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Lawyer, is founder and executive director of the Institute Choices. She started her career as a veterinarian working in several vet hospitals for 9 years. Since , she has worked in the reinsurance sector with emphasis on the agribusiness as senior agro underwriter for the Latin America region. Isabel has more than 10 years of experience in the insurance and reinsurance industry with focus on the agro sector and its sub lines of business such as crop, forestry, aquaculture, livestock and greenhouse.

Leonardo has over 20 years of experience in business, finance, and the social sector. He also spent 8 years at McKinsey, advising clients on strategic and operational challenges, with a focus on the telecommunications and energy sectors. He has over 15 years of professional experience in commercial, environmental and technical positions in various sectors, including agribusiness, metallurgy, health, construction, energy services, environmental consulting and environmental remediation.

His areas of expertise include supply chains, energy efficiency, corporate sustainability, project management and environmental research. Prior to the position at CAF, Mr. Maria is responsible for structuring innovative public-private mechanism to drive green investments in the Latin American region, including the development of capital markets and the role of public development. She also has more than 20 years of experience in the area of climate finance.

She also worked for more than 10 years at the Secretariat of the United Nations Framework Convention on Climate Change UNFCCC , where she was responsible for climate finance, and formed part of the team that established the institutional framework for implementing the carbon market mechanism set out in the Kyoto Protocol.

He has co-authored and contributed to numerous industry building publications. His monograph Solutions for Impact Investors: From Strategy to Implementation , is used as a roadmap for investors seeking guidance designing and implementing impact investing strategies. He also co-authored the follow up report, Evolution of An Impact Portfolio: From Implementation to Results , which demonstrates that impact investments can compete with, and at times outperform traditional asset allocation strategies while pursuing meaningful and measurable social and environmental impact results.

He has 20 years of experience, of which more than 15 years has been focused on sustainability in Europe and Latin America, supporting financial entities in the development of sustainable businesses, governments in the definition of regulatory frameworks that promote sustainability, and in the planning and construction of Zero Carbon real estate developments. Murilo Portugal Filho has a solid career in public sector economics and international organizations.

He was an adviser at the Civil Office, from to , and chief economic adviser at the General Secretary of the Presidency between and , when he coordinated the federal deregulation program. From to , Mr. Portugal was Secretary of the National Treasury. In , Mr. Portugal started his international trajectory. In March , Mr. Portugal returned to Brazil to take over the position of Executive Secretaryat the Ministry of Finance, remaining until March Laura coordinates projects in emerging markets that aim at providing scenario analysis tools and policy research to financial institutions, industry associations, supervisory authorities and policy makers.

Eighteen years of experience in environmental management and sustainable strategic planning. Since she has assumed her position as Head of Corporate Sustainability at HSBC El Salvador and Coordinator of Corporate Sustainability for Latin America, developing sustainability policies for the region and implementing global environmental education and management programs in 12 Latin American countries.

Passionate about the intersection between finance, sustainability and innovation. With almost 25 years of experience in international finance, I have held several management positions in commercial and investment banks in London, Madrid and New York, with a focus on Latin America and the Caribbean. I launched the first sustainability program for the financial sector in the region beyondBanking: banking on global sustainability ; I am a YPO, a First Mover Aspen Fellow; and I was named one of the 50 most influential women in Latin America by Latin Trade magazine.

He frequently serves as an exhibitor in national and international forums on topics related to the regulation and supervision of financial intermediaries, risk management and international standards. He is a Complementary Pension Specialist, licensed from Previc. He worked for 20 years in corporate finance. Atuou por 9 anos no ING Bank Brasil, como analista quantitativa, analista e gerente de risco de mercado.

He is a trained banker by profession and holds a Master Degree in Development Management and two Bachelor degrees. During the past years, he advised and trained ministries, regulators, banks and associations in short and long term missions. She worked for 11 years in the financial sector with different subjects: Financial Education and Social, Entrepreneurship and Environmental Risks and Opportunities, working directly with different Business Units: Investments, Credit and Insurance. Professional with strong background in risk management, compliance, process redesign and strategic projects.

He is responsible for the Swiss insurance stream, which includes projects to improve the environmental performance of mainly Swiss insurance companies and the integration of sustainability within their core business. Before joining WWF, he worked for 8 years as a fund analyst and selector at Generali Switzerland and 9 years in various positions within a Raiffeisenbank, including customer advising. Marco holds a bachelor in business administration with a specialization in Finance from the University of Applied Sciences and Arts Northwestern Switzerland.

Over the years UNEP FI has established some of the most important sustainability oriented frameworks within the finance industry, including the Principles for Responsible Investment , the Principles for Sustainable Insurance and work now underway on the establishment of Principles for Responsible Banking. Since joining in , Mr. Usher has focused on accelerating the deep integration of sustainability risks into financial practice, including addressing climate change, natural capital loss and human rights abuses, as well as building out the frameworks for positive impact finance needed to achieve the UN Sustainable Development Goals.

Usher has over twenty years of experience in the low carbon sectors, spanning technology commercialisation in Canada, solar rural electrification in Morocco and financial sector development across emerging markets. During , Mr. He was selected for a job as on-site inspector on the results of a public competitive examination. Previously, he had been a Navy Officer, after his graduation from the Naval Academy Since September , he has worked as Director of the Investor Protection and Assistance Department and has led various financial and investor education initiatives in Brazil as well as joint educational projects public and private sector for investor, journalists, professors and judges.

The focus of her research is the intersection of Finance, Sustainability and Economics. Some of her recent research projects include channeling private financial resources towards sustainable development, ecosystem services valuation; low carbon agriculture, financial risks associated with capital natural degradation in agricultural value chains; conservation finance.

Annelise has over 20 years of professional experience in feasibility studies valuation , credit analysis, portfolio management, mergers and acquisitions. Manages environmental and social risk for Santander globally. From to , was executive director of the Brazilian Recycling Commitment Cempre , an association dedicated to promoting waste recycling in Brazil.

Represented Brazil at international ISO negotiations on eco-labeling from to Conducted peer review for life cycle assessment studies. Siphonoglyph is conspicuous. No zooxanthellae present. The most important nematocyst characteristics observed in this species are large holotrichs holotrich 1 and the presence of p-mastigophores with a well-marked filament Table 1 and Figure 4. Due to the scarcity of material, we only observed nematocysts from one specimen.

Therefore, future observations could reveal new cnidae characteristics for this species. This species is found in association with the primnoid octocoral C. Only one specimen of H. Therefore, we assume that the prevalence of this association is rare. Fine mineral particles are incorporated in polyps. This species name is dedicated to another research yacht of Prince Albert I of Monaco, Princess Alice , used during his oceanographic campaigns in the Azores, and during which the homonymous seamount was discovered.

This colonial zoantharian is found at the surface of the stylasterid Errina dabneyi Figures 2I,J. Polyps are grouped by a thin coenenchyme and with a presence of encrustations in the ectoderm. There is an enlarged endodermal sphincter. Large holotrichs are present in all the tissues and there are special spirulae in most tissues.

No capitular ridges were visible. Ectoderm and outer mesogloea are encrusted with mineral particles. Mesenteries in macrocnemic arrangement, there are at least 30 mesenteries observed under the binocular. No specific number of mesenteries was possible due to scarcity of material.

Some mesenteries present well developed retractor in the middle of the mesenteries. No zooxanthellae are present. Numerous categories of nematocysts are the main character, with large holotrichs holotrich 1 in all the tissues including the presence of special b-mastigophores Table 1 and Figure 4.

It is found on the surface of the skeleton of the stylasterid E. There is no evidence of parasitic behavior. Diagnosis: Characterized by a simple mesogloeal sphincter muscle, this family includes the genera Epizoanthus Gray, , Paleozoanthus Carlgren, , and Thoracactis Gravier, The genus Palaeozoanthus has not been found or examined in detail since its original description Carlgren, , while Thoracactis topsenti Gravier, is the sole representative of its genus and is an epibiont on sponges at — meters around the Cape Verde Islands Gravier, The type genus of Epizoanthidae, Epizoanthus , includes species that have epibiotic associations with hermit crabs Ates, ; Reimer et al.

There are some cases of free-living species reported Epizoanthus lindahli Carlgren, , Epizoanthus vagus , Herberts, Polyps usually strongly encrusted with sand. In colonial species, polyps are linked by stolons or by a continuous coenenchyme. There are no symbioses with Symbiodinium zooxanthellae. Type Species: Dysidea papillosa Johnston, Diagnosis: Generally as for the family. Only one specimen is known from the genus Paleozoanthus and it was not conserved well enough for genus distinction Sinniger et al.

Furthermore, Epizoanthus is distinct from the genus Thoracactis by presenting polyps joined by a common coenenchyme, in contrast with Thoracactis which shows isolated polyps without apparent coenenchyme connection. Figures 2 — 4 , 6 , Table 1. Figure 6. Histological sections of Epizoanthus martinsae sp. A mesogloeal sphincter muscle MSM ; B retractor pennons P of the mesenteries; C tentacles musculature T ; D structure of the mesogloeal body wall showing enlarged lacunae with endodermic content.

Longitudinal section of contracted polyps at the region of the capitulum is shown in A. Cross sections at the region of the mesenteries are shown in B,D , and of the tentacles C. Nomenclatural act recorded at Zoobank: urn:lsid:zoobank. The name of this species is dedicated to Dr. Anthozoa, family Leiopathidae , fragment of approximately of 10 polyps formalin-fixed and 6 polyps ethanol-fixed , DOP Colonial zoantharian found at the surface of the black coral Leiopathes sp.

Figures 2K,L. Polyps with encrustations in the ectoderm and grouped by a thin coenenchyme forming a compact unit. It has a very short mesogloeal sphincter. Mesogloeal of the body wall has enlarged lacunae. Presence of large holotrichs in all tissues and special spirulae in the body wall. On live specimens, polyps are brown with two rows of transparent tentacles Figure 2L.

In preserved samples, dark brown polyps are connected by a thin coenenchyme growing over the Leiopathes sp. Living polyps have an oral disc 4—5 mm in diameter, column diameter 3—4 mm, height 6—8 mm; column cylindrical. In preserved specimens, contracted polyps are 3—4 mm in diameter and extend to 5—7 mm above the coenenchyme; column is cylindrical Figure 2L.

There are two rows of 14—17 transparent tentacles smaller, immature polyps can have fewer , with circular knobs at tips; length of tentacles is the same length than the expanded oral disk diameter. Pharynx is oval and whitish. Column is dark brown due to heavy encrustations of sand grains.

Encrustations are more abundant in the ectoderm, and extend into the mesoglea. Mesenteries are in a macrocnemic arrangement, distributed in 12 pairs of complete mesenteries, 12 pairs of incomplete mesenteries and several pairs of very low development mesenteries; we counted up to 31 mesenteries.

There is a very short mesogleal sphincter Figure 6A. Musculature poorly developed; retractor muscle present, with some pennons well arranged in the mesoglea Figure 6B ; the parietobasilar muscle is undistinguished. Ectodermal tentacles musculature is well developed and arranged in numerous small pennons Figure 6C. The mesogloea of the body wall presents enlarged lacunae with endodermic contents Figure 6D.

High diversity of nematocysts is characteristic in this species, the presence of large holotrichs holotrich 1 in all the tissues and the presence of small special b-mastigophores in the body wall being the key characters regarding the cnidae Table 1 and Figure 4. Found on skeletons of the black coral Leiopathes sp. In aquaria, it was found to colonize rocks and aquarium walls.

One of the four new species of zoantharians reported for the Azores was found to belong to the family Epizoanthidae and three species to the family Parazoanthidae Figure 7 , Supplementary Figures 1 , 2. Table 2. Bayesian posterior probabilities for selected clades inferred from phylogenetic trees reconstructed from different alignments: SIC alignment, indel events were coded as binary characters and appended to the existing alignment; NoSIC alignment, indel events were not additional coded and included to the existing alignment; GBlock alignment, potentially poorly aligned positions were excluded altogether from the alignment.

Figure 7. Bayesian phylogenetic tree obtained using concatenated sequences COI, 16S, and 12S and indel events coded as binary characters. Values at branches represent Bayesian posterior probabilities and bootstrap values obtained for the Maximum Likelihood tree. Nodes where Bayesian posterior probability was under 0. Inside Parazoanthidae, the first split corresponded to the separation of Isozoanthus Carlgren in Chun, from the rest of the family representatives.

Within Parazoanthidae, all octocoral-associated zoantharians grouped together in a well-supported monophyletic group distinct of Parazoanthus Haddon and Shackleton, , Umimayanthus and Antipathozoanthus species. The specimen DOP of Z. The genus Isozoanthus represented by the type species Isozoanthus giganteus Carlgren in Chun, 12S, 16S and Isozoanthus sulcatus Gosse, COI concatenated in a single sequence Supplementary Table 1 represented by a well-defined branch formed after a deep first split within Parazoanthidae.

This branch is well separated from Z. Phylogenetic analyses also indicate that the zoantharian associated with the primnoid octocoral Candidella imbricata constitutes a new species within the recently described genus Hurlizoanthus of the family Parazoanthidae Sinniger et al.

The Azores are considered a CWC biodiversity hotspot, particularly with regards to the sub-class Octocorallia e. Comparably less is known about the epifaunal organisms associated to octocorals and other coral taxa and the biological interactions symbiosis, competition, predation between these two groups of organisms Buhl-Mortensen et al.

Within this context, the present study contributes to increase the knowledge on the diversity of the zoantharians associated with CWCs, with the description of four new zoantharian species, and the nature of their association to their coral host. Similar to the pattern of increased biodiversity in azooxanthellate corals species in deeper waters Cairns, ; Watling et al. This diversity may be under-evaluated when descriptions are based on morphology only.

Indeed, genetic tools can help to distinguish between cryptic species that are species with similar morphologies but strong genetic differences. Phylogenetic reconstruction using three mitochondrial DNA markers COI, 16S, and 12S provided well-supported trees resolving the relationship within Parazoanthidae and Epizoanthidae families Figure 7 , with topology supported by ecological and morphological characteristics Sinniger et al.

These relationships were best resolved by concatenating the three markers and using phylogenetic information carried by indel events BI. This type of information was shown to be very important for zoantharian phylogeny as it allows distinguishing among some genera Sinniger et al. Strong phylogenetic signals were provided by the variable regions within the 16S and 12S fragments, portions of the genes reported in Sinniger et al.

Furthermore, our phylogeny confirms: 1 agreement with the re-classification and erection of the genus Hydrozoanthus as proposed by Sinniger et al. The genus Zibrowiu s, recently described from Hawaii Sinniger et al.

Carreiro-Silva et al. The re-classification of the latter species, which was originally classified as Isozoanthus , is in agreement with the revised morphology, anatomy and molecular phylogeny. This re-classification is supported by the association with a primnoid octocoral as is also the case with Hurlizoanthus hirondelleae sp.

The identification of Z. Unreliability of the morphological characters traditionally used to distinguish Isozoanthus i. The latter is a symbiotic zoantharian which was first described as belonging to the genus Isozoanthus but it was more related to Parazoanthus s. In an effort to integrate modern molecular data with traditional morphological research, Swain et al.

These authors identified traditionally targeted fifth mesenteries, marginal muscle arrangement, encircling sinus and novel fissure morphology, basal canals of the mesenteries morphological features, as the most important characters necessary for reunification and revision of Zoanthidea systematics. Useful morphological characters identified for the genus Isozoanthus included fissured capitulum and hypertrophic retractor muscles of the microdirectives, which should be targeted in future studies.

In addition, the characterization of the cnidome in Isozoanthus could also aid in the correct identification of the genus. The newly described zoantharian Zibrowius alberti sp. Molecular data indicated that this species is more closely related to Zibrowius ammophilus although in its appearance polyp size, color, and presence of particles in the body wall it is more similar to Z.

The genus Hurlizoanthus is characterized by zoantharians associated with the primnoid octocorals. Adding Hurlizoanthus hirondelleae sp. Furthermore, the color of H. In the original description of the genus Hurlizoanthus the diagnostic character was the association with primnoid corals. According to the most recently accepted definition of the genus Parazoanthus Low et al.

However, the new species Parazoanthus aliceae sp. The reason for this apparent incongruity may be related to the fact that P. Thus P. Indeed, P. However, molecular phylogenetics and especially short patristic distances clearly indicate a very close relationship with P. Such a discrepancy between molecular and morphological data for the genus Parazoanthus further emphasizes the need to identify and standardize the morphological characters used in Zoantharia systematics Swain et al.

The new species presents a large variety of nematocysts as the main character, including several b-mastigophore categories absent from the other species in the same clade large holotrichs in all tissues and at least two different categories of homotrich in all the analyzed tissues. In addition, P.

However, it differs from P. There are also other deep-sea Parazoanthus species for which sequences are not yet available, but which could possibly belong to this Parazoanthus s. Even though cnidae show great promise for taxonomic delimitation, they should be used with caution as some types of cnidae found in P.

There has been continuous effort with the aim of resolving the paraphyly of Parazoanthus with the description of new genera: Antipathozoanthus, Hydrozoanthus, Bergia , and Umimayanthus Sinniger et al. In our phylogenetic reconstruction there is a monophyletic clade also observed by Montenegro et al. This clade is well supported by the molecular data, but the morphology needs further investigation before proper taxonomic reassessment can be conducted for this clade.

However, according to the literature Reimer and Fujii, , P. Moreover, these species occur in geographic distinct areas P. From all the specimens' sequences included into the phylogenetic reconstruction, placement of representatives belonging to Umimayanthus varied the most when comparing phylogenetic trees built from the different alignments.

While all the representatives from these genera formed a well-supported clade according to the SIC alignment 0. As the formation of the genus Umimayanthus was mostly based on the presence of highly conservative and unique indels in the V5 region Montenegro et al. Namely, the SIC alignment also included indel events coded as binary characters, the NoSIC alignment did not utilize phylogenetic information carried by the indel events in such a way, while the Gblock alignment had such potentially poorly aligned positions excluded altogether.

The zoantharian Bergia puertoricense West, formerly known as Parazoanthus clade C sensu Sinniger et al. The genus Hydrozoanthus was described on the basis of substrate specificity and characteristic patterns in DNA sequences specific insertions and deletions in the 16S rDNA, especially in the V5 region as defined in Sinniger et al.

While limited support for such positioning also comes from the Gblocks alignment 0. However, reconstruction of higher level phylogeny within Zoantharia is beyond the scope of this study and should also include representatives from other families accompanied with a detailed revision of their morphology and anatomy.

Furthermore, to resolve the relationships within Parazoanthidae and to identify true paraphylies future efforts should be focused on establishing the correct species tree using a multilocus approach with a multispecies coalescent model and not just relying on mtDNA and limited number of nuclear loci. The zoantharian Epizoanthus martinsae sp. Substrate specificity corroborates the structure of the tree associating a unique type of host or typology of substrate to each of the monophyletic clade of zoantharians: antipatharians to Antipathozoanthus , sponges to Umimayanthus , hydrozoans to Hydrozoanthus , octocorals to the large clade housing Zibrowius, Hurlizoanthus, Corallizoanthus, Kulamanamana, Bullagummizoanthus, Kauluzoanthus , and Savalia Sinniger et al.

In contrast, species within the genus Epizoanthus do not appear to form monophyletic groups according to the type of substrate used, being often associated with gastropod shells regularly inhabited by pagurid crustaceans , tube worms, stalks of glass sponges or living on rocks or dead organisms e. The nature of the symbiotic association between zoantharians and their hosts varies along the mutualism-parasitism continuum.

While the association between zoantharians and hydrozoans is often thought to be commensal or even potentially mutualistic e. In many cases, coral colonization by zoantharians may result in the complete death of the host. This is the case of the zoantharians Antipathozoanthus cf. The parasitic behavior of the zoantharian Savalia savaglia Bertoloni, toward their host octocorals Paramuricea clavata, Eunicella spp. In this case, when the host is completely engulfed by the zoantharian, S.

The symbiotic association between zoantharians and octocorals observed in the present study and reported in Carreiro-Silva et al. Support for a parasitic behavior is based on the following observations: 1 zoantharians covering gorgonian polyps and coenenchyme damaging the host tissue and forcing the gorgonian polyps to live only on the axis; and 2 incorporation of gorgonian sclerites in the zoantharian tissue Figure 3 , see also Figure 3 in Carreiro-Silva et al.

Therefore, these observations suggest that zoantharians progressively eliminate gorgonian tissue, keeping the gorgonian axis as support, while coral sclerites are used for protection, although the complete overgrowth and death of the octocoral host was never observed. Nevertheless, observations made in aquaria confirm that these zoantharians are able to survive in the absence of their coral host, feeding on particles in the water column. In contrast, zoantharians associated with the antipatharian Leiopathes sp.

A similar parasitic relationship between a zoantharian and a deep-sea gorgonian has been described for Epizoanthus sp. Those authors observed what they classified as Epizoanthus sp. Mortensen et al. We found the greatest incidence of zoantharians to occur on C. However, based on our data, we cannot determine whether a similar relationship exists between gorgonian damage caused by fishing and the degree of colonization by zoantharians.

All octocoral specimens examined in this study were collected within fishing grounds, so we have no comparison with areas closed to fishing. Future studies comparing number of colonized C. Our study contributes to a better understanding of the zoantharian diversity associated with CWCs in deep-sea environments see also Carreiro-Silva et al.

Our phylogenetic reconstruction confirmed the relationship between zoantharian taxonomy and the type of substrate selected for settlement Sinniger et al. The phylogenic tree identified four well-supported monophyletic groups within the family Parazoanthidae, one representing all octocorals associated with zoantharians, one representing zoantharians associated with living colonies of antipatharians genus Antipathozoanthus , two representing zoantharians associated with sponges genus Umimayanthus , and genera Parazoanthus and Bergia and one with hydrozoans Hydrozoanthidae, Hydrozoanthus.

Furthermore, our phylogenetic reconstructions confirmed the conclusion of several other authors that Isozoanthus zoantharians associated with non-living substrates forms a well-defined lineage that should not be included within the Parazoanthidae Sinniger et al. Understanding the speciation processes and distribution of these zoantharians will contribute to a better knowledge of the past oceanographic patterns in the worldwide oceans as well as contributing to developing efficient impact assessment methods in relation to deep-sea resource exploitation.

In addition, given the parasitic behavior displayed by octocoral-associated zoantharians, a better knowledge on the effect of fishing and climate change on the severity of parasitic association is required to predict changes in octocoral populations in the future.

Because of difficult access, the deep-sea remains one of the least explored regions of the oceans Mora et al. The study was exempt from ethics committee because it used invertebrate organisms that are not endangered species. Molecular analyses: SS, DS. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. This work is a contribution to the ATLAS project and has received funding from the European Union's Horizon research and innovation programme under grant agreement no.

The paper reflects the authors' views and the European Union is not responsible for any use that may be made of the information it contains. Special thanks go to Lea-Anne Henry for her precious help proofreading English language and grammar, and Bert Hoeksema, Kei Matsuyama, and Max Wisshak for helping with nomenclature questions. Supplementary Figure 1. Values at branches represent Bayesian posterior probabilities. Supplementary Figure 2. Bayesian phylogenetic tree obtained using strict alignment of concatenated sequences COI, 16S, and 12S edited by Gblock Gblock alignment.

Supplementary Table 1. List of species with associated genbank accession number for each of the markers used to build the phylogenetic trees. Abel, E. Eine neue Krustenanemone der Adria, Epizoanthus paxi , nov. Ates, R. Ayres, D. BEAGLE: an application programming interface and high-performance computing library for statistical phylogenetics. Beaulieu, S. Life on glass houses: sponge stalk communities in the deep sea. Bell, F. Contributions of our knowledge of Antipatharian corals.

Bertoloni, A. Google Scholar. Bo, M. Black coral assemblages from Machalilla National Park Ecuador. Braga-Henriques, A. Diversity, distribution and spatial structure of the cold-water coral fauna of the Azores NE Atlantic. Biogeosciences 10, — Buhl-Mortensen, L.

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