THE MID-PALAEOZOIC BIOTIC CRISIS: SETTING THE TRAJECTORY OF TETRAPOD EVOLUTION

Lead Research Organisation: University of Cambridge
Department Name: Zoology

Abstract

This project will shed light on a key stage in the evolution of life on Earth. The advent onto land of limbed vertebrates (tetrapods) was an event that shaped the future evolution of the planet, including the appearance of humans. The process began about 360 million years ago, during the late Palaeozoic, in the early part of the Carboniferous Period. Within the 20 million years that followed, limbed vertebrates evolved from their essentially aquatic and fish-like Devonian predecessors into fully terrestrial forms, radiating into a wide range of body forms that occupied diverse habitats and ecological niches. We know this because we have an adequate fossil record of the earliest limbed vertebrates from the Late Devonian, contrasting with the terrestrial forms that lived significantly later in the Early Carboniferous, about 340 million years ago. It is also clear that a mass extinction event occurred at the end of the Devonian, following which life on land and in fresh water habitats had to be re-established. Unfortunately, the formative 20 million years from the end of Devonian times has remained almost unrepresented for fossil tetrapods and their arthropod contemporaries. Thus, we know little about how tetrapods evolved adaptations for life on land, the environments in which they did so, and the timing or sequence of these events. The evolutionary relationships among these early tetrapods and how they relate to modern forms are also unclear and controversial as a result of this lack of fossil information. The entire fossil hiatus has been called 'Romer's Gap' after the American palaeontologist who first recognized it. Now, for the first time anywhere in the world, several fossil localities representing this period have been discovered in south-eastern Scotland. They have already provided a wealth of new fossils of tetrapods, fish, invertebrates and plants, and our team is the first to have the opportunity to study this material and the environmental, depositional, and climatic context in which this momentous episode took place. We have a number of major aims. The existing fossil material will form a baseline for this study, but the project will augment this by further excavating the richest of the sites so far found and subjecting it to a detailed archaeological-style analysis. We will collect from other recently recognized sites and explore for further sites with relevant potential. The fossil material will be described and analysed using a range of modern techniques to answer many questions related to the evolution of the animals and plants. Not only that, using stratigraphical, sedimentological, palynological, geochemical and isotopic data, we will establish the conditions of deposition that preserved the fossils, the environments in which the organisms lived and died, and the precise times at which they did so. We will drill a borehole that will core through the entire geological formation in which these fossils have been found. Using this, we will integrate data from our fossil sites using the signals provided by the sedimentary record to build a detailed time line showing in which horizons the fossils were found, the age of each occurrence and their sequential relationship. We will compare and correlate our data with that from contemporaneous deposits in Nova Scotia, the only other locality with information sufficiently rich to be meaningful. Our data will allow us to infer changes to the environment and the evolutionary trajectories of the animals and plants during the deposition of this formation, covering the 20 million years following the end-Devonian mass extinction. Comparison with similar data for the Late Devonian will allow us to chart the changes around the time of the mass extinction, to infer its causes and consequences, and obtain a detailed record of exactly how changes to the environment correlated with changes to the fauna and flora.

Planned Impact

The earliest Carboniferous (Tournaisian) interval (360-340 Ma) has long formed a 'bottleneck' in studies of the evolution of terrestrial ecosystems, and particularly the earliest evolution of terrestrial tetrapods, because of the almost complete lack of fossil evidence from this key time. We are now able to populate this hiatus in fossil data as a result of recent discoveries in Scotland, and to place these specimens in a palaeoecological and stratigraphic context.
Given the current paucity of data previously available for study of the faunas and floras from this time, and the profound changes to terrestrial ecosystems that took place then, we fully expect our results to be literally and metaphorically ground breaking. The material we will discover and describe will be of international significance, and enhance the reputation of the UK as a centre of excellence for Palaeozoic tetrapod fossils and their study. We anticipate publication in high profile journals including Science and Nature. Our palaeontological results will benefit interrelated disciplines from anatomists through molecular phylogeneticists to palaeoecologists. Biomechanics of locomotion, feeding, and breathing will gain from the new insight into the basal anatomical conditions at the onset of terrestriality. Studies of the evolutionary development of skeletal systems, and the timing of key innovations in tetrapod morphological adaptations will use our findings. Molecular phylogenies will benefit from new calibration points for the origins of the tetrapod and actinopterygian crown groups. The geological aspects of the proposal will benefit those modelling ancient climates in deep time, environmental and sedimentary systems, and their influence on and relationship to key evolutionary events. The refined stratigraphical, isotopic and palynological data that our studies provide will augment the, so far, relatively poorly known picture of this key period. Our comprehensive dataset from the earliest Carboniferous of Scotland will provide a future standard of comparison for contemporary deposits in other parts of the world. The industry-standard geophysical log data will be of interest to petrophysicists in academia and their industry partners, from including civil engineers, the construction industry and the hydrocarbon industries, who are studying the characterisation and prediction of physical properties in UK rock formations. As a team, we are particular well placed for high impact in the press and other media, with two members situated in nationally and internationally renowned museums, and with close links to others. Both institutions have professional in-house officers who deal with outreach on a continuing basis. Our experience talking to media representatives, the general public and other non-academics including people local to our sites, suggests that this project is of wide public interest and appeal.

Publications

10 25 50
 
Title Fossil Hunters: Unearthing the story of life on land 
Description An exhibition based on our project from NE/J022713/1 and associated consortium grants opened at the National Museum of Scotland (see also Engagement Activities) included video footage, interactive displays and models generated for the exhibition 
Type Of Art Film/Video/Animation 
Year Produced 2016 
Impact Six months after the opening on February 19th 2016, we should have attendance figures, but results from the first few days have been encouraging. 
URL http://www.nms.ac.uk/about-us/research/research-now/the-tweed-project-evolution-s-missing-chapter/
 
Description A paper was published in Nature Ecology and Evolution describing new tetrapod taxa and their environment. We also show that oxygen levels in the Tournaisian were similar to those of the preceding Famennian and the succeeding Visean, indicating that and a proposed oxygen low at that time did not exist, and did not hamper the emergence of tetrapods. We were were also able to interpret the environment as an alternating drier and wetter one in a flood plain mosaic of rivers, lakes, ponds and dry land. This could have encourage the evolution of land-based life in tetrapods. We also noted a correlation between palaeosols, the overlying sandy silt ones from over bank flood deposits, suggest a relationship between land living and the tetrapods. We have also published papers on the arthropods, terrestrial palaeoenvironments, sedimentology and ichnology of the Ballagan Formation. The paper in Nature Ecology and Evolution is now highly cited with an Am score of 258.
One outcome not envisaged in the original proposal was the discovery of at least 21 new taxa of chondrichthyan (see below).
We have published further papers on vertebrates from the Tournaisian of of Scotland, explaining new insights from our studies. These include: the way tetrapods improved their efficiency to walk on land; discovery of tetrapod remains from a taxon formerly encountered only 20 million years later; the unexpectedly diverse array of chondrichthyan taxa from the Tournaisian; two major papers focusing two separate beds from the Bournemouth locality, showing the richness of the vertebrate fauna in each, and its faunal, sedimentological and environmental context.
A paper has been accepted on a new and very large lungfish taxon from about 30m above the Devonian-Carboniferous boundary, which shows a combination of typically Devonian and typically Carboniferous lungfish features.
Exploitation Route We hope to explore the next stage in the geological column, the early Visean. We also hope that others will now explore Tournaisian rocks for tetrapods elsewhere in the world. To that end, we have published a paper describing a new, specialised tetrapod from the Vision of Fife, that brings down the origin of the group to which it belongs by more than 3 million years. It comes from a newly discovered locality with great potential for exploring the Visean stage. We have a paper in review on the earliest heterodont and durophagous tetrapod, found in Visean rocks.
Sectors Education,Environment,Culture, Heritage, Museums and Collections

URL http://www.tetrapods.org
 
Description Part of the special exhibition which opened on February 19th at the National Museum of Scotland, placing our findings in the context of tetrapod evolution "Fossil Hunters: unearthing the mystery of life on land" was exhibited by the Biggar and Upper Clydesdale Museum 15th October 2016- 15th January 2017, and at the Montrose Museum 21st January - 1 April 2017. Our collections of fossils form the core of the exhibits and their interpretation. There are also videos and life reconstructions showing the context and importance of our material. The exhibition will travel to Stornoway 13th April - 24th June 2017. Project newsletter sent to 122 non academic subscribers; Facebook pages seen by between 300 and 2851 visitors per posting.
Sector Education,Culture, Heritage, Museums and Collections
Impact Types Cultural

 
Description Leverhulme Emeritus Fellowship
Amount £21,925 (GBP)
Organisation The Leverhulme Trust 
Sector Academic/University
Country United Kingdom
Start 01/2017 
End 04/2018
 
Description Expert on Carboniferous crustaceans 
Organisation University of Glasgow
Department Hunterian Museum and Art Gallery
Country United Kingdom 
Sector Academic/University 
PI Contribution Invited individual with expertise in identifying and describing early Carboniferous crustaceans and providing material collected by the consortium.
Collaborator Contribution Help with identifying and describing early Carboniferous crustaceans discovered by the consortium
Impact Material added to collections of museums in Cambridge and Edinburgh
Start Year 2014
 
Description Project partner 
Organisation University of Calgary
Country Canada 
Sector Academic/University 
PI Contribution Worked together at fossil locality to examine and describe new tetrapod limb elements from the Tournaisian site of Blue Beach in Nova Scotia. Travel to Nova Scotia and Halifax from Cambridge for discussions with partner.
Collaborator Contribution Jointly published a paper describing new tetrapod limb elements from the Tournaisian site of Blue Beach in Nova Scotia. Travel to Nova Scotia and Halifax from Calgary for discussions with partner.
Impact Paper published in PLoSOne
Start Year 2013
 
Description 4.Populating Romers' Gap : rebuilding terrestrial ecosystems after the end-Devonian mass extinction 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Regional
Primary Audience Public/other audiences
Results and Impact A talk introducing our project, to the Glasgow Geological Society. T. Neville George annual lecture.

The annual T. Neville George lecture: awarded the T. Neville George medal.
Year(s) Of Engagement Activity 2013
URL http://www.tetrapodworld.com
 
Description A new tetrapod world: laying the foundations: talk to Chirnside community 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Public/other audiences
Results and Impact Visits to our fossil excavation in the river Whiteadder at Willie's Hole, near Chirnside, Borders Region, by local teachers, archaeologists and fishermen.

Interest in continuing contact with the TW:eed project
Year(s) Of Engagement Activity 2015
URL http://tetrapods.org
 
Description Article describing the results of our project THE MID-PALAEOZOIC BIOTIC CRISIS: SETTING THE TRAJECTORY OF TETRAPOD EVOLUTION in Discover Magazine 
Form Of Engagement Activity A magazine, newsletter or online publication
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Discover Magazine published an article describing our project findings that appeared in Nature Ecology and Evolution
Year(s) Of Engagement Activity 2017
 
Description Atlantic Geoloscience Society Annual Meeting 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Type Of Presentation poster presentation
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact Aroused interest in early Carboniferous sites in Nova Scotia

The poster was later put up in the Nova Scotia Natural History Museum in Halifax, NS.
Year(s) Of Engagement Activity 2015
URL http://ags.earthsciences.dal.ca
 
Description Bridging Romer's Gap 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Other audiences
Results and Impact This was a talk to a Further Education College, in Sunderland. There is no slot for a FE audience group.
Year(s) Of Engagement Activity 2015
 
Description Chirnside community lecture: A new tetrapod world - laying the foundations. 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Public/other audiences
Results and Impact A talk to the local Chirnside community about the work of the TW:eed project, and the ongoing excavations at Willie's Hole on the nearby River Whitadder
Year(s) Of Engagement Activity 2015
URL http://tetrapods.org
 
Description Diversity among new Tournaisian tetrapods from Scotland 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Type Of Presentation paper presentation
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact A presentation to the annual meeting of the Palaeontological Association, Zurich

Interest from non-vertebrate-palaeontologists
Year(s) Of Engagement Activity 2013
 
Description Keynote speaker for Birmingham Biosciences Seminar, July 2017 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach National
Primary Audience Undergraduate students
Results and Impact "From Bugs in Birmingham to Beasts in the Borders: a charmed life in palaeontology. Describing my experiences in getting to where I am today.
Year(s) Of Engagement Activity 2017
 
Description Life, death and resurrection in the Late Palaeozoic 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Schools
Results and Impact Annual Bennett lecture given by J. A. Clack. About 100 school children (16-19) attended, plus other local audiences. Part of an open day at the University of Leicester.

Many school children attended a day-long session on vertebrates of the Carboniferous and enjoyed examining specimens of microfossils from the collections made by the consortium
Year(s) Of Engagement Activity 2014
URL http://www.tetrapods.org
 
Description Life, death and resurrection in the Late Palaeozoic 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Type Of Presentation keynote/invited speaker
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact A talk to staff and students at the Organismal Biology department of the University of Chicago

Part of a celebration of my being awarded an Honorary Doctorate of Science from the University of Chicago.
Year(s) Of Engagement Activity 2012
URL http://www.tetrapods.org
 
Description Life, death and resurrection in the Late Palaeozoic 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach Regional
Primary Audience Public/other audiences
Results and Impact A talk to the Oxford Colloquium in Geology, aimed at local enthusiasts and schools. The hour-long talk was part of a two-day activity in the University Museum.

Interest expressed by many members of the audience.
Year(s) Of Engagement Activity 2014
URL http://www.tetrapods.org
 
Description Lungfishes from Romer's Gap: experiments in tooth plate morphology 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Type Of Presentation paper presentation
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact Presentation to the annual Symposium of Vertebrate Palaeontology, Edinburgh

Offers of help with field work on the project
Year(s) Of Engagement Activity 2013
URL http://www.tetrapods.org
 
Description Manchester Geological Society. Repopulating a post-extinction world: The resurgence of vertebrate diversity in the early Carboniferous. 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Public/other audiences
Results and Impact Talk was enjoyed and appreciated by the audience of amateur and professional geologists.
Year(s) Of Engagement Activity 2016
 
Description Media interviews in Nova Scotia related to the TW:eed consortium project 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Interviews and filming for peak time main news programme in Nova Scotia, also with local politician, and filming for a documentary on the Nova Scotia Blue Beach fossil site.

We have yet to see the documentary or hear whether our interviews had any impact on local conditions.
Year(s) Of Engagement Activity 2014
URL http://www.tetrapods.org
 
Description New tetrapod and fish faunas from the earliest Carboniferous of Scotland 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Type Of Presentation paper presentation
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact Presented some of our preliminary results to an audience that included vertebrate anatomists as well as palaeontologists

Interest from representatives of high profile publications
Year(s) Of Engagement Activity 2013
URL http://www.tetrapods.org
 
Description Palaeontological and molecular approaches to early vertebrate evolution 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Type Of Presentation keynote/invited speaker
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact This interdisciplinary meeting aimed to further understanding and integration of palaeontological and molecular approaches to the study of evolution. Clack presented results of early tetrapod evolution based on new work from the TW:eed project

Enthusiastic responses from participants from other disciplines. A very successful meeting
Year(s) Of Engagement Activity 2015
URL http://www.kva.se/
 
Description Populating Romer's Gap: new arthropods, tetrapods and fishes from the Early Carboniferous of northern Britain 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Public/other audiences
Results and Impact A talk for the Cambridge Geology Club describing our project for non-specialist amateurs.

Interest from local amateur geologists
Year(s) Of Engagement Activity 2013
URL http://www.tetrapodworld.com
 
Description Populating Romer's Gap: rebuilding terrestrial ecosystems after the end-Devonian mass extinction 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Type Of Presentation keynote/invited speaker
Geographic Reach Regional
Primary Audience Professional Practitioners
Results and Impact A seminar for the Oxford Biosciences Society

Potential interest by the media
Year(s) Of Engagement Activity 2013
URL http://www.tetrapodworld.com
 
Description Populating Romers' Gap : rebuilding terrestrial ecosystems after the end-Devonian mass extinction 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Type Of Presentation keynote/invited speaker
Geographic Reach Regional
Primary Audience Public/other audiences
Results and Impact A lecture to the Leicester Literary and Philosophical Society. November 2012, describing the project

A talk to the society's members. Asked to contribute a piece to the LLPS newsletter
Year(s) Of Engagement Activity 2012
URL http://tetrapods.org
 
Description Populating Romers' Gap : rebuilding terrestrial ecosystems after the end-Devonian mass extinction 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Type Of Presentation paper presentation
Geographic Reach National
Primary Audience Public/other audiences
Results and Impact A presentation as part of the Geological Association's Festival of Geology, at UCL, November, introducing our project.

A day's conference/ workshop. Iain Stewart was another contributor.
Year(s) Of Engagement Activity 2012
URL http://tetrapods.org
 
Description Radio Wales interview 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach National
Primary Audience Media (as a channel to the public)
Results and Impact Inteview regarding the recent publication of our paper on the NERC-funded project THE MID-PALAEOZOIC BIOTIC CRISIS: SETTING THE TRAJECTORY OF TETRAPOD EVOLUTION in Nature Ecology and Evolution
Year(s) Of Engagement Activity 2016
 
Description Repopulating a post-extinction world: The resurgence of vertebrate diversity in the early Carboniferous. 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Public/other audiences
Results and Impact A talk to the Stamford and District Geological Society. Mainly consisting of older people, retired and interested in geology.
Year(s) Of Engagement Activity 2017
URL http://www.stamfordgeolsoc.org
 
Description Romer's Gap: a surprisingly productive period in vertebrate evolution 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact A presentation to the annual meeting of the Palaeontological Association in Zurich

Interest from non-vertebrate-palaeontologists
Year(s) Of Engagement Activity 2013
URL http://www.tetrapods.org
 
Description Royal Holloway college Lyell Society. A post-Hangenberg Event world: was the Tournaisian not so bad after all? 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach National
Primary Audience Undergraduate students
Results and Impact The annual Lyell Society Seminar consist of 6 speakers, this year on climate change. My talk was one of these.
Year(s) Of Engagement Activity 2017
 
Description Searching to close Romer's Gap: finding tetrapod fossils in the Early Carboniferous of northern Britain 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Other academic audiences (collaborators, peers etc.)
Results and Impact One of a series of seminars in the Zoology Department introducing members of the department to research that goes on there.

General interest from members of the Department of Zoology, Cambridge
Year(s) Of Engagement Activity 2013
URL http://www.tetrapodworld.com
 
Description TW:eed project newsletter 
Form Of Engagement Activity A magazine, newsletter or online publication
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Notable interest from amateur collectors who have contributed specimens and guidance to new localities.

New locality explored with significant finds made
Year(s) Of Engagement Activity 2012,2013,2014,2015,2016
URL http://www.tetrapods.org
 
Description Talk to Sunderland FE college: Bridging Romer's Gap 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Schools
Results and Impact An invited talk to college student about the work of the TW:eed project. Interest from students in the informal tea and snacks afterwards.

Interest from students in the informal tea and snacks afterwards, who were interested in pursuing courses in Geology.
Year(s) Of Engagement Activity 2015
 
Description The end-Devonian mass extinction, Romer's Gap and the Lilliput effect. 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Type Of Presentation paper presentation
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact Talk at a symposium: SVPCA Oxford

Interest in the Early Carboniferous generally
Year(s) Of Engagement Activity 2012
URL http://www.tetrapodworld.com
 
Description Trinity College Dublin. Repopulating a post-extinction world: The resurgence of vertebrate diversity in the early Carboniferous. 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Undergraduate students
Results and Impact A talk for staff, graduate and undergraduate students, Trinity College Dublin
Year(s) Of Engagement Activity 2016
 
Description University of Cambridge Sedgwick Club: Repopulating a post-extinction world: The resurgence of vertebrate diversity in the early Carboniferous. 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Regional
Primary Audience Undergraduate students
Results and Impact A student group for the Earth Sciences Dept, University of Cambridge expressed interest.
Year(s) Of Engagement Activity 2016
 
Description University of Hull. Repopulating a post-extinction world: The resurgence of vertebrate diversity in the early Carboniferous 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Regional
Primary Audience Postgraduate students
Results and Impact A talk for the Environmental Sciences group, staff, graduate and undergraduate students, University of Hull
Year(s) Of Engagement Activity 2013,2016
 
Description Window onto Romer's Gap 
Form Of Engagement Activity Participation in an open day or visit at my research institution
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Public/other audiences
Results and Impact Exhibition in UMZC for Science Week 2013

An exhibition of specimens illustrating our finds of fossils from the earliest Carboniferous, helping fill 'Romer's Gap'. Stayed in place until the museum closed on 1 June 2013.
Year(s) Of Engagement Activity 2013
URL http://tetrapods.org
 
Description Yorkshire Geological Society open day with presentations 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Type Of Presentation keynote/invited speaker
Geographic Reach Regional
Primary Audience Public/other audiences
Results and Impact Much interest from participants - the open day included posters as well as several invited talks, and a visit to the drill-core storage at BGS. Audience included professionals, local amateurs and other members of the general public.

Stimulated other regional groups to request follow-up talks.
Year(s) Of Engagement Activity 2015