Research Ideas and Outcomes :
Workshop Report
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Corresponding author: Covadonga Orejas (cova.orejas@ieo.csic.es), Marina Carreiro-Silva (carreirosilvamarina@gmail.com), Christian Mohn (chmo@ecos.au.dk), James Davis Reimer (jreimer@sci.u-ryukyu.ac.jp), Toufiek Samaai (toufiek.samaai@gmail.com), A. Louise Allcock (louise.allcock@gmail.com), Sergio Rossi (sergio.rossi@unisalento.it)
Received: 13 Oct 2022 | Published: 11 Nov 2022
© 2022 Covadonga Orejas, Marina Carreiro-Silva, Christian Mohn, James Reimer, Toufiek Samaai, A. Louise Allcock, Sergio Rossi
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Orejas C, Carreiro-Silva M, Mohn C, Reimer JD, Samaai T, Allcock AL, Rossi S (2022) Marine Animal Forests of the World: Definition and Characteristics. Research Ideas and Outcomes 8: e96274. https://doi.org/10.3897/rio.8.e96274
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The term Marine Animal Forest (MAF) was first described by Alfred Russel Wallace in his book “The Malay Archipelago” in 1869. The term was much later re-introduced and various descriptions of MAFs were presented in great detail as part of a book series. The international research and conservation communities have advocated for the future protection of MAFs and their integration into spatial plans and, in response, there are plans to include the characteristics of MAFs into national policies and international directives and conventions (i.e. IUCN, CBD, OSPAR, HELCOM, Barcelona Convention, European directives, ABJN policies etc.). Some MAF ecosystems are already included in international and national conservation and management initiatives, for instance, shallow water coral reefs (ICRI, ICRAN) or cold-water coral reefs and gardens and sponge aggregations (classified as Vulnerable Marine Ecosystems, VMEs), but not as a group together with other ecosystems with similar ecological roles. Marine Animal Forests can be found in all oceans, from shallow to deep waters. They are composed of megabenthic communities dominated by sessile suspension feeders (such as sponges, corals and bivalves) capable of producing three-dimensional frameworks with structural complexity that provide refuge for other species.
MAFs are diverse and often harbour highly endemic communities. Marine animal forests face direct anthropogenic threats and they are not protected in many regions, particularly in deep-sea environments. Even though MAFs have been already described in detail, there are still fundamental knowledge gaps regarding their geographical distribution and functioning. A workshop was dedicated to clarifying the definition of MAFs, characterising their structure and functioning, including delineating the ecosystem services that they provide and the threats upon them. The workshop was organised by Working Group 2 of the EU-COST Action “MAF-WORLD” (hereafter WG2), which is responsible for collating and promoting research on mapping, biogeography and biodiversity of MAFs, to identify and reduce these knowledge gaps. Herein, we report on this workshop and its outputs.
Marine Animal Forest, structural role, functionality, ecosystem services, threats, conservation, policy
Ricardo Aguilar, Suchana Apple Chavanich, Nur Arafeh Dalmau, Genuario Belmonte, Meri Bilan, Simona T . Boschetti, Lorenzo Bramanti, Carlo Cerrano, Giovanni Chimienti, Federica Constantini, Thanos Dailianis, Vianney Denis, Julian Evans, Charlotte Havermans, Paulo Horta, Kerry Howell, Georgios Kazanidis, Darina Koubínová, Piotr Kuklinski, Vesna Mačić, Francesco Mastrototaro, Monica Montefalcone, Declan Morrisey, Marco Palma, Alexa Parimbelli, Slavica Petović, Massimo Ponti, Maria Rakka, Buki Rinkevich, Maria Salomidi, Maria Sini, Eva Turicchia, Şükran Yalçin Özdilek
The workshop was held online on 10 March 2022.
In total, 41 participants from 19 different countries with diverse professional backgrounds and affiliations participated in the workshop. Dr. Nur Arafeh Dalmau, an external expert with experience in the policy-science interface, also attended the Workshop, while Prof. David Johson (GOBI, Seascape consultant) and Dr. Ellen Kenchington (DFO, Canada), both of whom have extensive expertise in science and policy, provided comments on the workshop’s results. The majority of the participants were from European countries, but there were also participants from Brazil, Japan, Thailand, Australia, Taiwan and South Africa (Table
List of participants (in alphabetical order) in the workshop, including affiliation and country.
Participant name |
Affiliation |
Country |
|
1 |
Ricardo Aguilar |
OCEANA |
Spain |
2 |
Louise Allcock |
NUI Galway |
Ireland |
3 |
Suchana Apple Chavanich |
Chulalongkorn University |
Thailand |
4 |
Nur Arafeh Dalmau |
University of Queensland |
Australia |
5 |
Genuario Belmonte |
University of Salento |
Italy |
6 |
Meri Bilan |
University of Salento |
Italy |
7 |
Simona T. Boschetti |
Joint Research Center - EC consultant |
Italy |
8 |
Lorenzo Bramanti |
Observatoire Oceanologique Banyuls sur Mer |
France |
9 |
Marina Carreiro Silva |
University of the Azores |
Portugal |
10 |
Carlo Cerrano |
Universita Politecnica delle Marche |
Italy |
11 |
Giovanni Chimienti |
Department of Biology, University of Bari |
Italy |
12 |
Federica Costantini |
University of Bologna |
Italy |
13 |
Thanos Dailianis |
Hellenic Centre for Marine Research |
Greece |
14 |
Vianney Denis |
National Taiwan University |
Taiwan |
15 |
Julian Evans |
University of Malta |
Malta |
16 |
Charlotte Havermans |
Alfred-Wegener-Institute |
Germany |
17 |
Paulo Horta |
Universidade Federal de Santa Catarina |
Brazil |
18 |
Kerry Howell |
University of Plymouth |
UK |
19 |
Georgios Kazanidis |
University of Edinburgh |
UK |
20 |
Darina Koubínová |
University of Neuchâtel |
Switzerland |
21 |
Piotr Kuklinski |
Institute of Oceanology, Polish Academy of Sciences |
Poland |
22 |
Vesna Mačić |
University of Montenegro |
Montenegro |
23 |
Francesco Mastrototaro |
University of Bari |
Italy |
24 |
Christian Mohn |
Aarhus University |
Denmark |
25 |
Monica Montefalcone |
University of Genoa |
Italy |
26 |
Declan Morrissey |
NUI Galway |
Ireland |
27 |
Covadonga Orejas |
IEO-CSIC |
Spain |
28 |
Marco Palma |
Universita Politecnica delle Marche |
Italy |
29 |
Alexa Parimbelli |
NUI Galway |
Ireland |
30 |
Slavica Petović |
University of Montenegro |
Montenegro |
31 |
Massimo Ponti |
University of Bologna |
Italy |
32 |
Maria Rakka |
University of the Azores |
Portugal |
33 |
James Reimer |
University of the Ryukyus |
Japan |
34 |
Buki Rinkevich |
National Institute of Oceanography |
Israel |
35 |
Sergio Rossi |
University of Salento |
Italy |
36 |
Maria Salomidi |
Hellenic Centre for Marine Research |
Greece |
37 |
Toufiek Samaai |
University of the Western Cape |
South Africa |
38 |
Maria Sini |
University of the Aegean |
Greece |
39 |
Eva Turicchia |
University of Bologna |
Italy |
40 |
Şükran Yalçin Özdilek |
Canakkale Onsekiz Mart University |
Turkey |
Marine Animal Forests (MAF) is a term used to describe some of the world's most biologically important (i.e. extremely diverse, productive and distinctive) yet frequently under-studied marine benthic habitats. Marine Animal Forests are benthic ecosystems dominated by megabenthic invertebrates (> 1 cm), mostly suspension feeders, that form three-dimensional frameworks of different sizes and sometimes canopies*
In their structural role and partially in their functionality, these communities resemble the structure of terrestrial forests (Fig.
Some examples of Marine Animal Forests in Danish waters: (A) Community of Porifera with scattered red algae in the area of the Great Belt bridge, (B) Common seastar (Crossaster papposus) with assemblages of sea urchins (Strongylus centrotus) and polychetae (Pomacoteros triqueter) at Schultz’s Grund in the Southeast Kattegat, (C) Reef structure made by bubbling methane of Frederikshavn in Kattegat dominated by the sea anemone Metridium senile, (D) Community of soft corals, hydrocoans and the red algae Phycodrys rubens (Kattegat, ‘Chinese Wall’, 18 m depth). Images were kindly provided by Karsten Dahl (Aarhus University; Copyright Karsten Dahl with permission to publish under the terms of the Creative Commons Attribution 4.0 License).
Due to their high vulnerability to anthropogenic impacts, some MAFs, such as shallow water coral reefs and deep-sea habitats like cold-water coral reefs and gardens and sponge aggregations, are already included in conservation and management initiatives. Shallow coral reefs have been particularly impacted by anthropogenic impacts due to their proximity to shore and human population dependency on the resources they provide. They are also one of the most species diverse marine ecosystems and as such protected under several international initiatives (e.g. International Coral Reef Initiative (ICRI) and International Coral Reef Action Network (ICRAN)). Deep-sea species have particular life history characteristics, such as long-lived, slow growth and low reproductive output that hampers their recovery from human disturbance. Concerns raised on the impacts of fishing activities (specially bottom trawling) and long recovery times of these animals have resulted in the consideration of the habitats they form (e.g. cold-water coral reefs and gardens, sponge aggregations) as Vulnerable Marine Ecosystems (VMEs) (UNGA Resolution 61/105,
As such, one aim of Working Group 2 (WG2: Underwater mapping, biogeography and biodiversity) within the “MAF-WORLD” COST Action is to produce a list of criteria helpful to identify which ecosystems may be defined as MAFs for consideration in national, regional and international management and conservation policies, thus providing a common basis to draw attention to their importance.
The workshop's main goal was to bring together marine specialists from around the world and gather information and expertise from them to better define MAFs. We particularly focused on the aims below:
Covadonga Orejas and Marina Carreiro Silva led the online event on 10 March 2022, with Toufiek Samaai, Christian Mohn, James Reimer and Louise Allcock providing assistance. Christian Mohn provided technical support for the workshop, which took place via the Zoom interactive platform. Topics 2, 3 and 4 were run in parallel sessions in separate Zoom breakout rooms (Topic 2: C. Orejas as moderator and T. Samaai as rapporteur; Topic 3: L. Allcock as moderator and M. Carreiro-Silva as rapporteur; Topic 4 Christian Mohn as moderator and James Reimer as rapporteur), with discussions, examples, experiences, terms and criteria delivered using the Zoom interactive platform. A final general discussion and poll completed the workshop.
The agenda consisted of a general introduction that provided background to the workshop, followed by discussions about the four major topics in breakout groups. The list of topics, as well as a background document, was shared with all participants before the workshop. The assigned moderators and rapporteurs guided and made notes of all the discussions and these were collated and used to produce the current report. The four topics with key aspects as guidance for discussions are listed below.
Topic 1: Defining criteria “what are MAFs? vs. Ecologically and Biologically Significant Areas (EBSAs) and Vulnerable Marine Ecosystems (VMEs)”
Topic 2: Structural and functional roles of MAFs
Topic 3: MAFs as suppliers of ecosystem services
Topic 4: Threats to MAFs
Agenda timing
Time (CET)
Activity
10:00 – 10:10 Short introduction, goals.
10:10 – 10:40 Round 1 – Discuss Topic 1 (all participants work in breakout groups).
10:40 – 10:55 Rapporteurs present the major outcomes from discussions in each breakout group.
11:00 – 11:30 Round 2 – Discuss Topics 2, 3 & 4 (all participants work in breakout groups).
11:30 – 12:00 Share reports from each breakout group.
Lunch Break
13:30 – 14:00 Round 2 – Discuss Topics 2, 3 & 4 (all participants work in breakout groups).
14:00 – 15:00 Final discussions Topic 1. Finding consensus on criteria defining MAFs at the end of the workshop via poll and discussion.
15:00 Close of the workshop.
Key outcomes of discussions for Topic 1
The first session considered the descriptions of MAFs included in the book from
For the discussions, three potential descriptions of MAFs were considered. Lack of data to quantify the dominant habitat forming species and variation in size and density of dominating species in different MAFs led to a focus on functionality as a defining characteristic. Specific characteristics, such as “unique”, “rare”, “fragile” and “long-lived” were not accepted as applying to all MAFs. Redundancy of terms used in previous descriptions was also discussed: there was a preference for a short, simple and clear definition that would be accessible to policy-makers and non-experts.
When discussing criteria that best define MAFs, workshop participants were generally of the opinion that a description should give particular emphasis to the role of MAFs as ecosystem engineers and to the importance of their three-dimensional structure in providing shelter and functional complexity to other species. The size, geographical extent and depth ranges were not considered as suitable criteria for distinction of MAFs, but it was considered that the MAF description should mention that MAFs may encompass differently-sized organisms and occur over wide depth and geographic ranges.
It was noted that MAFs:
Key outcomes of discussions in breakout sessions Topics 2 – 4
Topic 2
The structural and functional roles of MAFs were discussed in Topic 2. The key elements, including three-dimensional structure, in the make-up of a MAF were discussed, as well as how MAFs may be quantified. We debated whether to include canopy, as defined in terrestrial forests, in the MAF description. It was decided to exclude it from the MAF criteria due to a lack of understanding of the term canopy in terms of animal forest.
In addition, some MAFs do not form a true “canopy” (e.g. sea pens, mussel beds, sponge aggregations), but still substantially increase three-dimensional complexity and have important functional roles as habitat providers or in nutrient and carbon cycling. Participants discussed "what species make a MAF" because it is inevitable that species and habitats must be conserved.
The role of microbes (microbiomes) was discussed, giving consideration to the concept of the holobiont, the assemblage of a host and the many other species living in or around it, which together form a discrete ecological unit. Within this framework, we further discussed ecosystems services including Blue Carbon budgets and carbon sequestration, generation of microclimates, roles in the bentho-pelagic coupling and biogeochemical cycles, as well as the importance of considering any MAF definition also encompassing ephemeral animal forests.
Microbial actions, benthopelagic coupling, biogeochemical cycling and microclimate modifications were considered to be important functions of MAFs. The scientific debate about whether MAFs can be considered Blue Carbon ecosystems similarly to coastal wetlands (seagrasses, salt marshes, mangroves) is still ongoing. Certain MAFs, such as tropical coral reefs, sequester CO2 in an indirect manner, whereas others do not. In calcifying species, such as mussels and corals, the majority of the carbon is retained in their carbonate skeletons as opposed to being sequestered through organic C and fluxes.
The last topic of discussion was what makes MAFs unique. Based on group discussion, the following are some of the main defining characteristics we consider need to be included when generating a set of criteria:
Topic 3
We discussed ecosystem services initially under the Common International Classification of Ecosystem Services (CICES) framework. The group identified and discussed different categories of services, including provisioning, regulating, cultural and supporting services. Most of the identified services were associated with well-studied or emblematic MAFs, such as shallow-water coral reefs, oyster reefs and mussel beds. The group recognised the need to identify and describe services and functions of less well-known MAFs (e.g. bryozoans, hydrozoans) in future efforts. The group also discussed the need to better value cultural services provided by MAFs, particularly their aesthetic importance in terms of spiritual value and human well-being. People will better support the need to preserve and protect MAFs if they can spiritually connect with them. Images of MAFs available through educational materials and websites may be very useful tools. We agreed that it is important to determine who benefits from these ecosystem services and who loses when any of these ecosystem services are lost. A list of ecosystem services provided by MAFs was produced.
Provisioning services:
Regulation and maintenance:
Supporting services:
Natural-cultural heritage:
We discussed the function of MAFs in terms of habitat cascades, recognising that a habitat becomes a MAF when the animals within it are sufficiently dense to function as one. We acknowledged that not all MAFs provide the same ecosystem services. In order to provide clear guidance to policy-makers and conservation organisations, there is a need to identify knowledge gaps and conduct further research to describe the ecosystem services which different MAFs provide. The group identified it as important to describe ecosystem services and their functional role by MAF type, depth (shallow, mesophotic, deep sea) and region. Blue Carbon is a particular area of importance in this regard that need to be explored. Determining which MAFs can be considered carbon sinks and over what time scales should be a priority, although it should be taken into account that there currently is a debate on the potential role of MAFs as carbon sinks.
Finally, we discussed the stable environments (although some are also ephemeral) provided by some MAFs and their importance as providers of new biodiversity across evolutionary time scales.
Topic 4
Possible threats and their effects and impacts on MAFS were discussed in Topic 4. In addition, existing EU and other key policies addressing potential threats were identified.
What are the threats to MAFs?
We gathered a list of a wide variety of threats to MAFs detailed below:
What are the effects of these threats to MAFs?
How can these threats affect the diversity, structure and function of MAFs?
EU and other key policies
Key outcomes of final discussion of the breakout session Topic 1
The last session was devoted to a group participatory discussion, focused on delineating the criteria and description of MAFs. As well, there was discussion on overcoming obstacles and moving forward from this workshop.
The Chair of this session prepared the new description of MAFs, based on the advice and opinions from the morning breakout groups on this topic.
The MAF description, as defined in
1. Megabenthic invertebrates that build three-dimensional (3-D) living structures, often including fragile and long-living species, that support high levels of biodiversity and are found throughout the world’s oceans, at all depths from polar to tropical areas (
2. MAFs are biogenic habitats created by megabenthic faunal species acting as autogenic ecosystem engineers that change the environment by creating a complex, 3-D biophysical structure. The high biophysical complexity of MAFs provides new ecological niches and additional surface area for colonisation by associated biota, resulting in an elevated species diversity compared to adjacent areas with similar seabed geomorphology, but lacking MAFs.
3. Benthic communities characterised mostly, but not exclusively, by suspension and filter feeder animals (e.g. sponges, anthozoans, bivalves, hydrozoans, bryozoans) that build 3-D structures, similar to trees, bushes or meadows in a terrestrial forest/ecosystem (
The MAF workshop resulted in a modified description.
New description of MAF: 3D structure formed by benthic animals acting as autogenic ecosystem engineers which provides new ecological niches and colonisation surfaces for other organisms and results in increased provision of functions and services.
This description of MAFs, while developed ‘de novo’ during the workshop, encompasses two out of five criteria used to define Vulnerable Marine Ecosystems, namely structural complexity and functional significance (
A live poll of all participants determined that 76% (= 26/34) of participants favoured this new definition (Table
● We organised and conducted an internationally attended workshop that reached a broad audience with > 40 participants from 19 countries.
● Participants were experts in various fields of MAF biology (i.e. benthic invertebrate ecologists, taxonomists, marine scientists and policy-makers) and this allowed a broad discussion on what MAFs are and are not.
● MAFs have ecological importance and occur from shallow water to the deep oceans and from the polar regions to the tropics.
● We developed a set of criteria and an operational definition of MAFs.
● Participants agreed that it will be important for policy-makers to understand MAFs and their roles in the marine environment.
This publication is based upon work from COST ACTION Marine Animal Forests of the World (MAF-WORLD), CA20102, supported by COST (European Cooperation in Science and Technology). We would like to express our sincere gratitude to our institutions for allowing us to host and attend the MAF workshop. This MAF workshop was a multi-institutional collaborative effort and we are thankful to all participants for taking the time to attend. The workshop would not have been possible without the participants and we thank them for engaging with us on the MAF subject. C. Orejas is grateful to the Hanse Wissenschaftskolleg – Institute for Advanced Study for the wonderful opportunity to spend ten months with a fellowship, which has allowed her to contribute to the development of new scientific ideas and projects, such as the MAF COST action. M. Carreiro-Silva acknowledges funding from Program Stimulus of Scientific Employment (CCCIND/03346/2020) from the Fundação para a Ciência e Tecnologia.
Many thanks to Ellen Kenchington, David Johnson and Nur Arafeh for acting as external reviewers of both the workshop (Nur Arafeh) and workshop report (Ellen Kenchington, David Johnson and Nur Arafeh) and helping to improve the WS outcomes with their suggestions and experience.
The forest canopy is a structurally complex and ecologically important subsystem of the forest. It is defined as “the aggregate of all crowns in a stand of vegetation, which is the combination of all foliage, twigs, fine branches, epiphytes as well as the interstices (air) in a forest” (