britain and Ireland. The surveys around Plymouth were continued until the mid 1980s, with one
site continued until 1998.
Limpet Surveys
Steve Hawkins made quantitative surveys on the boreal limpet Patella vulgata and lusitanian limpet Patella
depressa in Wales and south-west Britain (Fig. 4c). A resurvey of all the Crisp & Southward (Crisp and
Southward 1958, unpub) limpet sites in England and Wales were made between 1980 and 1984, with
intermittent surveys of a small selection of sites from the mid 1980s onwards.
Coastal Surveillance in the 1970s
Extensive studies of the lusitanian topshell species Gibbula umbilicalis and Osilinus lineatus were initiated in
the 1970s as part of the work by the NERC Rocky Shore Surveillance Group under the leadership of Jack
Lewis (Fig. 4c; Kendall and Lewis 1986, Kendall 1987). Data collected by this unit was archived by Kendall
following its closure in the mid 1980s. In situ counts of topshells were made using replicated timed
collections, amongst boulders, underlying gravel and bedrock where juvenile abundance was estimated to be
greatest. The basal diameter of each individual in the sample was recorded and the animals returned to the
shore.
During the 1960s and 1970s additional long term programmes were prompted by concerns about pollution,
especially in relation to the growing oil industry. Thus time series were started at Shetland, Orkney, north
Scotland, north-east England and mid Wales. There was a realisation that background, natural variability
needed to be measured (Lewis 1976, Lewis 1996). MarClim also collated details of these datasets and is the
custodian of data from Scotland and north Wales collected by the Coastal Surveillance Unit at Menai Bridge
between 1973 and 1985 (data also collected from 1993-1994 Holt et al. 1995). A paper based on the
Shetland data collected by the Shetland Oil Terminal Environmental Advisory Group was published showing
spatial concordance in changes of abundance in a broad suite of species, including barnacles (Burrows et al.
2002). This work showed how barnacles integrate processes over a wide spatial scale.
15
Fig. 4. Location of sites in the United Kingdom for which historic data exists for (a) broadscale semi-quantitative surveys (b) quantitative
barnacle surveys (c) quantitative limpet surveys on the proportion of the southern limpet Patella depressa to the northern limpet Patella
vulgata and (d) topshell population data. Sources: Work by Crisp, Southward, Hawkins, Kendall, Williamson and Lewis
(
a
)
Broadscale surve
y
s (b) Quantitative barnacle surveys
(
c
)
Quantitative lim
p
et surve
y
s (d) Quantitative topshell surveys
16
6. The Basis of the MarClim Project: Resurvey Data
Historical baselines have been continued under MarClim by
way of a large-scale biogeographic resurvey of Britain.
Over a four-year period the biogeographic resurvey of intertidal species for which long-term data exist has
been coordinated by two separately funded MarClim programmes. Surveys in Britain, the Isle of Man and
Channel Islands have been made by the Marine Biological Association (MBA), in conjunction with the Scottish
Association for Marine Science (SAMS), Plymouth Marine Laboratory (PML) and the University of Plymouth
(UoP), assisted by selected agency staff. University College Cork (UCC) has surveyed sites in the Republic
of Ireland (MarClim Ireland). A joint survey by UCC and the MBA in conjunction with the Department for the
Environment (Northern Ireland) was also made in Northern Ireland to allow researchers to cross-calibrate data
collection protocols.
The primary objective of both survey programmes was to resurvey locations for which past data were held
using the original protocols. The MarClim resurvey included 33 of the species examined in historical
broadscale surveys and added a further 23 species thought to be equally sensitive to climate variability in
Britain (Table 1). The aims were: (1) to resurvey sites where historical broadscale surveys had been made; (2)
to resurvey sites where regular quantitative surveys had occurred to form time-series; (3) to survey additional
sites close to and beyond historical recorded range edges and (4) to include sites within or adjacent to areas
of designated conservation status. All the resurvey sites were restricted to moderately-exposed and exposed
shores of natural substratum except for surveys conducted beyond the recorded range edge of species,
where less-exposed shores and artificial substrata were examined. The current abundances of all 56 species
were mapped at sites around the British coastline (Fig. 5a to see sites sampled). Based on past records,
quantitative surveys of barnacles (northern Semibalanus balanoides, southern Chthamalus montagui and
Chthamalus stellatus, and the invasive Elminius modestus; Fig. 5b), limpets (relative proportions of northern
Patella vulgata and southern Patella depressa; Fig. 5c) and topshells (southern Osilinus lineatus and Gibbula
umbilicalis; Fig. 5d) were carried out, whilst MarClim Ireland restricted surveys to barnacles and trochids only,
due to the absence of P. depressa in Ireland. In addition, both survey programmes increased the replication
level for quantitative counts used in the historical surveys under the resurvey, to facilitate both modern
statistical analyses and to provide a more rigorous baseline for any future comparison. By adopting the same
protocols of data collection at locations for which past data are held, MarClim has produced new, but directly
comparable, baselines of the distribution and abundance of rocky shore biota in Britain and Ireland (requests
for data can be made by contacting the Marine Environmental Change Network (MECN) co-ordinator at
www.mba.ac.uk/MECN).
Additional surveys were also made by both teams: MarClim Ireland surveyed a number of locations in addition
to those with past records (semi-enclosed bays, sea lochs and harbours) in order to assess the status of E.
modestus (Allen et al. in press). MarClim (SAMS team) quantitatively surveyed a range of locations in
western Scotland to examine the effects of the level of limpet grazing on patterns of community structure on
both large-scales and in relation to local conditions of wave exposure. The SAMS team supplemented the
survey protocols with digital photography of 0.5x0.5m areas at two shore levels, measuring sizes and
densities of limpets in each area along with estimates of cover of barnacles, mussels and macroalgae, and
17
counts of other gastropods. Both programmes also trialled digital photography as a modern survey technique
for rapidly collecting replicated quantitative data on barnacle population abundance and structure.
Table 1. Temperature-sensitive rocky intertidal species surveyed by MarClim
Species
Common name
Notes on
biogeographical
distribution
Codium spp. Velvet Horn
Laminaria hyperborea Tangle Northern
Laminaria digitata Oarweed Northern
Laminaria saccharina Sugar Kelp Northern
Laminaria ochroleuca Golden Oarweed Southern – northern limit
S.W. Britain
Alaria esculenta Dabberlocks Northern
Himanthalia elongata Thongweed Northern
Sargassum muticum Wireweed Invasive
Ascophyllum nodosum Knotted Wrack Northern
Pelvetia canaliculata Channeled Wrack Northern
Fucus spiralis Spiral Wrack Northern
Fucus vesiculosus Bladder Wrack Northern
Fucus serratus Toothed Wrack Northern
Fucus distichus Wrack Northern – southern limit
British Isles
Cystoseira spp. Rainbow Wrack Southern
Halidrys siliquosa Sea Oak Northern
Bifurcaria bifurcata A Brown Seaweed Southern
Mastocarpus stellatus False Irish Moss Northern
Chondrus crispus Carrageen Northern
Lichina pygmaea Black Lichen Southern
Halichondria panicea Breadcrumb Sponge Ubiquitous
Anemonia viridis Snakelocks Anemone Southern
Aulactinia verrucosa Gem Anemone Southern
Actinia fragacea Strawberry Anemone Southern
Actinia equina Beadlet Anemone Ubiquitous
Sabellaria alveolata Honeycomb Worm Southern
Sabellaria spinulosa Ross Worm Ubiquitous
Chthamalus stellatus Poli’s Stellate Barnacle Southern
Chthamalus montagui Montagu’s Stellate Barnacle Southern
Semibalanus balanoides Acorn Barnacle Northern
Balanus crenatus Acorn Barnacle Northern
Balanus perforatus Volcano Barnacle Southern
Elminius modestus Australasian Barnacle Invasive
Campecopea hirsuta Southern
Clibanarius erythropus Red Clawed Hermit Crab Southern
Haliotis tuberculata Green Ormer Southern
Tectura testudinalis Tortoiseshell Limpet Northern
Patella vulgata Common Limpet Northern
Patella depressa Black Footed Limpet Southern
Patella ulyssiponensis China Limpet Southern
Gibbula umbilicalis Flat Topshell Southern
Gibbula pennanti Pennant’s Topshell Southern
Gibbula cineraria Grey Topshell Ubiquitous
Osilinus lineatus Toothed Topshell Southern
Calliostoma zizyphinum Painted Topshell Southern
Littorina littorea Common Periwinkle Northern
Littorina saxatilis agg. Rough Periwinkle Northern
Littorina neglecta Obscure Periwinkle Southern
Melarhaphe neritoides Small Periwinkle Southern
Nucella lapillus Dogwhelk Northern
Onchidella celtica Celtic Sea Slug Southern
Mytilus spp. Common Mussel Ubiquitous
Asterias rubens Common Starfish Northern
Leptasterias mulleri Northern Starfish Northern
Paracentrotus lividus Purple Sea Urchin Southern
Strongylocentrotus droebachiensis Northern Sea Urchin Northern
Note: Ubiquitous species occur throughout the British Isles
18
Fig. 5. Location of sites where (a) semi-quantitative broadscale resurvey, (b) quantitative barnacle resurvey (c) quantitative limpet
resurvey and (d) topshell population resurveys by the U.K. team were made in 2001-2005. Please note additional surveys are ongoing
and these maps represent sites that have been visited up to October 2005.
(d) Topshell resurvey locations
(c) Limpet resurvey locations
(b) Barnacle resurvey locations
(a) Broadscale resurvey locations
19
7. Main Findings: Changes in the Geographic Distribution of Intertidal Species in Response to Climate
Change and the Biological Mechanisms Underlying these Changes
MarClim has identified shifts in the geographical limits of
species and potential biological mechanisms causing the
observed responses of species to climate change.
Southern species advancing
Topshells
Gibbula umbilicalis
In 2002 a breeding population of the flat topshell, Gibbula umbilicalis was recorded at Fresgoe, north Scotland
(Fig. 6). This was the first time that animals have been recorded at this location despite intensive searches
carried out in the 1970s and 1980s. The establishment of the Fresgoe population constitutes a northern
range extension of approximately 55km beyond the previous range edge population at Skerray, north
Scotland since 1985. Two individuals were also found 80km past Skerray at Murkle Bay in 2003 (Fig. 6),
where occasional individuals had been found during previous searches in the 1980s. Since the current period
of climate warming began in the late 1980s, populations of Gibbula umbilicalis near their northern range limits
have shown statistically significant increases in abundance (Fig. 7). Analysis of recruitment and sea surface
temperature (SST) data for the 1970s, 1980s and 2000s shows that the increased abundance is likely to be
the result of increases in the frequency and strength of recruitment success within populations at and close to
the northern range edge in response to warmer temperatures. Winters have become warmer and mean sea
temperatures have increased by ~0.5ºC along the north Scotland coastline in the past two decades (Fig. 8).
A significant, positive relationship between winter SST and recruitment success in populations close to their
range limits in Scotland exists in the 1970s, 1980s and 2000s, with much higher numbers of newly settled
juveniles surviving the milder winters during the 2000s in populations close to their northern range edge,
resulting in adult breeding populations developing (Mieszkowska et al. in review).
The eastern range limit has also extended in the English Channel since the mid 1980s (Fig. 9b) via the
colonisation of artificial structures, such as sea defences. The limit was previously set at the Isle of Wight,
where there is no clear relationship between population success and sea temperature in the 1970s, 1980s or
the 2000s (Mieszkowska et al. in review), indicating that this species eastern range limit was set by a lack of
suitable rocky substrate beyond the Isle of Wight. Colonisation of sea defences has been rapid; Gibbula
umbilicalis was not found on sea defences at Elmer in 2001; one or two individuals were found in 2002;
occasional individuals were present in 2003 and it had become common by 2004. Breeding populations
showing signs of recent recruitment were also found on natural rocky shores east of Brighton in 2004 and
2005 with occasional individuals being found at Beachy Head, showing that artificial structures can act as
stepping stones between areas of suitable natural habitat which were previously too distant to allow
successful colonisation.
20
Fig. 6. Extension of the northern range limit of the breeding population of Gibbula umbilicalis at Fresgoe in north Scotland between 1985
and 2002 (from Mieszkowska 2005). Two individuals were found at Murkle Bay in 2003.
0
5
10
15
20
25
Fresgoe
Skerray
Rispond
Eriboll
Culkein
Clashnessie
Reiff
location
mean abundance per min
1985
2002
2003
2004
6.0
6.5
7.0
7.5
8.0
8.5
9.0
9.5
10.0
10.5
1975
1978
1981
1984
1987
1990
1993
1996
1999
2002
Year
Mean Winter SST °C
59 N, 4 W
59 N, 5W
59 N, 6W
Fig. 7. Abundance of Gibbula umbilicalis at sites close to
their northern range edge in north-eastern Scotland (from
Mieszkowska 2005).
Fig. 8. Mean winter SST (± 1 SE) 1975-2004 for the grid
squares at the north-eastern range edge of G. umbilicali
s
on the north coast of Scotland. Data provided by the
Hadley Centre, UK Met Office (HadISST, Version 1.1).
(from Mieszkowska 2005)
21
Species distribution (1950s, 1980s) Range extension (2000-2004) Range limit
N
40 km
Key:
Not found
Rare
Occasional
Frequent
Common
Abundant
Gibbula umbilicalis Osilinius lineatus
Patella ulyssiponensis (aspera)
Patella depressa
Melaraphe neritoides Balanus perforatus
a)
e)
d)
c)
b)
g)
f)
Species distribution (1950s, 1980s) Range extension (2000-2004) Range limit
N
40 km
Key:
Not found
Rare
Occasional
Frequent
Common
Abundant
Species distribution (1950s, 1980s) Range extension (2000-2004) Range limit
N
40 km
N
40 km
N
40 km
Key:
Not found
Rare
Occasional
Frequent
Common
Abundant
Key:
Not found
Rare
Occasional
Frequent
Common
Abundant
Gibbula umbilicalis Osilinius lineatus
Patella ulyssiponensis (aspera)
Patella depressa
Melaraphe neritoides Balanus perforatus
a)
e)
d)
c)
b)
g)
f)
Fig. 9. Abundances and range extensions for six species along the English Channel.
Osilinus lineatus
In the English Channel the previous range limit of Osilinus lineatus was located at Lyme Regis (Kendall 1987),
with isolated individuals recorded past this point (Hawthorne 1965, Kendall 1987). In 2002, 2003 and 2004 a
breeding population was present at Osmington Mills, 55km east of Lyme Regis (Fig. 10). Individuals were
also found at Lulworth Cove, and Freshwater Bay on the Isle of Wight, 60km and 120km respectively further
east (Fig. 9f). A detailed study of the range edge population in Dorset has been made by local collaborators
aided by some MarClim funding.
Populations of the toothed topshell Osilinus lineatus were eradicated or reduced in the cold winter of 1962-63
from the northern range edge on Anglesey to the south of the Lleyn Peninsula (Figs. 11, 12; Crisp 1964), and
only occasional specimens have since been recorded by Hawkins and co-workers at Rhosneigr, on the south-
west coast of Anglesey during the cooler 1980s and early 1990s (Fig. 13). Resurveys as part of MarClim
showed that Osilinus lineatus had recolonized all sites in north Wales up to Cemlyn Bay, Anglesey, in
response to climate warming since the late 1980s, and populations are now present on the Lleyn Peninsula at
locations where previous records show apparent absence or only isolated individuals (Fig. 14).
There has also been a significant increase in the abundance of Osilinus lineatus at many sites covering four
degrees of latitude (approx 450km; Fig. 15), accompanied by an increase in successful recruitment at all
locations for which data from the 1980s exist (Mieszkowska 2005). Inter-annual variation in abundance
during the 2000s was significantly less than the change in abundance between the 1980s and 2000s,
indicating that major decadal-scale increases in population abundance have occurred in response to the
decadal-scale changes in climate. Recruitment is still not occurring every year at the northern range edges in
north Wales (Fig. 17) although gonads are known to develop (Lewis 1964, Bowman 1986, Williamson et al. in
22
press), suggesting that the climate is still not warm enough for successful reproduction or survival of the
juvenile cohort in every year.
Warmer or longer summers are likely to increase the period available for reproduction resulting in increased
population sizes (Bode et al. 1986, Kendall 1987). Any latitudinal cline in the length of the reproductive cycle
and spawning periodicity would suggest a correlation with environmental temperature, in this case warming,
which decreases along a latitudinal gradient from the tropics to the poles. Preliminary studies on the
reproductive output of Osilinus lineatus have shown that there is no apparent gradient in timing and periodicity
of gonad cycles from populations sampled from their northern range edge to populations at the centre of their
range (covering 4.7 degrees of latitude), but reproduction occurs earlier in warmer years such as those
experienced in the 2000s (Mieszkowska 2005) and the last warm period in the 1950s (Williams 1964, Williams
1965, Desai 1966). The earlier arrival of juvenile recruits on the shore in warm years provides them with a
longer time period to build up metabolic reserves prior to the onset of winter. This may act in conjunction with
the increased survival of juveniles that has been observed when experimentally exposed to increased winter
temperature (Fig. 16). These results support the conclusions that observed re-colonization and range
extensions are being driven by increased survival of juveniles within populations close to range edges,
leading to increased numbers of gametes being produced (Mieszkowska 2005).
Fig. 10. Eastern range extension (black line) of Osilinus lineatus along the English Channel, 1986-2002
(Crisp & Southward 1958, Kendall 1987; from Mieszkowska 2005)
23
0
5
10
15
20
25
30
35
40
Aberaeron
West Angle Bay
Hartland Quay
Wellcombe
Widemouth
Wembury
Noss Mayo
Prawle
Crackington Haven
Lyme Regis
Churstone
Locquirec
Le Guersit
Ille Callot
Roscoff
Brignogan
site
mean abundance per minute
1986
2002
0
20
40
60
80
100
cold ambient warm
treatment
percentage survival
4ºC 9ºC 13ºC
Fig. 15. Abundance of Osilinus lineatus at sites close to its
northern limits & further south into the range in England & France
(± 1SE) (from Mieszkowska, 2005).
Fig. 16. Survival of Osilinus lineatus ‘0’ cohort juveniles
exposed to winter sea temperatures (± 1 SE) (from
Mieszkowska 2005)
Fig. 11. Abundance of Osilinus lineatus at the northern range
edge, north Wales in the 1950s (Crisp and Knight-Jones 1954).
Fig. 12. Abundance of Osilinus lineatus at the northern range
edge, north Wales in 1964 after the cold winter 1962/63 (Crisp
1964).
Fig. 13. Abundance of Osilinus lineatus at the northern range
edge, north Wales in the 1980s (Hawkins unpublished).
Fig. 14. Abundance of Osilinus lineatus at the northern range
edge, north Wales in the 2000s (Mieszkowska 2005)
24
Fig. 17. Population structures of Osilinus lineatus at northern range limits in Wales in the 2000s (from Mieszkowska 2005).
Limpets
The southern limpet Patella depressa is now much more common at many locations in Britain compared to
the cooler early 1980s. This is particularly the case in north Cornwall and Cardigan Bay. In north Wales,
numbers have been steadily increasing at sites close to the northern range edges throughout the 2000s, and
a breeding population is present at Cricceith, north Wales, where previous surveys in the 1980s and 1990s
found only isolated individuals. Elsewhere abundances have not recovered to levels seen in the warm 1950s
(Kendall et al. 2004). Therefore it would appear that local factors can override climatic influences in this
species. Evidence would, however, suggest that more of the Patella depressa population is spawning in
recent years (Moore 2005). Spawning events for P. depressa are happening earlier in the year and for longer
periods than previous records suggest, with evidence of gonad redevelopment in south-west Britain (Fig18;
Moore 2005). Resurveys show that the eastern range limit in the English Channel of Patella depressa has
recently extended 30km east from the Isle of Wight to Hayling Island, where a small population has been
found on artificial sea defences (Fig. 9d). The range of Patella ulyssiponensis has also extended eastwards
in the English Channel with a breeding population recorded at Seaford in August 2004, 120km further east
than Bembridge, Isle of Wight where only isolated individuals had been found in the 1980s (Fig. 9e). A
breeding population has become established at Bembridge since the mid 1990s. A Patella ulyssiponensis
was also recorded further east at Beachy Head.
25
Fig. 18. The proportion of P. vulgata and P. depressa in advanced gonad states (gonad stages 4 & 5 after Orton et al. 1956) for a)
Kingsand and b) Andurn Point south-west Britain (from Moore 2005).
Winkles
Melarhaphe neritoides used to only be found as far east as the Isle of Wight. It is now found throughout the
eastern English Channel on artificial substrata such as sea walls, piers and sea defences as well as less
commonly on chalk cliffs (Fig. 9c). The common factor in the extensions of winkles and limpets along the
eastern basin of the English Channel is the colonisation of recently constructed artificial coastal defences.
These species have used the structures as stepping stones across areas where unsuitable soft substrata
occurs. Although these increases in range have occurred during the current period of climate change, at this
stage it cannot be concluded that they are climate-driven.
Barnacles
MarClim resurveys found an increased abundance in northern populations of the warm water barnacles
Chthamalus montagui and Chthamalus stellatus along the Atlantic coastline of northern Scotland since the
1950s. Isolated individuals of both Chthamalus montagui and Chthamalus stellatus have been found further
south along the North Sea coastline of north-east Scotland during each survey year of the 2000s. In 2005
Chthamalus montagui was found as far south as Pittenweem in Fife. These range extensions have coincided
with warmer Atlantic waters entering the North Sea (Hulme et al. 2002).
Infilling of gaps between isolated individuals and breeding populations of both species have also been
recorded. In 2003 Chthamalus montagui were found on artificial substrata on much of the Wirral coastline,
where previously only one or two isolated individuals had been recorded during the 1950s (Southward, pers
comm.). In 2004 Chthamalus stellatus was recorded for the first time on the Isle of Man, 100km from the next
nearest known population in North Wales. Between the 1960s and the 2000s the eastern range limit of
Balanus perforatus in the English Channel has extended 120km east, with individuals found on artificial
substrata at Hastings and subsequently on natural rock at Fairlight Cove and Sandgate, near Hythe in Kent
(Herbert et al. 2003). There has been an increase in the abundance of Balanus perforatus at 18 out of 38
sites resurveyed in the 1990s compared to surveys carried out in the 1950s in the English Channel (Herbert et
al. 2003). At 17 sites there was no difference in the abundance between surveys, while at three sites there
was a decrease in abundance (Herbert et al. 2003).
Macroalgae
The most easterly population of the southern brown alga Bifurcaria bifurcata has remained near Dartmouth,
Devon close to the 6°C winter sea surface isotherms, for the last 100 years (Fischer-Piette 1936, Crisp and
Kingsand
0
0.2
0.4
0.6
0.8
1
ASONDJFMAMJJASONDJF
Month
Proportion of limpets
in advanced gonad
states
P. vulgata P. depressa
An du r n P o in t
0
0.2
0.4
0.6
0.8
1
ASONDJFMAMJJASONDJF
Month
Proportion of limpets
in advanced gonad
states
P. vulgata P. depressa
26
Southward 1958, Lewis 1964). Dubious records do however exist from the 19th century suggesting a more
easterly range limit (Harvey 1846-1851, Johnstone and Croall 1860). In 2002 Bifurcaria bifurcata was found
growing at Portland Bill, Dorset, indicating a possible eastern range extension of approximately 150km. The
plants, usually restricted to rock pools near their range limits, were located on open rock on the low shore, a
habitat previously only observed to be used by plants in south-west England (Mieszkowska et al. in prep).
Northern species retreating
Resurveys by MarClim have found less evidence of a reduction in the ranges of northern species, although
some species have not been found on shores where they have been previously recorded.
Algae
Although the absolute range limits of the northern brown alga Alaria esculenta have not changed, this species
has suffered severe declines in abundance or localised extinctions on shores around the coasts of south-west
Britain since previous surveys in the 1950s (Mieszkowska et al. in prep). Continued absence from most of
these shores was confirmed by the MarClim team between 2001-2004. Transplant experiments failed to re-
establish Alaria esculenta at sites where it had been recorded in the 1950s (Vance 2004). Alaria esculenta
has also disappeared from Robin Hood’s Bay, Northumberland between the 1980s and the 2000s.
Barnacles
Although the biogeographic range of Semibalanus balanoides in Britain has not changed it is less abundant in
the 2000s than during the cooler climatic periods of the 1960s and 1970s on shores of south-west England.
The southern species of barnacles (Chthamalus montagui and Chthamalus stellatus) are now more abundant
on shores of the south-west than Semibalanus balanoides. In recent years there have been more years
where Semibalanus balanoides settlement has failed on shores in south-west Britain, particularly in 2004
when there was little or no settlement of this species on shores around Plymouth (Moore 2005). Work from
further south in its range, in Galicia, suggests it has disappeared from its previous southern limit in Europe
(Wethey, pers. comm.)
Limpets
The relative proportions of the northern limpet Patella vulgata have reduced in comparison to the southern
species of limpet, Patella depressa, since the 1980s in south-west Britain. These two species have different
bioeographical origins but occupy similar niches on the shore in Britain and northern Europe. In 2004, less
than 10% of the Patella vulgata population on semi-exposed to exposed shores around Plymouth reached
gonad states at which spawning could occur (Fig. 18; Moore 2005). This is in direct contrast to the increase
in spawning success of the southern limpet Patella depressa in recent years (Fig. 18; Moore 2005). The
tortoiseshell limpet Tectura testudinalis was not found on the Isle of Man in the 2000s where it had been
previously recorded during the 1970s and early 1980s (Hawkins pers. ob.).
There may be various reasons for the apparent lack of northern species range contractions. Many of the
northern/boreal species surveyed by the MarClim team reach their range limits slightly further south than
Britain and Ireland (Fischer-Piette 1936, 1948, Fischer-Piette and Prenant 1956, Southward et al. 1995).
Climatic conditions in Britain and Ireland may not yet have reached a threshold beyond which these species
27
will contract their range. In many cases these species will initially respond to increased warming by
decreasing in abundance and becoming limited to habitats which provide more amelioration from adverse
climatic conditions, such as under macroalgal canopies (Moore in review). The lack of evidence for
northern/boreal species ranges being restricted polewards may also be the result of changed climatic
conditions affecting the most sensitive stages in a species life history. Such changes may take many years
before they become apparent at the population level due to the longevity of many intertidal species. It is likely
that northern/boreal populations will become increasingly dominated by adults as mortality of the more
susceptible juvenile stages becomes more frequent in warmer conditions (Lewis 1996) and as recruitment
fails (Svensson et al. 2005).
Table 2. Summary of changes in the range limits of species with northern and southern biogeographic ranges
a) Species with southern biogeographic distributions b) Species with northern biogeographic distributions.
Species Previous
Eastern limit
Range Extension Previous
Northern limit
Range Extension Comments
Gibbula umbilicalis
Osilinus lineatus
Patella depressa
Patella ulyssiponensis
Chthamalus montagui
Chthamalus stellatus
Balanus perforatus
Bifurcaria bifurcata
Isle of Wight
Lyme Regis
Isle of Wight
Isle of Wight
Isle of Wight
Isle of Wight
Isle of Wight
Dartmouth,
Devon
Beachy Head
(125km)
Osmington Mills
(55km)
Hayling Island
(30km)
Seaford (120km)
None evident
None evident
Fairlight Cove,
Kent (170km)
Portland Bill
(150km)
Skerray (1985)
Lleyn Peninsula
(1964)
Porth Oer (1952)
Orkney
Shetland
South Wales
Wales
Fresgoe (55km)
North Anglesey
(40km)
Cricceith
(cut back after cold
winter 1962/63)
None evident
Pittenweem
(140km)
Cove Bay,
Aberdeenshire
(40km)
Under verification
Isolated
individuals as far
east as Isle of
Wight
Isolated
individuals on
Anglesey prior to
1962/3 & 1980s
Northern limit
Norway
Now Common in
Orkney
Now Abundant in
Orkney
Previous records
incorrect, range
being verified
Species Previous limit Range
Contraction
Previous
Southern limit
Range Contraction Comments
Alaria esculenta
Tectura tessulata
Salcombe
Dodman Point,
South Cornwall
(120km)
Present in Robin
Hood’s Bay in
1980s
Dublin and
Anglesey
Not present in Robin
Hood’s Bay 2004.
Loss of species
from south of Isle of
Man since 1980s
Disappeared
from much of the
western English
Channel in
1950s. Did not
recover post
1963.
a
)
b
)
28
8. Prediction of Future Changes: Modelling Species Populations
Comparison of long time series of population abundance with
climatic data can be used to model past and future distributions
of key intertidal species in relation to climate.
The MarClim project has constructed population models of competing pairs of northern and southern species
of barnacle. Forecasting the response of species to shifts in environmental conditions is important for scoping
the impacts of climate change, but predicting how a species will react is complicated by a number of factors.
For example, the presence of competing species or fluctuations in the abundance of predator or prey species
may all mediate climatic influence. Untangling these factors can be problematic, particularly where there are
interactions between several species. However, long-term data sets and a sound knowledge of the
underlying biology of the species allow for biologically realistic predictions of species responses to climate
change.
In the British Isles, the intertidal zone on open coasts is dominated by one northern (Semibalanus balanoides)
and two southern (Chthamalus montagui and Chthamalus stellatus) species of barnacles. Semibalanus
balanoides is a midshore boreo-arctic species, while the lusitanian C. montagui and C. stellatus extend further
up the intertidal zone. Intertidal barnacles are ideal for an investigation of climatic influence on population
dynamics. In the UK all three species reach the edges of their biogeographical distributions, are relatively
unimpacted by human activities and their population ecology and interactions with other species are well
known. Populations at range edges should be most sensitive to fluctuations in the environment. Indeed, a
record of recent changes in relation to environmental variation in the second half of the 20th century is
available to MarClim as a long time series of population data (35 years) collected by Alan Southward (see The
basis of the MarClim project – Historical data), restarted by Hawkins in 1997.
The historical barnacle time series revealed that during a cold period in the late 1960s and early 1970s
Semibalanus balanoides increased in numbers while Chthamalus spp. decreased (Fig. 19). Two statistical
techniques identified environmental sensitivities of key processes of population regulation in barnacle
populations. Firstly, general linear models were used to calculate average numbers of barnacles at the mid
shore level for each year of the 35-year series. Cross-correlation of the annual abundance of Semibalanus
balanoides, with monthly time-lagged local sea surface temperature, showed a strong negative association
between counts of adults and temperatures in spring of the previous year. Adult counts of barnacles are
dominated by the youngest adult age class (1+) so this association suggested a critical period for survival in
the first few months after settlement. In contrast high sea surface temperature values were associated with
low counts. For the chthamalid species, the strongest association was also with sea temperature the previous
spring, but the association was positive indicating that chthamalids are favoured by a warm spring.
29
Secondly, the statistical technique of formal path analysis revealed that while temperature may be acting
directly on Semibalanus balanoides, the climatic influence on Chthamalus spp. is probably mediated by the
abundance of the competitively superior Semibalanus balanoides (Connell 1961). A hierarchy of age-
structured population models were constructed to explore whether climate acts directly on each species or if
inter-species competition is important. Three levels of model, each of increasing complexity, were developed:
(1) temperature influences juvenile Semibalanus balanoides; (2) as 1 but with adult numbers influencing
Semibalanus balanoides recruitment; (3) as 2 but with Semibalanus balanoides juveniles competing directly
with young Chthamalus spp. In models 1 and 2, competition was expressed as simple space pre-emption.
Models were assessed to determine the optimal complexity; if a model is too simple, key components driving
population regulation may be excluded. The explanatory power of the three models was assessed using an
index that combined the goodness of fit of each model to the data with the number of free parameters in the
model (AIC: Akaike’s Information Criterion). The model with the largest AIC was selected as the best of those
considered.
Adding complexity increased the fit of the models. Models started to reproduce the large fluctuations
observed in the historical time series for both species once competition was introduced, showing that simple
temperature dependent models may be inadequate in explaining changes in population numbers (Fig. 20).
The final task for this section of modelling was to run the models forward using temperatures forecast under
different emissions scenarios from the UKCIP02 scenarios to determine the probability of extinction of
Semibalanus balanoides in south-west England over the next 50 years (Fig. 21). MarClim has shown that
successful and realistic hindcasts and forecasts can be made of species responses to climatic change if key
elements of the biology and ecology of the species are incorporated into models that build on observations of
change (Poloczanska et al. in review).
Chthamalus spp.
0
2
4
6
8
10
1950 1956 1962 1968 1974 1980 1986 1992 1998 2004
no
pe
r
2
AJS
SJH/ MarCli
HWN MTL LWN
HWN MTL LWN
Semibalanus
spp.
Chthamalus
spp.
Semibalanus spp.
‘Southern’
‘Northern’
0
2
4
6
8
10
1950 1956 1962 1968 1974 1980 1986 1992 1998 2004
HWN MTL LWN
HWN MTL LWN
Chthamalus
spp.
0
2
4
6
8
10
1950 1956 1962 1968 1974 1980 1986 1992 1998 2004
no per cm
2
HWN MTL LWN
HWN MTL LWN
Semibalanus balanoides
Chthamalus
spp.
Fig. 19. Long-term changes in northern (Semibalanus balanoides) and southern (Chthamalus spp) barnacles averaged fo
r
several shores on the South coast of Devon & Cornwall (from Southward & Hawkins, unpublished; Southward 1991,
Southward et al. 1995).
30
6.0
7.0
8.0
9.0
10.0
11.0
12.0
13.0
14.0
1903 1913 1923 1933 1943 1953 1963 1973 1983 1993
Year
Temp
0
C
0.00
0.20
0.40
0.60
0.80
1.00
1.20
1903 1913 1923 1933 1943 1953 1963 1973 1983 1993
Count Adult S. balanoide
s
Model
Historical
0.00
0.50
1.00
1.50
2.00
2.50
3.00
3.50
1903 1913 1923 1933 1943 1953 1963 1973 1983 1993
Count Adult Chthamalus s
p
Model
Historical
a
)
Mean SST off Pl
y
mouth
b
)
Semibalanus balanoides
c
)
Chthamalus s
pp
.
Fig. 20. (a) Annual sea surface temperature (SST) in May from the western English Channel including station E1 [50º 2’N 4º
22’W] off Plymouth Sound (b) and (c) historical counts of adult barnacles from south west England and counts produced by two-
species population model with Semibalanus balanoides recruitment dependant on SST in May and stock size and with inter-
specific competition between juveniles: (b) Semibalanus balanoides and (c) Chthamalus spp (based on Poloczanska et al. in
review).
31
Fig.21. (top) The number of Semibalanus balanoides and Chthamalus spp. per cm
2 (bottom) Model predictions and observations of
fluctuations at high shore levels in SW England (MHWN) from 1940 to 2080, using observed (1940-2003) and projected temperature
series generated using the UKCIP02 Medium-High emissions scenario, a linear increase in mean temperature to 2080, and interannual
variability as observed from 1900 to 2000 (from Poloczanska et al. in review).
1940 1960 1980 2000 2020 2040 2060 2080
0.0
1.0
2.0
3.0
1940 1960 1980 2000 2020 2040 2060 2080
Numbe
r
per cm²
Model S. balanoides
Historical S. balanoides
Model Chthamalus spp.
Historical Chthamalus spp.
10
12
14
16
18
Temperature (°C)
Sea Surface Temperature, June
32
9. Prediction of Future Changes – Modelling Species Distributions
Broadscale abundance and environmental data can be used to
predict species distributions both now and in the future.
Using the extensive spatial broadscale resurvey data, we have modelled the likelihood of species reaching
particular abundance categories at specific locations in the British Isles in relation to environmental variables
such as wave action and sea surface temperature. GIS-based models have been developed that estimate
wave fetch for 500m grid cells over the entire European coastline (e.g. Fig. 22), whilst sea surface
temperature (SST) data have been extracted from the British Atmospheric Data Centre (GOSTA dataset)
(Bottomley et al. 1990) and from the UKCIP02 scenarios 1961-90 baseline (UKCIP02) (Hulme et al. 2002)
(Fig. 23). Some climate forecast scenarios predict stormier seas (Hulme et al. 2002).
Generalised linear models with cumulative logit link functions have been used to establish statistical
relationships between environmental variables and the categorical abundance data for key species. Wave
fetch and February SST successfully predict present day distributions for many species (Fig. 24). Where the
match is less good the discrepancy between predicted and observed distributions highlights possible
hydrographical barriers to dispersal or past historical events during recolonization of the Great Britain and
Ireland after the last ice age. For example, the models predict that present day climate is suitable for Osilinus
lineatus to occur around the west and south coasts of the Outer Hebrides but present day distribution is
restricted to south-west England and Wales and around much of the Irish coast. Osilinus lineatus may not
have made the ‘jump’ across to the Outer Hebrides from Ireland as conditions warmed after the last glacial
maximum, or hydrographical barriers may exist which impede its spread northwards. These models have
been extended to include effects of occupancy of neighbouring sites to account for vicariance effects.
It has also been possible to combine these statistical models with future climate change scenarios to forecast
species distributions. Sea surface temperature projections for the 2020s, 2050s and 2080s have been
incorporated into simulation models based on the statistical models including occupancy to predict likely
future changes in species ranges. As an example, given present day relationships between abundance and
wave fetch and SST (Fig. 24 top centre) and a 2 to 3°C rise in average February temperatures by the 2080s,
the models predict an expansion of the range of the southern barnacle Chthamalus stellatus into the entire
North Sea wherever suitable rocky habitat is present, a range extension over many 100s of km (Fig. 24 top
right). In the same scenario, the northern species of kelp, Alaria esculenta, is predicted to retreat from coasts
of south-west England and the coasts of western and southern Ireland (Fig. 24 bottom right).
The modelling exercise emphasizes the need to extend the surveys to the east coasts of Scotland and
England for which there is little baseline data. This is essential in order to monitor future responses to climate
change. Such an approach could also be used to model the outcomes of the NERC RAPID programme
which is exploring the consequences of changes in the Atlantic Thermohaline Circulation.
33
Fig. 22. Local wave exposure for 500m coastal grid squares calculated as the sum of the wave fetch in 16 sectors around each cell.
Local wave action is an important predictor of abundance of most rocky shore species and may also vary as climate changes.
9.0
7.0
6.0
10.0
8.0
7.0
8.0
9.0
8.0
12.0
11.0
8.0
9.0
10.0
11.0
9.0
9.0
11.0
10.0
10.0
Fig. 23. Sea surface temperature in February (left) for 1961 to 1990, and (right) projected for the 2080s under the UKCIP02 Medium-
High emissions scenario, showing a less than 2°C change on western coasts and over 3°C in the North Sea. Data are shown as
downscaled isotherms from 50sq km gridded data provided by UKCIP.
February SST 2080 – UKCIP02-
medium-high scenario
February 1961-1990 observed baseline
34
8
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Observed Predicted Forecast – 280 high
a) Chthamalus stellatus, a southern barnacle found at wave-exposed sites
b) Alaria esculenta, a northern seaweed found at wave-exposed sites
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Observed Predicted Forecast – 280 high
a) Chthamalus stellatus, a southern barnacle found at wave-exposed sites
b) Alaria esculenta, a northern seaweed found at wave-exposed sites
Fig. 24. Observed distributions (left), distributions predicted from statistical models with February temperature and wave exposure
(middle), and forecast distributions from models using the UKCIP 2080s high emissions scenario (right). Size of symbols indicates
observed or likely abundance categories at each site surveyed between 2002 and 2004, and for simulated sites along un-surveyed
coasts (grey circles).
35
10. Conclusions and Project Legacy
Main Conclusions:
The MarClim project has demonstrated the importance of the rescue of historical broadscale and
quantitative time series, available for rocky shores, in providing a baseline for the previously warm
period (the 1950s up to the severe winter of 1962/63) against which to measure responses of
biodiversity to current rapid climate change.
Marine species, including plankton (Beaugrand et al. 2002, Beaugrand and Ibanez 2004, Edwards
and Richardson 2004, Richardson and Schoeman 2004) and fish (Genner et al. 2004, Sims et al.
2004, Coombs et al. 2005, Perry et al. 2005, Southward et al. 2005) show rapid responses to
alterations in climate. Such changes are also clearly seen and easily quantified for intertidal species
(Herbert et al. 2003, Hiscock et al. 2004, Southward et al. 2005, Mieszkowska et al. 2006), providing
sensitive indicators of environmental change.
The rate at which the biogeographic limits of southern intertidal species are extending northwards and
eastwards towards the colder North Sea is up to 50km per decade (e.g. Gibbula umbilicalis has
extended its northern range limit along the north-east cost of Scotland by over 55km and by over
125km along the eastern English Channel since the 1980s), far exceeding the global average of
6.1km per decade in terrestrial systems (Parmesan and Yohe 2003, Root et al. 2003).
It is likely that the northward range extensions observed in North Wales and those along the north
coast of Scotland down into the North Sea have occurred in response to climatic warming increasing
reproductive effort and juvenile survival success allowing these species to establish on suitable
habitats.
It is likely that range extensions along the eastern English Channel have occurred due to a
combination of the proliferation of artificial sea defences along this coast providing suitable habitat
where none was previously present and greater recruitment success of southern species in response
to climatic warming.
The increased abundance of some southern species, such as trochids, limpets and barnacles, is likely
to have been the result of the earlier commencement (and in some cases prolonged) annual
reproductive cycles in response to warmer springs, coupled with increased survival of newly settled
recruits exposed to milder, shorter winters on the shore. These species may also be out-competing
northern equivalents as climatic conditions become more suitable for their survival and less suitable
for species with cold water affinities.
Differential rates of range extensions and contractions are likely to result in a short-term increase in
biodiversity on rocky shores close to the biogeographic boundary between cool/boreal and
warm/lusitanian waters in Britain. However, as the climate continues to warm biodiversity is likely to
36
return to previous levels as northern cold-water species ranges retract to be replaced by southern
warm-water species resulting in different species compositions.
Only sustained broadscale and long-term decadal observations can separate global environmental
change from regional and localised impacts from the intrinsic natural spatial and temporal variability of
marine ecosystems. Rocky shores provide an ideal sentinel system to monitor such changes in a
cost effective manner.
Project Legacies
Creation of the MarClim Data Archive as a central repository for historical and resurvey data records
over a 60 year period, held in a standardised electronic format with site locations fully geo-referenced
using the WGS84 reference frame. This dataset will expanded over time with repeat surveys and
monitoring (data can be accessed via the National Biodiversity Network at www.searchnbn.net).
All original data records have undergone extensive quality assurance checks in conjunction with the
original data custodians or their surviving collaborators.
Historical data for 4400 site visits at over 1000 locations are included in the MarClim electronic
database (requests for data can be made by contacting the Marine Environmental Change Network
(MECN) co-ordinator at www.mba.ac.uk/MECN).
Between 2002 and 2004 the MarClim team completed a total over 800 site visits at over 470
individual site locations. Of these resurveys 177 were undertaken at sites for which historical data
exist. Targeted work to fill gaps was done at over 40 sites in 2005.
MarClim has ensured the long-term availability of data by storing all data records in National
Biodiversity Network (NBN) format via the Marine Recorder front-end. This information will be made
available through the MarClim Website that is maintained by MarLIN at the Marine Biological
Association (www.mba.ac.uk/marclim).
MarClim data can be imported to the NBN Gateway for dissemination to the wider scientific
community through the Marine Life Information Network (MarLIN).
Sampling protocols have been developed from the original methodology and trialled during the field
phase of MarClim. They are available from the project website: www.mba.ac.uk/marclim.
A fit-for-purpose network of 80 designated sites for which long-term data exists has been designed for
future monitoring as part of a wider British marine monitoring programme. The monitoring network
documents are available form the project website: www.mba.ac.uk/marclim
37
Countryside Council for Wales have incorporated MarClim monitoring sites into their annual marine
intertidal survey programme, and have adopted MarClim survey protocols to ensure continuation of
the long-term data collection.
Twenty peer-review manuscripts are published or have been submitted for publication (Appendix 2).
Over 40 presentations have been made at international scientific conferences and policy orientated
meetings and workshops (a selection of these presentations are given in Appendix 2; see the website
for a full list).
Models have been constructed to predict species current distribution patterns and forecast future
changes in abundance and distribution based on sea surface temperature and wave action.
Models of northern and southern barnacle populations have been constructed which broadly match
historical datasets and can be used as a general proxy for forecasting other changes in biodiversity.
38
11. The Implications of Climate Change for Marine Stewardship Policy
This study highlights the scale and scope of climate change
impacts on rocky shore biota over the last 60 years and in the
future. These changes will need to be matched by changes in
marine conservation policy.
The MarClim project has already generated increased awareness of marine climate change issues that has
stimulated the production of climate change scenarios for the marine environment, as well as the creation of
the Marine Climate Change Impact Partnership (MCCIP), which was officially launched by Elliot Morley (MP)
in March 2005. A summary of the key messages is given below with more detail being provided by the
accompanying report: “The MarClim Project, key messages for decision makers and policy advisors, and
recommendations for future administrative arrangements and management measures” (Laffoley et al. 2005).
British & Irish leadership in assessing the impacts of climate change. The MarClim database
provides a globally unique cover of more than 50 temperature-sensitive seashore species over six
decades, across the geographical scale of Britain and Ireland. Used alongside other sources of
information, such as the Continuous Plankton Recorder (CPR) run by the Sir Alistair Hardy
Foundation for Ocean Science (SAHFOS), this provides Britain and Ireland with a unique opportunity
to become world leaders in the understanding of climate change induced trends and impacts on
marine ecosystems.
The impacts of climate change are real, measurable and growing in extent. MarClim
demonstrates that climate-induced changes are occurring for a variety of intertidal and fish species
and that it is a general effect not limited to smaller forms of marine life, such as plankton.
Provision of a robust baseline of information. Whilst there are many other species whose
distributions are temperature related, other than data for plankton, none have anywhere near the
extensive and robust baseline of MarClim from which to determine future climate change trends and
impacts. Results from MarClim, and other work on offshore benthic community datasets, suggest that
changes to invertebrate and fish species distributions are also occurring in both coastal and offshore
environments, and that these may affect offshore biodiversity as well as species of economic
importance for the UK.
Offshore waters. A priority for future study should be to extend the approach of MarClim into offshore
waters, and predict the consequences of climate change for key habitats and species of economic
and nature conservation importance.
The value and necessity of long-term data sets. MarClim has demonstrated the considerable value
of long-term data sets in understanding changes in marine ecosystems and the fundamental
39
importance of a long-term commitment to continued data-collection for analysing climate induced
changes in species geographic ranges and abundance.
The benefits of a consortium approach with a broad geographic coverage. The broad
geographic coverage enabled by the MarClim consortium, in bringing together relevant agencies and
research institutes in the United Kingdom and the Republic of Ireland, has enabled north-south and
east-west trends to be validated in a way that a narrower national territorial approach could not have
achieved. These lessons are being transmitted to a wider European audience via the European
Platform on Biodiversity Strategy Research, the marine board of the European Science Foundation,
Climate Change Impacts on Marine Ecosystems working group and via networks developed as part of
the E.U. Framework Programme Six of the Marine Biodiversity and Ecosystem Functioning network of
excellence.
Developing indicators for marine and coastal biodiversity. Intertidal species studied by MarClim
should be included as part of the key indicator set for marine and coastal biodiversity: they are easy
and cost effective to measure, are very responsive indicators of climate change signals, and are
supported by extensive baseline information geographically dispersed across the islands of Great
Britain and Ireland.
Factoring in marine climate change into day-to-day business. The predicted scale and nature of
marine climate change impacts needs to be built into how human activities are managed,
conservation sites established, monitored, assessed and reported on the status of habitats, species
and ecosystems, as well as the structuring of underlying legislation and policies.
A national priority for an integrated marine climate impacts monitoring programme. The
underlying monitoring work, which provides information used to understand and predict impacts of
climate change, needs to be properly resourced and structured as a core element of a fully integrated
programme. Given the importance of tracking climate change impacts and the sensitivity of the United
Kingdom and the Republic of Ireland to changes in north Atlantic circulation patterns, developing and
funding such an initiative should be seen as a national priority.
Future monitoring network. Continued monitoring of rocky shore indicators of biodiversity
responses to climate change should form an integral part of any monitoring network developed via the
Marine Environmental Change Network (MECN) programme.
Developing streamlined reporting to Government. The unique value of MarClim data can be
increased further by drawing it together with the results of other relevant marine climate change
impact projects and programmes (such as SAHFOS for plankton) to provide a streamlined,
coordinated and effective reporting framework to deliver key messages quickly to Government and
the general public on a regular and possibly annual basis – the concept of ‘annual report cards’
40
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43
Appendix 1. ACFOR abundance categories used by Southward & Crisp (1954b) and by the MarClim
project for their broadscale resurveys.
Abund ance Cate gory Species
Abund ant Common Frequent O ccasional Rar e
Barnacles
Chthamalus stellatus,
Chthamalus montagui,
Semibalanus
balanoides, Elminius
modestus
More than 1 per
cm2; rocks well
covered
0.1-1.0 per cm2;
up to 1/3 of rock
space covered
0.01-0.1 per cm2;
individuals never
more than 10cm
apart
0.0001-0.01 per
cm2; few within
10cm of each other
Less than 1 per m2;
only a few found in
30 min searching
Balanus perforatus Over 0.1 per cm2;
close groups on
most vertical
faces, often up to
MTL
0.01 to 0.1 per
cm2; adjacent
groups, not
always above
LWN
Less than 0.01 per
cm2; adjacent in
crevices
Less than 0.01 per
cm2; rarely
adjacent even in
crevices
Only a few found in
30 min searching
Limpets
Patella vulgata, Patella
depressa, Patella
aspera
Over 50 per m2 or
more than 50% of
limpets at certain
levels
10-50 per m2,
10% to 50% at
certain levels
1 to 10 per m2, 1%
to 10% at certain
levels
Less than 1 per m2
on average, less
than 1% of
population
Only a few found in
30 min searching
Topshells
Osilinus lineatus,
Gibbula umbilicalis,
Gibbula pennanti
Exceeding 10 per
m2 generally
1-10 per m2,
sometimes very
locally over 10
per m2
Less than 1 per m2,
locally sometimes
more
Always less than 1
per m2
Only a few found in
30 min searching
Periwinkles
Littorina saxatilis agg.,
Littorina
obtusata/mariae
Melaraphe neritoides
Over 1.0 per cm2
at HW, extending
down the
midlittoral zone
0.1-1.0 per cm2,
mainly in the
supralittoral fringe
Less than 0.1 per
cm2, in crevices
Only a few found in
30 min searching
Littorina littorea More than 50 per
m2
10 – 50 per m2 1 – 10 per m2 Only a few found in
30 min searching
Anemones
Actinia equina, Actinia
fragracea, Anemonia
viridis, Bunodactis
verrucosa
Many in almost
every pool and
damp place
Groups in pools
and damp places
Isolated specimens
in few pools
A small number,
usually under 10,
found after 30 min
searching
Algae
>30% 5-30% <5% Scattered
individuals
Few plants 30 min
search
44
Appendix 2. Publications by the MarClim team 2001-2005 and additional climate research by the MBA
and Collaborators.
Direct support by the MarClim project and SNH, MAFF/Defra precursors are asterisked.
Refereed Papers
* Burrows, M.T., Moore, J., & James, B. 2002. Spatial scale synchrony of population changes in rocky shore
communities in Shetland: implications for monitoring. Marine Ecology Progress Series 240: 39-48.
* Coombs, S.H., Halliday, N.C., Southward, A.J. & Hawkins, S.J. 2005. Distribution and abundance of sardine
(Sardina pilchardus) eggs in the English Channel from Continuous Plankton Recorder sampling, 1958-1980.
Journal of the Marine Biological Association of the UK, 85: 1243-1247
* Genner, M.J., Sims, D.W., Wearmouth, V.J., Southall, E.J., Southward, A.J., Henderson, P.A. & Hawkins,
S.J., 2004. Regional climatic warming drives long-term community changes of British marine fish.
Proceedings of the Royal Society of London, Biological Sciences, 271: 655-661.
* Hardman-Mountford, N.J., Allen, J.I., Frost, M.T., Hawkins, S.J., Kendall, M.A., Mieszkowska, N.,
Richardson, K.A. & Somerfield, P.J. (2005). Diagnostic monitoring of a changing environment: An alternative
U.K. perspective. Marine Pollution Bulletin 50: 1463-1471.
* Hawkins, SJ, Southward, A.J. & Genner M.J. (2003). Detection of environmental change in a marine
ecosystem-evidence from the western English Channel. Science of the Total Environment, 310: 245-56.
* Hawkins, S.J., Gibbs, P.E., Pope, N.D. Burt, G.R., Chesman, B.S., Bray, S., Proud, S.V., Spence, S.K.,
Southward, A. J. & Langston, W.J. (2002). Recovery of polluted ecosystems: the case for long-term studies.
Marine Environmental Research, 54: 215-222.
* Herbert, R.J.H., Hawkins, S.J., Sheader, M. & Southward, A.J. (2003). Range extension and reproduction of
the barnacle Balanus perforatus in the eastern English Channel. Journal of the Marine Biological Association
of the UK, 83: 73-82.
* Herbert, R.J.H., Southward, A.J., Sheader, M. & Hawkins, S.J. (in press). Influence of recruitment and
temperature on distribution of intertidal barnacles in the English Channel. Journal of the Marine Biological
Association of the UK.
* Hiscock, K., Southward, A.J., Tittley, I. & Hawkins, S.J. (2004). Effect of changing temperature on benthic
marine life in Britain and Ireland. Aquatic Conservation 14: 333-362.
* Kendall, M.A., Hawkins, S.J., Burrows, M.T. & Southward, A.J. (2004). Predicting the effects of marine
climate change on the invertebrate prey of the birds of rocky shores. Ibis (Special Edition) 146: 40-47.
45
* Lima, F.P., Queiroz, N., Ribeiro, P.A., Hawkins, S.J. & Santos, A.M. (in press). Geographic expansion of a
marine gastropod, Patella rustica Linnaeus, 1758, and its relation with unusual climatic events. Journal of
Biogeography
* Mieszkowska, N, Kendall, M.A., Hawkins, S.J., Leaper, R., Williamson, P., Hardman-Mountford, N.J. &
Southward, A.J. (2006). Changes in the range of some common rocky shore species in Britain - a response
to climate change? Hydrobiologia 555: 241-251.
* Sims, D.W., Genner, M.J., Southward, A.J. & S.J. Hawkins, 2001. Timing of squid migration reflects North
Atlantic climate variability. Proceedings of the Royal Society of London - Series B 268: 2607-2611.
* Sims, D.W., Wearmouth, V.J., Genner, M.J., Southward, A.J. & Hawkins, S.J., 2004. Low-temperature-
driven early spawning migration of a temperate marine fish. Journal of Animal Ecology 73: 333-341.
* Southward, A.J., Langmead, O., Hardman-Mountford, N.J., Aiken, J., Boalch, G.T., Dando, P.R., Genner,
M.J., Joint, I., Kendall, M.A., Halliday, N.C., Harris, R.P., Leaper, R., Mieszkowska, N., Pingree, R.D.,
Richardson, A.J., Sims, D.W., Smith, T., Walne, A.W. & Hawkins, S.J. (2005). Long-term oceanographic and
ecological research in the western English Channel. Advances in Marine Biology 47: 1-105.
* Simkanin, C.S., Power, A-M., Davenport, J., Myers, A.A. & McGrath, D. (2003). Monitoring intertidal
community change in a warming world. The Irish Scientist 2003 Yearbook May 2003. Samton Limited.
Svensson, C.J., Jenkins, S.R., Hawkins, S.J. & Aberg, P. (2005). Population resistance to climate change:
modelling the effects of low recruitment in open populations. Oecologia 142: 117-126.
* Thompson R.C., Crowe, T.P, & Hawkins, S.J. (2002). Rocky intertidal communities: past environmental
changes, present status and predictions for the next 25 years. Environmental Conservation. 29(2): 168-191.
Submitted Papers
* Mieszkowska, N., Kendall, M.A., Lewis, J.R., Richardson, A.J. & Hawkins, S.J. (in review). Range
expansion of the southern trochid gastropod Gibbula umbilicalus during recent climate warming. Journal of
Biogeography
* Moore, P., Hawkins, S.J. & Thompson, R.C. (in review). Different behaviours of congeneric species
modulate the role of habitat amelioration in a changing climate. Marine Ecology Progress Series
* Poloczanska, E.S., Hawkins, S.J., Southward, A.J. & Burrows, M.T. (in review). Modelling intertidal barnacle
population response to climate change: Hindcast, forecast and prediction. Global Change Biology
46
In Preparation
Kendall, M.A., Mieszkowska, N., Burrows, M.T., Laffoley, D. & Hawkins, S.J. (in prep). Can we use intertidal
species as indicators of climate change. Invited paper to Marine Pollution Bulletin.
Mieszkowska, N., Boalch, G.T. & Hawkins, S.J. (in prep). Changes in the biogeographic range of Bifurcaria
bifurcata in the English Channel.
Moore, P., Hawkins, S.J. & Thompson, R.C. (in prep). Species identity affects the outcome of grazer
interactions with canopy forming macroalgae.
Poloczanska, E.S., Hawkins, S.J., Mieszkowska, N, Kendall, M.A., Leaper, R, Southward, A.J., Simkanin, C.,
Power, A-M. and Burrows, M.T. (in prep). Distributions of intertidal sea anemones around British and Irish
coasts: past, present and future?
Theses
Mieszkowska, N. (2005). Changes in the biogeographic distribution of the trochid gastropods Osilinus
lineatus (da Costa) and Gibbula umbilicalis (da Costa) in response to global climate change: range dynamics
and physiological mechanisms. Ph.D. Thesis, University of Plymouth. p146
Moore, P.J. (2005). The role of biological interactions in modifying the effects of climate change on intertidal
assemblages. Ph.D. Thesis, University of Plymouth. p126
Vance, T. (2004). Loss of the northern species Alaria esculenta from Southwest Britain and the implications
for macroalgal succession. MRES Thesis, University of Plymouth.
Reports and Semi-popular Publications
Cannell, M., Brown, T., Sparks, T., Marsh, T., Parr, T., George, G., Palutikof, J., Lister, D., Dockerty, T. &
Leaper, R. (2003). Review of UK Climate Change Indicators. DEFRA Contract Report No. EPG 1/1/158.
Frost, M. T., Leaper, R., Mieszkowska, N., Moschella, P., Murua, J., Smyth, C. & Hawkins, S. J. (2004).
Recovery of a Biodiversity Action Plan species in northwest England: possible role of climate change, artificial
habitat and water quality amelioration. Confidential Report to English Nature, The Marine Biological
Association.
Kendall, M. A. (2002). MarClim – Marine Biodiversity and Climate Change. Report to BIOMARE Newsletter,
p10, Autumn 2002 Issue.
Laffoley, D., Baxter, J., O'Sullivan, G., Greenaway, B., Colley, M., Naylor, L. & Hamer, J. 2005 The MarClim
Project: Key messages for decision makers and policy advisors, and recommendations for future
administrative arrangements and management measures. English Nature Research Reports, No. 671.
47
Leaper, R. (2003). Intertidal species as indicators responses of biodiversity to rapid climate change in UK
marine ecosystems. Report to the Marine Biological Association Newsletter, p8, April 2003 Issue.
Mieszkowska, N., (2003). Osilinus lineatus. Thick top shell. Marine Life Information Network: Biology and
Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United
Kingdom.
Moore, P., Hawkins, S.J., Hiscock, K. & Southward, A.J. (Winter 2006). A Sea of Change. The Edge – The
magazine of CoastNET.
Presentations: Lectures, seminars, workshops and public events
Results from the MarClim project were disseminated via presentations to a broad audience including the
wider scientific community, policymakers, stakeholders and the general public. Below is a selected list of
presentations given by the MarClim team over the course of the project. A full list of presentations given can
be viewed at www.mba.ac.uk/marclim.
P.J. Moore, S.J. Hawkins, M. J. Genner, D. W. Sims, M. Burrows, E. S. Poloczanska, N. Mieszkowska, K.
Hiscock & A. J. Southward. ‘Climate change – impact on biodiversity and ecosystems’. Devon Living Coasts
Conference, Met Office, Exeter, Oct 2005.
S.J. Hawkins, G. Beaugrand, P.C. Reid, M. J. Genner, D. W. Sims, M. Burrows, E. S. Poloczanska, N.
Mieszkowska, P.S. Moschella & A. J. Southward. ‘Climate Change Impacts on Marine Biodiversity and
Ecosystems’. European Platform for Biodiversity Research Strategy, Hosted by DEFRA and the Scottish
Executive as part of the UK presidency of the EU, Aviemore, Scotland, Oct 2005
S.J. Hawkins, M. J. Genner, D. W. Sims, M. Burrows, N. Mieszkowska, S. R. Jenkins, R. Leaper, M. Frost, P.
S. Moschella, P. Masterson & A. J. Southward. ‘The combined influences of global environmental change
and regional and local impacts on English Channel ecosystems’. Invited Speaker on section to inform Marine
Bill considerations. British Ecological Society Annual Meeting, University of Hertfordshire, Sept 2005.
M.T. Frost, K. Hiscock, N. Mieszkowska & S.J. Hawkins ‘Long-term monitoring and data collection projects
hosted by the MBA: The UK Marine Environmental Change Network (MECN), Marine Life Information
Network for Britain and Ireland (MarLIN) and Marine Biodiversity and Climate Change project (MarClim)’.
Long Term and Large Scale Management of Marine Biodiversity Information: MarBEF workshop, Helgoland.
Mar 2005.
N. Mieszkowska, S.J. Hawkins, M.T. Burrows, M.A. Kendall, R. Leaper, E. S. Poloczanska & A.J. Southward.
‘Accounting for climate change in monitoring intertidal communities in Wales’. Invited Speaker. Countryside
Council for Wales Marine Monitoring Workshop, Aberystwyth Oct 2004.
48
N. Mieszkowska, S.J. Hawkins, M.T. Burrows, M.A. Kendall, R. Leaper, E. S. Poloczanska & A.J. Southward.
‘MarClim Marine Biodiversity and Climate Change research in Britain’. National Science Foundation (USA)
network to co-ordinate research on the North Atlantic (CORONA), Plymouth July 2004.
S.J. Hawkins, M.J. Genner, D.W. Sims & A.J. Southward. ‘Climate driven change in the Western English
Channel Ecosystem’. Invited presentation. Climate Change and Aquatic Systems: Past, Present & Future.
British Ecological Society, Marine Biological Association and Freshwater Biological Association, University of
Plymouth, U.K. 21st-23rd July 2004
R. Leaper & A.J. Southward. ‘Detecting a signal from noise: retrospective analysis of intertidal communities
over a period of major climate fluctuation’. Climate Change and Aquatic Systems: Past, Present & Future.
British Ecological Society, Marine Biological Association and Freshwater Biological Association, University of
Plymouth, U.K., July 2004
N. Mieszkowska, S.J. Hawkins, R.C. Thompson & M.A. Kendall. ‘Is climate change driving increased
reproductive success in British trochid gastropods?’ Climate Change and Aquatic Systems: Past, Present &
Future. British Ecological Society, Marine Biological Association and Freshwater Biological Association,
University of Plymouth, U.K., July 2004.
E.S Poloczanska, M.T Burrows & S.J Hawkins. ‘Barnacles and Climate Change’. Climate Change and
Aquatic Systems: Past, Present & Future. British Ecological Society, Marine Biological Association and
Freshwater Biological Association, University of Plymouth, U.K., July 2004
P. Moore, S.J. Hawkins & R.C. Thompson. ‚Ecological complexity places climate envelope in the shade’.
Climate Change and Aquatic Systems: Past, Present & Future. British Ecological Society, Marine Biological
Association and Freshwater Biological Association, University of Plymouth, U.K., July 2004
S.J. Hawkins, M.J. Genner, D.W. Sims & A.J. Southward. Gaia and Global Change hosted by James
Lovelock: ‘Changes in marine life in the English Channel in response to climate and fishing pressure’. Invited
speaker. Dartington, UK, Jun 2004.
N. Mieszkowska, S.J. Hawkins, M.T. Burrows, M.A. Kendall, R. Leaper & A.J. Southward. ‘Application of the
Marine Biodiversity and Climate Change Project research to marine policy making’. UKCIP United Kingdom
Climate Impacts Programme workshop for marine decision-makers – climate change impacts – what do we
need to know? Invited speaker. London, Apr 2004
S.J. Hawkins, N. Hardman-Mountford, D.W. Sims, M.J. Genner & A.J. Southward. ‘Long-term changes in the
western English Channel by the Marine Biological Association: making the case for time series’. Keynote
speaker. International conference hosted at the Linnaean Society on the use of long-term databases for the
prediction of ecological change, London, Oct 2003.
49
N. Mieszkowska, M.A. Kendall & S.J. Hawkins. ‘Detecting the effects of climate change on intertidal diversity
using long-term datasets’. International conference hosted at the Linnaean Society on the use of long-term
databases for the prediction of ecological change, London, Oct 2003.
M.J. Genner, D.W. Sims, V. Wearmouth, E. Southall, S.J. Hawkins & A.J. Southward. ‘Climate and fishing
drive a century of change in a marine fish community’. International conference hosted at the Linnaean
Society on the use of long-term databases for the prediction of ecological change, London, Oct 2003.
R.C. Thompson, T.P. Crowe & S.J. Hawkins. ‘Rocky intertidal communities: past environmental changes,
present status and predictions for the next 25 years’. International Conference on Environmental Future,
Zurich, Switzerland, Mar 2003.
S.J. Hawkins, M.T. Burrows, M.A. Kendall, R. Leaper, N. Mieszkowska, P. Moore, A.J. Southward & R.C.
Thompson. ‘Climate change and temperate reef ecosystems: integrating space and time’. International
Temperate Reefs Symposium. Invited convenor of climate session. Christchurch, New Zealand, Jan 2003.
R. Leaper, M.T. Burrows, M.A. Kendall, N. Mieszkowska, P. Moore & A.J. Southward. ‘Can intertidal species
be used as indicators of global climate change? Measuring and predicting responses of marine ecosystems
in the North East Atlantic’. Invited speaker. International Temperate Reefs Symposium, Christchurch, New
Zealand, Jan 2003.
S.J. Hawkins, K. Hiscock, R. Leaper, N. Mieszkowska, M. Kendall, R.C. Thompson, D. Sims, M. Genner, & J.
Hill, M.T. Burrows, A.J. Southward. ‘The impacts of climate change on marine species and ecosystems
around the British Isles and Ireland’. Invited speaker. Marine Conservation Society, Oct 2002
S.J. Hawkins, M.T. Burrows, M.A. Kendall, A.J. Southward & R.C. Thompson. ‘Climate change and
biodiversity in long-term work by the MBA’. Invited seminar. Tyndall Workshop, University of East Anglia, Oct
2001.
International Working Groups
S.J. Hawkins Intergovernmental Panel on Climate Change reviewer
S.J. Hawkins European Science Foundation, Marine Board Working Group on Climate Change Impacts
on Marine Ecosystems.
50
Appendix 3. MarClim Project Deliverables
Summary of Project Progress in Year 1 (April 2001 - June 2002).
Project Tasks Product
1. Identification
& extraction of
data
Spreadsheet of all known datasets and stewards Spreadsheet
Secure access to priority data sets Update
spreadsheet
2. Archiving
databases Website and database progress Activity
report
3. Analyses of
data
Southward, Hawkins & Burrows barnacle datasets – Progress to
November 2001 Report
Crisp and Southward broadscale data – The European geographical
distribution of intertidal indicators
Report /
maps
Southward, Lewis SOTEAG, MBA, & Lewis co-workers data – The
responses of intertidal indicators: Barnacles, limpets and topshells
Activity
report
4. Re-survey Resurvey of selected sites to test and train team – Progress to October
2001
Activity
report
Preliminary survey of Lewis sites – Progress to November 2001 Activity
report
Resurvey with new MarClim team
5. Monitoring
network
Targeted quantitative sampling at a selection of sites for future low cost
monitoring
Activity
report
MarClim sampling protocols Protocol
report
6. Prediction
Impacts of climate change on seabed wildlife in Scotland: Hiscock, K.,
Southward, A., Tittley, I., Jory, A. & Hawkins, S. 2001. The impact of
climate change on subtidal and intertidal benthic species in Scotland.
Edinburgh, Scottish Natural Heritage (Survey and Monitoring Series)
Report
8.
Communication
Conference papers at University College London – MarClim: Marine
Biodiversity and Climate Change
Conference
paper
Conference papers at SAFHOS and Edinburgh – Long term change in
the western English Channel
Conference
paper
Moore, P., Hawkins, S.J., Hiscock, K. & Southward, A.J. (Winter 2006).
A Sea of Change. The Edge – The magazine of CoastNET.
Semi-
popular
article
Herbert, R. J. H., S. J. Hawkins, et al. (2003). Range extension and
reproduction of the barnacle Balanus perforatus in the eastern English
Channel. Journal of the Marine Biological Association of the U.K. 83: 73-
8
Manuscript
End of Phase Report (1 & 2) Final Report
9. Policy
application Draft report of the MarClim policy workshop Brief report
51
Summary of Project Progress in Year 2 (July 2002 - June 2003).
Module Project Tasks Document
1. Identification
& extraction of
data
Secure access to priority data sets and assess quality Update
spreadsheet
2. Archiving
databases Data entry Activity
Report
3. Analyses of
data
Kendall, M.A., Mieszkowska, N. & Hawkins, S.J (in prep). Long-term
changes in the distribution, size, age and longevity of some near-limit
populations of Trochid Gastropod Osilinus lineatus
Manuscript
Poloczanska, E.S., Hawkins, S.J., Southward, A.J. and Burrows, M.T. (in
review). Modelling intertidal barnacle population response to climate
change: hindcast, forecast and prediction. Global Change Biology
Manuscript
MarClim phase 3 update: October 2002 to February 2003 Activity report
4. Re-survey Resurvey by MarClim team
Broadscale re-survey Activity report
Re-survey of Kendall and Lewis sites Activity report
5. Monitoring
network Proposals for establishment of the MarClim monitoring network Activity report
MarClim monitoring network: Provisional sampling strategy and standard
operational procedures Activity report
7. Providing
data access Establishment of data access vehicle on the website – awareness report Awareness
report
8.
Communication
Herbert, R.J.H., Southward, A.J., Sheader, M. and Hawkins, S.J. (in
press). Influence of recruitment and temperature on distribution of
intertidal barnacles in the English Channel. Journal of the Marine
Biological Association of the UK
Manuscript
Mieszkowska, N., Kendall, M.A., Richardson, A.J., Lewis, J.R. &
Hawkins, S.J. (in review) Changes in biogeographic range of the
southern trochid gastropod Gibbula umbilicalis during the current period
of climate warming. Journal of Biogeography
Manuscript
Summary of existing datasets Link website
to MarLin
52
Summary of Project Progress in Year 3 (July 2003 - June 2004).
Module Project Tasks Document
2. Archiving
databases Transfer data onto databases and provide metadata Activity report
3. Analyses of
data
Hiscock, K., Southward, A.J., Tittley, I. & Hawkins, S.J. (2004).
Effects of changing temperature on benthic marine life in Britain and
Ireland. Aquatic Conservation: Marine and Freshwater Ecosystems.
14: 333-362
Manuscript
Kendall, M.A., Burrows, M.T., Southward, A.J. & Hawkins, S.J.
(2004). Predicting the effects of marine climate change on the
invertebrate prey of the birds of rocky shore. Ibis 146(1): 40-47
Manuscript
4. Re-survey Resurvey by MarClim team
Completion of broadscale Crisp and Southward survey – Broadscale
re-survey Activity report
Kendall & Lewis Trochid re-surveys: Final report Report
5. Monitoring
network Long-term monitoring network (Oct 2003) Activity report
Long-term monitoring network (Dec 2003) Activity report
8. Communication
Mieszkowska, N., Kendall, M.A., Hawkins, S.J., Leaper, R.,
Williamson, P., Hardman-Mountford, N.J. & Hawkins, S.J. (2006).
Changes in the range of some common rocky shore species in
Britain – a response to climate change? Hydrobiologia. 555(1): 241-
251
Manuscript
End of Phase III Report Final report
53
Summary of Project Progress in Year 4 (July 2004 - October 2005).
Module Project Tasks Document
3. Analyses of
data Southward, Hawkins & Burrows barnacle and limpet datasets Final report
Recovery of a biodiversity action plan species in northwest England:
possible role of climate change, artificial habitat and water quality
amelioration
Final report
5. Monitoring
network Planning future survey and low-cost monitoring Workshop
and report
Targeted quantitative sampling at a selection of sites Final report
6. Prediction Prediction and modelling of climate change impact scenarios Activity
report
8.
Communication
Southward, A.J., Langmead, O., Hardman-Mountford, J., Aiken, J.,
Boalch, G.T., Dando, P.R., Genner, M.J., Joint, I., Kendall, M.A.,
Halliday, N.C., Harris, R.P., Leaper, R., Mieszkowska, N., Pingree,
R.D., Richardson, A.J., Sims, D.W., Smith, T., Walne, A.W. & Hawkins,
S.J. (2005). Long-term oceanographic and ecological research in the
western English Channel. Advances in Marine Biology 47: 1-105
Manuscript
Thompson, R.C., Crowe, T.P. & Hawkins, S.J. (2002). Rocky intertidal
communities: past environmental changes, present status and
predictions for the next 25 years. Environmental Conservation. 29(2):
168-191.
Manuscripts
Moore, P., Hawkins, S.J. & Thompson, R.J. (in review). The role of
biological habitat amelioration in altering the relative responses of
congeneric species to climate change. Marine Ecology Progress Series
Mieszkowska, N., Boalch, G. & Hawkins, S.J. (in prep). Changes in the
biogeographic range of Bifurcaria bifurcata in the English Channel.
9. Policy
application Strategic policy issues arising from data and climate change scenarios Interim report
Marine climate change indicator species and applications as
sustainability indicators Report
Implications of predicted changes to marine biota of Britain and Ireland Final report
End of Phase IV Report Final report
