N° Spé : UNOC 2025

Arts et sciences

www.openscience.fr/Arts-et-sciences

La revue Arts et sciences présente les travaux, réalisations, réflexions, techniques et prospectives qui concernent

toute activité créatrice en rapport avec les arts et les sciences. La peinture, la poésie, la musique, la littérature, la

fiction, le cinéma, la photo, la vidéo, le graphisme, l’archéologie, l’architecture, le design, la muséologie etc. sont

invités à prendre part à la revue ainsi que tous les champs d’investigation au carrefour de plusieurs disciplines telles

que la chimie des pigments, les mathématiques, l’informatique ou la musique pour ne citer que ces exemples.

Rédactrice en chef

Marie-Christine MAUREL

Sorbonne Université, MNHN, Paris

marie-christine.maurel@sorbonne-universite.fr

Membres du comité

Jean AUDOUZE

Institut d’Astrophysique de Paris

audouze@iap.fr

Georges CHAPOUTHIER

Sorbonne Université

georgeschapouthier@gmail.com

Ernesto DI MAURO

Università Sapienza, Italie

dimauroernesto8@gmail.com

Jean-Charles HAMEAU

Cité de la Céramique Sèvres et

Limoges jean-charles.hameau

@sevresciteceramique.fr

Ivan MAGRIN-CHAGNOLLEAU

Chapman University, États-Unis

magrinchagnolleau@chapman.edu

Joëlle PIJAUDIER-CABOT

Musées de Strasbourg

joelle.pijaudier@wanadoo.fr

Bruno SALGUES

APIEMO et SIANA

bruno.salgues@gmail.com

Ruth SCHEPS

The Weizmann Insitute

of Science, Israël

rscheps@hotmail.com

Hugues VINET

IRCAM, Paris

hugues.vinet@ircam.fr

Philippe WALTER

Laboratoire d’archéologie

moléculaire et structurale

Sorbonne Université Paris

philippe.walter@upmc.fr

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Arts et Sciences, une longue alliance source de

réciprocité créatrice

Marie-Christine Maurel

Depuis l'Antiquité, Arts et Sciences sont congénialement liés. Aristote, déclarait l’art comme

esthétique dont : “Les formes les plus hautes du beau sont l'ordre, la symétrie, le défini, et c'est

là surtout ce que font apparaître les sciences mathématiques. » (Métaphysique, 1078b).

A la Renaissance, le lien épistémique de tout art se renforce encore, avec des figures telles

que Piero della Francesca, grand scientifique, mathématicien éminent, et artiste exceptionnel.

Pensons aussi au Perugino, à son penchant naturaliste et à son élève le génial Raphael. Les arts

visuels ont ainsi particulièrement servi de "passerelles" entre les différentes formes artistiques

et les disciplines scientifiques. Léonard de Vinci polymathe incarne parfaitement cet

universalisme en tant que peintre, sculpteur, mathématicien et poète. Bernard Palissy,

céramiste, sculpteur et savant, a tenté de fusionner l’art et les sciences de la nature à travers ses

représentations de « natures mortes », still life. Diderot a plus tard affirmé dans l’Encyclopédie,

combien l'histoire de la nature est incomplète sans celle des arts. Les exemples sont

nombreux… Malgré une certaine distanciation entre arts et sciences au XXe siècle, en partie

due à la ferveur industrielle, cette séparation s'estompe progressivement aujourd’hui, en

particulier avec l'émergence des arts numériques mais aussi par des lectures qui re-pensent la

modernité de textes dits « classiques ».

Il est essentiel de comprendre comment un scientifique peut aider un artiste, mais aussi

comment un artiste peut apporter sa contribution à un scientifique. Les méthodes, l'imagination

et l'invention sont au cœur des processus scientifiques et artistiques. Cette convergence est

évidente dans la création et dans la scénographie théâtrale, rappelant par exemple le visuel des

mises en scène extraordinaires de Jérôme Bosch, véritable retour à un paradis perdu ou à un

enfer selon les perspectives.

Au-delà des sujets abordés, la créativité commune entre l'art et la science est le fil

conducteur. Les écrits de Diderot Le Rêve de d’Alembert et La Lettre sur les Aveugles à l’usage

de ceux qui voient, les œuvres de Molière traitant de considérations politico-religieuses (voir le

Tartuffe ou l’imposteur) annonciateur de l’imposture créationniste, sont autant d'exemples de

l’actualité et de la convergence entre arts et sciences.

Enfin rappelons que pour Einstein la véritable source de tout art et science réside dans le

mystère et dans l’engagement commun envers l'inconnu : « La plus belle chose dont nous

puissions faire l’expérience est le mystère – la source de tout vrai art, de toute vraie science ».

Aristote Métaphysique, traduction (éd. de 1953) de J. Tricot (1893-1963) Éditions Les Échos du Maquis (ePub, PDF), v.:

1,0, janvier 2014

Diderot Denis. 1769. Le rêve de D’Alembert. GF – Philosophie. Poche, 2002.

Diderot Denis. 1749. Lettre sur les aveugles à l’usage de ceux qui voient. Folio-Poche.

Molière. Le Tartuffe ou l’Imposteur. 1669. Librio-Poche.

Einstein, Albert. Textes écrits entre 1930-1935. Comment je vois le monde. Flammarion-Champs Sciences

© 2024 ISTE OpenScience – Published by ISTE Ltd. London, UK – openscience.fr

Raphael ou l’innovation artistique et scientifique dans : L’Ecole d’Athènes (1509-1511).

Fresque 550x770 cm(18x25ft) Salles Raphael, Musée-Cité du Vatican.

Les personnages représentés ont été identifiés comme suit : Au centre, Platon tenant le

Timée pointe le ciel, illustrant sa théorie des formes idéales et immuables qui existent au-delà

du monde physique. La connaissance, la transcendance va de la réalité à la vérité. A ses

côtés, Aristote, qui tient l’Ethique à Nicomaque, étend sa main vers le sol, symbolisant

l’immanence, la réalité concrète et les phénomènes naturels. La vérité ne peut résider qu’icibas,

dans la réalité.

Le visage de Platon est représenté par Raphael sous les traits de Léonard de Vinci.

A gauche au 1er plan et au bas de la fresque Pythagore et le groupe des géomètres.

Hypathie (philosophe, mathématicienne et astronome d’Alexandrie en Égypte

du IVe au Ve siècle), vêtue de blanc au centre est à proximité de Pythagore.

À l’opposé du côté d’Aristote, la géométrie représentée par la figure d'Euclide et son compas

est entouré d'étudiants. On reconnaît également l'architecte Bramante.

Au-dessus d’Euclide, les astronomes Ptolémée, et Zoroastre soutiennent chacun une

sphère céleste en hommage à leurs contributions en astronomie.


Arts et sciences

2025 - Volume 9

Numéro spécial UNOC

‣ Art & Science, Tradition & Modernity on the French Riviera: A Special Issue on the Occasion

of the United Nations Ocean Conference (UNOC) in Nice .................................................................1

Christian Sardet

DOI : 10.21494/ISTE.OP.2025.1286

‣ June 2025: Nice, capital of the world ocean ......................................................................................3

Jean-Pierre Gattuso, François Houllier

DOI : 10.21494/ISTE.OP.2025.1287

‣ Sailing to create, art in motion ...........................................................................................................9

Myriam Thomas

DOI : 10.21494/ISTE.OP.2025.1288

‣ Ernst Haeckel’s Radiolarians and Medusa: The influence of his visits to Villefranche

on his science and his art ...................................................................................................................17

John R. Dolan

DOI : 10.21494/ISTE.OP.2019.0420

‣ On the Works of Albanis Beaumont (1747-1810), a key contribution to the establishment

of the region of Nice as a favorite destination ...................................................................................31

John R. Dolan

DOI : 10.21494/ISTE.OP.2025.1289

‣ Two centuries of arts and science in Nice and Villefranche sur Mer:

1) Pioneers: 1800 to 1900 ..................................................................................................................46

Christian Sardet

DOI : 10.21494/ISTE.OP.2025.1290

‣ Two centuries of arts and science in Nice and Villefranche sur Mer:

2) Modern era: 1960 to 2024 .............................................................................................................68

Christian Sardet

DOI : 10.21494/ISTE.OP.2025.1291

‣ The Université Internationale de la Mer, Arts and Sciences ..............................................................90

Jean-Eric Aubert

DOI : 10.21494/ISTE.OP.2025.1292


Arts et sciences

2025, vol. 9, n° spécial UNOC 2025, 1-2 pages, DOI : 10.21494/ISTE.OP.2025.1286 ISTE OpenScience

Art & Science, Tradition & Modernity on the French

Riviera: A Special Issue on the Occasion of the United

Nations Ocean Conference (UNOC) in Nice

Art & Science, Tradition & Modernité sur la Côte d’Azur : Numéro spécial

à l’occasion de la Conférence des Nations Unies sur l’Océan (UNOC)

Christian Sardet 1

1

Sorbonne University, CNRS, Laboratoire de Biologie du Développement (LBDV), Institut de la Mer de Villefranche-sur-

Mer (IMEV), 06230, France, christian.sardet@imev-mer.fr

ABSTRACT. This special issue highlights the convergence of art and science on the French Riviera in celebration of the

3rd United Nations Ocean Conference (UNOC) and the One Ocean Science Congress (OOSC) hosted in Nice in June

2025. It explores the region’s rich legacy of marine research and naturalist traditions alongside contemporary cultural

initiatives such as the Biennale des Arts et de l’Océan. The issue pays tribute to historical figures, pioneering scientists,

and modern researchers whose work continues to shape our understanding of marine life and ocean sustainability.

RÉSUMÉ. Ce numéro spécial met en lumière la rencontre entre l’art et la science sur la Côte d’Azur, à l’occasion de la 3e

Conférence des Nations Unies sur l’Océan (UNOC) et du Congrès One Ocean Science (OOSC), organisés à Nice en juin

2025. Il retrace la riche tradition naturaliste et scientifique de la région tout en valorisant les initiatives culturelles

contemporaines comme la Biennale des Arts et de l’Océan. Ce dossier rend hommage aux figures historiques, aux

pionniers de la biologie marine et aux chercheurs d’aujourd’hui qui contribuent à une meilleure compréhension et

préservation des océans.

KEYWORDS. Nice, ocean, science, art, marine, UNOC, OOSC.

MOTS-CLÉS. Nice, ocean, science, art, marine, UNOC, OOSC.

In June 2025, the city of Nice will proudly host two major international events: the 3rd United

Nations Ocean Conference (UNOC) and the One Ocean Science Congress (OOSC). These gatherings

will unite thousands of researchers, environmental advocates, and policymakers dedicated to

addressing the degradation of ocean health and the alarming loss of marine biodiversity. Jean-Pierre

Gattuso and François Houllier highlight the global significance of these events, particularly in the

context of climate change, biodiversity conservation, and urgent societal challenges. Their shared

vision culminates in the Nice Ocean Action Plan, an ambitious framework designed to promote a

healthy, resilient ocean for future generations (1).

To celebrate this pivotal moment, the city of Nice has chosen an oceanic theme for its 2025 cultural

program (2). The Biennale des Arts et de l’Océan, titled La Mer Autour de Nous—inspired by Rachel

Carson’s influential book The Sea Around Us — will feature 11 exhibitions across 9 institutions, along

with 6 site-specific art installations throughout the city. One highlight is Becoming Ocean, a

contemporary art exhibition at Villa Arson, presented within the broader context of artist residencies,

as described by Myriam Thomas of the Tara Oceans Foundation (3).

Nestled between Alpine peaks and the azure Mediterranean, the French Riviera has long been

admired for its dramatic natural beauty. At the turn of the 19th century, writer and illustrator Albanis

Beaumont helped define this picturesque image through detailed travel and art books. His evocative

engravings and writings drew the attention of the British aristocracy, contributing to the Riviera's

© 2025 ISTE OpenScience – Published by ISTE Ltd. London, UK – openscience.fr Page | 1

emergence as a sought-after winter destination. John Dolan revisits Beaumont’s life and legacy in this

issue (5).

Nice also has a deep-rooted naturalist tradition, closely linked to the region’s rich biodiversity. In

the early 1800s, pharmacist-naturalists such as Antoine Risso and Jean Baptiste Vérany conducted

extensive studies of local marine and terrestrial species, aided by fishermen and local farmers. Their

pioneering efforts included some of the first scientific and artistic representations of marine organisms.

In 1810, zoologist François Péron and his friend, the renowned artist Charles-Alexandre Lesueur

visited Nice to document pelagic species, a topic explored in Christian Sardet’s article (6).

By the mid-19th century, supported by Vérany, distinguished German and Swiss biologists such as

Johannes Müller, Carl Vogt, and the young Ernst Haeckel came to Nice and Villefranche-sur-Mer to

study pelagic marine life. Their research flourished amid the revolutionary scientific atmosphere of

Darwin’s theory of evolution and the newly emerging cell theory of Schleiden and Schwann.

Meanwhile, Jean-Baptiste Vérany and Jean-Baptiste Barla — assisted by the talented local painter

Vincent Fossat—produced exquisite painted collections of fish and fungi, still preserved today as

exsiccata and molds at the Natural History Museum of Nice (6).

This legacy of scientific and artistic exploration continues today, especially at the Villefranche-sur-

Mer marine station, founded in the 1880s by pioneering biologists Hermann Fol — the discoverer of

fertilization — and Jules Barrois (6). In the late 19th and early 20th century, Russian scientists took

control of the station. They were originally attracted by the presence of the Russian Tsar’s family and

aristocracy wintering in Nice and the Russian marine’s frequent presence in the plankton-rich bay of

Villefranche.

Today, the Institut de la Mer de Villefranche (IMEV), under the joint direction of Sorbonne

University and the French CNRS, upholds this heritage. World-renowned researchers and students

work collaboratively to deepen our understanding of marine organisms, planktonic ecosystems, and

oceanic processes, drawing continuous inspiration from the scientific pioneers who came before them

(7). Finally, Jean-Eric Aubert traces the history of the Université Internationale de la Mer ( IUM) a

local institution created by his father raised awarness about marine and maritime isssues among

generations of local and foreign students.

References

(1) GATTUSO J.P., HOULLIER F., “June 2025: Nice, capital of the world ocean”, Arts et sciences, vol 9, n° spécial UNOC,

p. 3-8, 2025

(2) La Biennale des Arts et de l’Océan : https://anneedelamer.nice.fr/biennale-des-arts-et-de-locean/

(3) THOMAS M., “Sailing to create, art in motion”, Arts et sciences, vol 9, n° spécial UNOC, p. 9-16, 2025

(4) DOLAN J., “Ernst Haeckel’s Radiolarians and Medusa: The influence of his visits to Villefranche on his science and

his art”, Arts et sciences, vol 9, n° spécial UNOC, p. 17-30, 2025

(5) DOLAN J., “On the Works of Albanis Beaumont (1747-1810), a key contribution to the establishment of the region of

Nice as a favorite destination”, Arts et sciences, vol 9, n° spécial UNOC, p. 31-45, 2025

(6) SARDET C., “Two centuries of arts and science in Nice and Villefranche sur Mer: 1) Pioneers: 1800 to 1900”, Arts et

sciences, vol 9, n° spécial UNOC, p. 46-67, 2025

(7) SARDET C., “Two centuries of arts and science in Nice and Villefranche sur Mer: 2) Modern era: 1960 to 2024”, Arts

et sciences, vol 9, n° spécial UNOC, p. 68-89, 2025

(8) AUBERT J. E., “The Université Internationale de la Mer, Arts and Sciences”, Arts et sciences, vol 9, n° spécial

UNOC, p. 90-96, 2025


Arts et sciences


June 2025: Nice, capital of the world ocean

Juin 2025 : Nice, capitale de l’océan

Jean-Pierre Gattuso 1,2 and François Houllier 3

1

Sorbonne Université, CNRS, Laboratoire d'Océanographie de Villefranche, 181 chemin du Lazaret, F-06230 Villefranche-sur-Mer,

France

2

Institute for Sustainable Development and International Relations, Sciences Po, 27 rue Saint Guillaume, F-75007 Paris,

France

3

IFREMER, 1625 route de Sainte-Anne, F-29280 Plouzané, France

ABSTRACT. We outline the critical importance of the ocean for planetary health, economic prosperity, and human wellbeing,

while highlighting the urgent threats it faces from climate change, pollution, overfishing, and poor governance. It

presents the One Ocean Science Congress (OOSC) and the 3rd United Nations Ocean Conference (UNOC3), scheduled

for June 2025 in Nice, France, as pivotal events aimed at promoting ocean sustainability through science-based policy,

innovation, and global cooperation. The OOSC will deliver scientific recommendations to Heads of States and Governments,

while additional high-level events—the Ocean Rise and Coastal Resilience Summit and the Blue Economy and

Finance Forum—will address climate adaptation and sustainable investment. The outcome of these efforts will be the

Nice Ocean Action Plan, comprising a political declaration, voluntary commitments, and strategic priorities focused on

multilateral processes, financial mobilization, and enhanced scientific knowledge to advance the Sustainable Development

Goal 14 and ensure a resilient, thriving ocean for future generations.

RÉSUMÉ. Nous soulignons l’importance cruciale de l’océan pour la santé de la planète, la prospérité économique et le

bien-être humain, tout en mettant en lumière les menaces urgentes auxquelles il est confronté : changement climatique,

pollution, surpêche et gouvernance insuffisante. Il présente le One Ocean Science Congress (OOSC) et la 3e Conférence

des Nations Unies sur l’Océan (UNOC3), qui se tiendront en juin 2025 à Nice, comme des événements clés pour

promouvoir la durabilité de l’océan à travers des politiques fondées sur la science, l’innovation et la coopération internationale.

Le congrès OOS formulera des recommandations scientifiques à l’attention des chefs d’États et de gouvernements,

tandis que deux autres forums de haut niveau — le Sommet sur l’élévation du niveau de la mer et la résilience côtière,

et le Forum sur l’économie bleue et la finance — aborderont respectivement l’adaptation au des zones littorales et

l’investissement durable. L’ensemble de ces efforts aboutira au Plan d’action de Nice pour l’océan, structuré autour d’une

déclaration politique, d’engagements volontaires et de priorités stratégiques centrées sur le renforcement des processus

multilatéraux, la mobilisation des financements et l’accroissement des connaissances scientifiques afin de faire progresser

l’Objectif de Développement Durable 14 et garantir un océan résilient et prospère pour les générations futures.

KEYWORDS. Ocean, sustainability, climate change, blue economy, Governance, United Nations, Science, Policy.

MOTS-CLÉS. Océan, Durabilité, Changement climatique, Économie bleue, Gouvernance marine, Nations Unies,

Science, Politique.

Introduction

The ocean is crucial for our survival and underpins sustainable development. It covers more than

70% of the Earth’s surface and is home to diverse ecosystems, from lagoons and coral reefs to deep-sea

habitats, providing food security for over three billion people and supporting industries like fisheries,

aquaculture, tourism, and maritime transportation. The ocean also enriches our cultural heritage and

offers opportunities for recreation and scientific discovery. It absorbs 25% of anthropogenic CO₂

emissions and stores 90% of the excess heat due to human activities and faces increasing threats from

overfishing, pollution, habitat destruction, and climate change, particularly through ocean acidification

and warming

Annually, the ocean's ecosystem services – benefits to people – contribute approximately $2.5

trillion to the global economy. Coastal ecosystems such as seagrass and mangroves sequester carbon,

and shield coastlines from strengthening and increasingly frequent storm surges, reducing disaster’s


risks. The ocean’s potential for renewable energy through offshore wind, tidal, and wave power aids in

decarbonization efforts. Its rich biodiversity fosters advancements in marine biotechnology, enhancing

medicine, food security, and the development of sustainable materials. The ocean plays a vital role in

promoting mental health, recreation, and spiritual well-being.

However, the ocean is facing significant threats from human activities. Climate-warmed waters

cause coral bleaching and alter marine ecosystems and currents. Ocean acidification and

deoxygenation pose risks to coral reefs and shellfish, destabilizing food webs. Overfishing, pollution

(e.g. plastic, excess nutrients, and chemicals), oxygen loss, and habitat destruction (e.g. coastal

development, deep-sea mining) are pushing marine ecosystems toward collapse. Weak governance and

unequal resource access hinder effective conservation and sustainable management. This changing

ocean creates knowledge gaps which expand faster than science can address them.

Despite these challenges, the ocean holds immense potential for solutions to global crises. The

scientific community enhances the understanding of our ocean’s resources and ecosystems and the

intrinsic value to preserve and use them sustainably. Embracing the blue economy to support the

application of these solutions improves livelihoods, economic growth, and ocean health. Prioritizing

ocean-based solutions increases resilience, reduces greenhouse gas emissions, unlocks economic

opportunities, and safeguards vital ecosystems for future generations.

Sustainable Development Goal 14 and United Nations Ocean Conferences

The Sustainable Development Goals 1 (SDGs) are a set of 17 global goals adopted by the United

Nations in 2015 as part of the 2030 Agenda for Sustainable Development. They provide a roadmap for

addressing the world’s most pressing challenges, including poverty, inequality, climate change,

environmental degradation, peace, and justice. The SDGs emphasize a balanced approach to economic

growth, social inclusion, and environmental protection, aiming to create a more sustainable and

equitable future for all.

Sustainable Development Goal 14 2 (SDG 14: Life Below Water) aims to conserve and sustainably

use the oceans, seas, and marine resources for sustainable development. Key targets under SDG 14

include reducing marine pollution, protecting marine and coastal ecosystems, ending illegal,

unreported, and unregulated (IUU) fishing, and enhancing scientific knowledge and technology

transfer for ocean sustainability. Global cooperation is essential, as the ocean is globally connected,

and the health of marine ecosystems directly affects livelihoods, especially in small island developing

states (SIDS) and coastal communities that rely on marine resources.

Achieving SDG 14 requires integrated policies, effective marine governance, and sustainable blue

economy strategies that balance conservation with economic development. Efforts such as Marine

Protected Areas (MPAs), ocean-based climate solutions, and strengthened international agreements are

crucial to ensuring a resilient and thriving ocean for future generations.

United Nations Ocean Conferences (UNOCs) are high-level global gatherings focused on

accelerating action for SDG 14. The Third Conference will be co-hosted by France and Costa Rica and

held in Nice, France, from 9 – 13 June 2025. It will build on the previous UN Ocean Conferences,

hosted by Sweden and Fiji in 2017 in New York and by Portugal and Kenya in 2022 in Lisbon.

UNOC3 will be preceded by three United Nations Special events: the One Ocean Science Congress,

the Ocean Rise and Coastal Resilience Coalition Summit and the Blue Economy and Finance Forum.

Numerous other activities will take place between 28 May and 16 June, making the city of Nice the

capital of the ocean (Figure 1).

1 https://sdgs.un.org/goals

2 https://sdgs.un.org/goals/goal14


Figure 1. Chronological view of the events that will be held in Nice (France) and Monaco in June 2025.

One Ocean Science Congress

The One Ocean Science Congress (OOSC) is a high-level international gathering of leading ocean

scientists, policymakers, and stakeholders dedicated to advancing ocean knowledge and its role in

addressing global challenges. As the scientific prelude to the 3rd United Nations Ocean Conference

(UNOC), the OOSC will provide a crucial platform for showcasing the latest scientific advancements,

interdisciplinary research, and innovative solutions for ocean sustainability.

Scheduled for 2025, the congress will focus on key themes such as climate change, biodiversity loss,

ocean acidification, deoxygenation, marine pollution, and sustainable ocean governance. It will

emphasize the role of science in shaping evidence-based policies, fostering international collaboration,

and supporting ocean-based solutions for climate mitigation and adaptation.

The OOSC aims to bridge the gap between scientific research and decision-making, ensuring that

robust, policy-relevant knowledge informs global ocean governance. By convening world-renowned

experts, early-career scientists, and representatives from governments, intergovernmental

organizations, and civil society, the congress will facilitate discussions on the United Nations Decade

of Ocean Science for Sustainable Development (2021–2030) and its contributions to the Sustainable

Development Goals (SDGs).


Ultimately, the One Ocean Science Congress will serve as a catalyst for strengthening the sciencepolicy

interface, fostering international cooperation, and driving impactful ocean action.

The International Scientific Committee of the One Ocean Science Congress (OOSC) proposes

recommendations for the Heads of State and Government at the Third United Nations Ocean

Conference (Box 1). These recommendations are based on published scientific literature, major UN

reports, and initial outcomes of the United Nations Decade of Ocean Sciences for Sustainable

Development (2021-2030).

Grounded in scientific evidence, the International Scientific Committee of the One Ocean Science

Congress advances a set of 10 thematic recommendations to shape discussions at the 2025 United

Nations Ocean Conference. These recommendations are summarized below; the extensive version is

available 3 , with specific actions detailed in the following sections:

1. Inspire responsibility and respect for the ocean, integrating across knowledge systems.

2. Enable effective, equitable and environmentally safe ocean-based approaches to achieve the

mitigation and adaptation goals of the Paris Agreement.

3. Effectively protect and restore marine and coastal ecosystems through equitable and sustainable

management.

4. Pause harmful activities in the deep ocean while improving knowledge to enable sustainable and

equitable uses.

5. Ensure equitable sharing of benefits derived from marine genetic resources.

6. End illegal, unreported and unregulated fishing and improve transparency.

7. Ensure sustainable, equitable, and safe ocean-based food systems.

8. Adopt comprehensive measures to end marine plastic pollution.

9. Decarbonize shipping and reduce the environmental impact of maritime transport.

10. Ensure ambitious investments in inclusive fundamental transdisciplinary knowledge generation

to inform ocean action.

Five cross-cutting recommendations are outlined to underpin these efforts:

- Reinforce support to multilateral organizations to support fairness, inclusiveness, ecological

safety, and precautionary principles.

- Implement the already existing regulations and international commitments to protect the ocean

and its vital services to society.

- Eliminate subsidies that harm climate and biodiversity.

- Strengthen oceanographic research, particularly by enhancing observation and modeling

capacity and facilitating knowledge sharing.

- Sustain and build on the momentum that will be generated by the Third United Nations Ocean

Conference to strengthen the link between science and policy.

These recommendations serve as a framework for action, to ensure that science remains at the heart

of efforts to secure a sustainable future for the ocean and humanity.

Box 1

3 Gattuso J.-P., Houllier, F., Adams J., Amon D., Bambridge T., Cheung W., Chiba, S., Cortes, J., Duarte C., Frölicher T. L., Gelcich

S., Gephart J., Gjerde, K., Haugan P., Li D., Takoko M., Tuda A., 2025. Recommendations to Heads of State and Government from

the International Scientific Committee of the One Ocean Science Congress, Nice, 3-6 June 2025.

https://doi.org/10.5281/zenodo.14361191.


Ocean Rise and Coastal Resilience Summit

The Ocean Rise and Coastal Resilience Summit 4 is a pivotal international forum addressing the

urgent challenges posed by sea level rise, coastal erosion, extreme weather events, and climate-driven

ocean changes. Bringing together leading scientists, policymakers, urban planners, engineers, and

community leaders, the summit fosters interdisciplinary collaboration to develop innovative, sciencebased

solutions for enhancing coastal resilience.

As global temperatures rise, accelerating ice melt and thermal expansion are driving unprecedented

sea level rise, threatening coastal populations, infrastructure, and ecosystems. The summit will

highlight the latest research on ocean dynamics, climate modeling, nature-based solutions, and

adaptive governance, providing a platform to translate scientific insights into actionable strategies.

Key themes include climate adaptation, resilient urban planning, managed retreat, blue carbon

ecosystems, and financing mechanisms for coastal resilience. Special attention will be given to

vulnerable communities, particularly Small Island Developing States (SIDS) and low-lying coastal

cities, where the impacts of ocean rise are most acute.

The Ocean Rise and Coastal Resilience Summit aims to catalyze global action by fostering policy

dialogues, sharing best practices, and strengthening international cooperation. By bridging science,

policy, and local action, it will contribute to the development of sustainable, climate-resilient coastal

systems worldwide.

Blue Economy and Finance Forum

The Blue Economy and Finance Forum 5 is a high-level platform dedicated to advancing sustainable

ocean-based economies through innovative financial mechanisms, investment strategies, and policy

frameworks. It brings together leaders from finance, government, industry, science, and civil society to

explore how responsible investment can drive the transition to a regenerative and climate-resilient blue

economy.

The ocean contributes trillions of dollars annually to the global economy, supporting industries such

as fisheries, aquaculture, shipping, tourism, offshore renewable energy, and marine biotechnology.

However, overexploitation, pollution, and climate change threaten the long-term sustainability of these

sectors. The forum focuses on scaling up blue finance, including blue bonds, sustainable ocean funds,

impact investment, and public-private partnerships, to ensure economic growth aligns with marine

conservation and climate adaptation goals.

Key discussions will address de-risking ocean investments, financial innovations for marine

protected areas, regulatory frameworks, and the role of financial institutions in driving sustainable

ocean business models. Special emphasis will be placed on mobilizing capital for SIDS and coastal

communities, ensuring inclusive and equitable economic benefits.

By fostering collaboration between financial actors and ocean stakeholders, the Blue Economy and

Finance Forum aims to unlock sustainable investments that support a thriving ocean and resilient

coastal economies.

4 https://sdgs.un.org/events/unoc-2025-special-event-ocean-rise-and-coastal-resilience-coalition-summit-56788

5 https://beff2025monaco.org/


United Nations Ocean Conference

The 3rd United Nations Ocean Conference 6 (UNOC3) is a global platform dedicated to advancing

ocean sustainability and strengthening international cooperation to achieve SDG 14: Life Below Water.

Co-hosted by UN member states, this high-level event brings together heads of state, policymakers,

scientists, businesses, NGOs, and civil society to address the most pressing challenges facing the

ocean, including climate change, pollution, overfishing, habitat destruction, and declining marine

biodiversity.

UNOC3 will build on the outcomes of previous editions, aiming to accelerate transformative actions

for a sustainable blue economy, marine conservation, climate resilience, and equitable ocean

governance. Discussions will focus on science-based solutions, innovative technologies, policy

reforms, and financial mechanisms to protect marine ecosystems while supporting livelihoods and

economic growth.

Key themes include scaling up ocean-based climate mitigation and adaptation, strengthening marine

protected areas, combating illegal fishing, reducing plastic and chemical pollution, and ensuring

sustainable ocean financing. The conference will also highlight the importance of indigenous

knowledge, local communities, and youth engagement in ocean stewardship.

By fostering global commitments and partnerships, the UN Ocean Conference aims to drive impactful policy

changes and investments that ensure a healthy, resilient, and productive ocean for future generations.

Conclusion

The Nice Ocean Action Plan is the anticipated outcome of UNOC3. It aims to serve as a pivotal

framework for global efforts in ocean conservation, fostering collaboration and concrete actions to

ensure the health and sustainability of marine ecosystems for future generations.

The key components of the Nice Ocean Action Plan are:

- The Political Declaration: A concise, action-oriented declaration reflecting the collective

commitment of UN Member States to enhance ocean conservation efforts.

- Voluntary Commitments: A compilation of pledges from governments, international

organizations, civil society, the private sector, and other stakeholders, detailing specific actions

and resources dedicated to achieving SDG 14.

Thee key strategic priorities are:

- Multilateral ocean processes: Advancing international agreements and initiatives related to

ocean protection to bolster global conservation efforts.

- Mobilizing Financial Resources: Securing investments to support SDG 14 and fostering the

development of a sustainable blue economy.

- Enhancing Marine Scientific Knowledge: Strengthening Ocean Science, and the dissemination

of marine scientific data and knowledge to inform and improve policy-making.

6 https://unocnice2025.org/en/


Arts et sciences


Sailing to create, art in motion

Naviguer pour créer, l’art en mouvement

Myriam Thomas 1

1

Head of Ocean Literacy department, Tara Oceans Foundation, 8 Rue de Prague, 75012, Paris

myriam@fondationtaraocean.org / https://fondationtaraocean.org/en/home/

ABSTRACT. Science has always been a source of inspiration for artists. In this spirit, the Tara Ocean Foundation works

closely and organize exhibitions with artists to cultivate fresh perspectives on its scientific expeditions and on the Ocean

itself. Through residencies aboard the Tara schooner and Tara Polar Station, artists can harness their unique vision and

creativity to capture and reinterpret the richness of the Ocean, the essence of scientific exploration, and the rhythms of

life at sea. The goal is to awaken public awareness of the Ocean’s beauty and essential role in our world.

RÉSUMÉ. Science et connaissance ont toujours été une source d’inspiration pour les artistes. C’est dans cet esprit que

la Fondation Tara Océan collabore étroitement et organise des expositions avec des artistes, pour cultiver un autre

regard sur ses expéditions scientifiques et l’Océan. En résidences artistiques à bord de la goélette et la station polaire, à

travers leurs regards uniques et leurs créativités, ils capturent et réinterprètent la richesse de l’Océan, la recherche

scientifique et la vie quotidienne à huis clos. L’objectif est de sensibiliser le public à la beauté et l'importance de l'Océan.

KEYWORDS. Artists, residencies, science, public awareness, oceans, pollutions, Tara Foundation.

MOTS-CLÉS. Artistes, residences, science, diffusion du savoir, oceans/ pollutions, Fondation Tara.

Conversation métabolite by Antoine Bertin.

Photo ©Fondation Tara_104_bfougeirol 2025


One Ocean, Many Stories..

Knowledge has always been transmitted through stories. From the discovery of new territories and

the establishment of the first commercial maritime routes in the 16th century, to the great scientific

expeditions like Darwin’s, tales of exploration and seafaring adventures have shaped and spread our

understanding of the world. Our personal relationship with the Ocean reflects the incredible diversity

of cultures across the globe. These perspectives — whether marked by anxiety, serenity, or awe —are

rooted in the intimate, often mysterious connection each of us shares with this vast and largely

unexplored realm.

The Tara Ocean Foundation (1) is not only a hub for scientific research through its expeditions (2),

but also a platform for artistic exploration (3). Thanks to the enduring commitment of agnès b. and

Etienne Bourgois, more than 50 artist residencies have taken place aboard the schooner since 2004 (2).

Men and women from diverse backgrounds have set sail to document and interpret the richness of the

ocean through their unique creative lenses — sharing insights into the scientific work undertaken on

board.

Among the first to embark on this journey were renowned artists such as Pierre Huyghe, Xavier

Veilhan, and Sebastião Salgado.

Creativity Without Borders

Artistic residencies aboard the schooner offer a journey of discovery. Each artist must adapt to the

unpredictable rhythms of the sea, using its ever-changing environment as inspiration to build, reflect,

and complete their work. Painters, illustrators, photographers, sculptors, writers, sound artists, and

filmmakers all contribute to revealing what often remains hidden — offering fresh ways to perceive the

Ocean. Rather than categorizing this artistic abundance by subject matter, the editorial vision of the

residencies is shaped around five major themes: Living, Landscapes, Pollutions, Sensible, and

Travelogues.

Under Living, Christian Sardet & The Macronauts’s Plankton chronicles installation : Ballet du

Plancton (4), Manon Lanjouère’s Particules (5), and Aurore de La Morinerie’s Cells & Salps (6)

reveal the microscopic life that forms the basis of our ecosystem.

Landscapes, as seen through the eyes of Nicolas Floc’h (7), Yann Bagot (8), and Emmanuel Régent

(9), offer windows onto the marine world — each using photography, intricate line drawing, or ink to

portray different oceanic vistas.

Through the lens of Pollutions, works by Samuel Bollendorff (10), Laure Winants (11), and

Robertina Šebjanič (12) make visible the often-invisible threats facing the ocean, confronting us with

the environmental challenges of our time.

In Sensible, Elsa Guillaume’s Slices (13) navigates the boundary between attraction and repulsion,

while Noémie Sauve (14) highlights coral’s delicate fragility — set against the subtle soundscapes of

Antoine Bertin’s plankton voices (15).

Finally, Travelogues — through video diaries, illustrations, paintings, and photographs — trace the

transformation of each artist through their experience at sea. These works remind us that they all

originate from a shared stage: the Ocean itself.


Left: Heliogravures - Aurore de la Morinerie

Right: Un monde sculptural - Cécile Fouillade – SIQOU.

Photo ©Fondation Tara_104_bfougeirol2025

Plankon Ballet – Christian Sardet & The Macronauts.


Les larmes de sirènes – Samuel Bollendorf offers a dialogue between the

beauty of seascapes and the sad reality of samples taken on site.



Art in Common, at the Heart of Social Ties with the Ocean

A landmark exhibition « La Grande Expédition » at Le Centquatre (104) in Paris (16) and the

publication of L’Art et la Science pour Révéler l’Océan (17) by The Eyes Publishing offered the public

a chance to experience singular creations born of these residencies.

The goal was to reconnect us with the ocean, reveal its hidden biodiversity, underscore its central

role in regulating the climate, and confront us with the pollution that threatens it — all through the eyes

of artists.

The works also posed deeper, universal questions:

• What is humanity’s relationship with life?

• How do we leave our mark on the landscape?

• What invisible forces endanger living beings?

• Can we perceive the ocean through the senses?

Today, sharing these questions is essential. By doing so, we open a collective dialogue about the

future of the Ocean — and of life itself.

To reach broader audiences, the Tara Ocean Foundation is bringing these artistic visions to venues

across many regions, making the ocean’s story accessible to all.

Part of the representation of the Ocean and the expeditions of the schooner Tara as seen by the artists

collective Ensaders. Photo ©Fondation Tara_104_bfougeirol202

A new chapter opened in Nice with the exhibition « Becoming Ocean » (18), at the Villa Arson, coproduced

by Villa Arson (19), TBA21–Thyssen-Bornemisza Art Contemporary (20), and the Tara

Ocean Foundation.


The remarkable exhibition echoes the 3rd United Nations Ocean Conference (UNOC) (21), which

takes place in Nice in June 2025. It brings together the voices of diverse artists to reflect on the

Ocean’s present and future.

The exhibition is one of 11 events taking place in the city of Nice’s 6th Biennale des Arts (22) –

« La Mer Autour de Nous », a title in hommage to Rachel Carson’s famous book « The sea around

us » (23). All events are centered on the Ocean and two monumental works by artists associated with

the Tara Ocean Foundation have taken center stage in the city of Nice on this occasion (23).

Nicolas Floc’h, artist, navigator, and diver, presents a light installation on the façade of the Bellanda

Tower, tracing the swirling patterns of the Gulf Stream, which connects the Gulf of Mexico to Europe.

Emmanuel Régent’s CETACEA unfurls along the Nice harbor breakwater, using a line of blue light

to mark the presence of cetaceans and dolphins near the Pelagos Sanctuary —reminding us of the

invisible inhabitants that traverse these waters.

Both works resonate with the pressing issues discussed at the UN summit UNOC and the One

Ocean Science Congress (21, 24), illuminating the necessity of ocean protection.

Becoming Ocean exhibition at Villa Arson in Nice


Nicolas Floc’h neon light installation « Gulf Stream, 2017-2025 »,

Bellanda tower in Nice. Photo Nicolas Floc’h

Emmanuel Régent « CETACEA » light line installation (with Miraceti),

Nice Harbor. Photo Emmanuel Régent

New Residencies on the Horizon

In 2026, new territories of artistic exploration will emerge. Once again, painters, illustrators,

photographers, sculptors, writers, and audiovisual artists will be invited to propose projects aligned

with the Foundation’s scientific missions.


March 2026 will see the continuation of the Tara Pacific mission on coral ecosystems, while the

Foundation’s new vessel, Tara Polar Station (25), will embark on a stationary expedition into the

Arctic Ocean — delving deep into the workings of the climate system. As always, artists will

accompany the voyage, offering their interpretations of these fragile and vital environments.

Onboard the vessels of the Tara Ocean Foundation — the Tara schooner and the Tara Polar

Station (26) — each residency becomes an act of creation and reflection. The ships are both laboratory

and studio an ever-evolving environment where art and science converge.

Scientists and artists alike seek to understand and translate the ocean’s mysteries. Though their

methods differ, they share a common goal: to awaken a deeper collective awareness of the Ocean’s

essential role in sustaining life on Earth.

The vessels of the Tara Ocean Foundation.

Photo @ Fondation Tara Océans Valentin Lauféron

Acknowledgements

We would like to thank Christian Sardet for initiating and translating this article and for his

extraordinary plankton images, which have allowed the public to discover the ocean’s invisible world.

We would also like to thank all the artists who took part in the artistic residencies aboard the Tara

schooner.

We would like to express our sincere gratitude to Ms. Hélène Guénin and Mr. Jean-Jacques

Aillagon for their dedicated curatorship of the Nice Biennale, and to Ms. Marie-Ann Yemsi and Ms.

Francesca Thyssen-Bornemisza for their visionary work on the Villa Arson exhibition.


References

1) Tara Oceans Foundation : https://fondationtaraocean.org/en/home/

2) Tara expeditions : https://fondationtaraocean.org/en/expedition/

3) Artists in residence - Tara Oceans Foundation https://fondationtaraocean.org/en/art-science/

4) Christian Sardet & The Macronauts : https://planktonchronicles.org/en/

5) Manon Lanjouère : https://manonlanjouere.com/Les-Particules

6) Aurore de la Morinerie : https://www.auroredelamorinerie.com/

7) Nicola Floc’h : https://www.nicolasfloch.net/

8) Yann Bagot : https://yannbagot.com/

9) Emmanuel Régent : https://fondationtaraocean.org/artistes/emmanuel-regent/

10) Samuel Bollendorff : http://www.samuel-bollendorff.com/fr/

11) Laure Winants : https://laurewinants.com/

12) Roberta Sebjanic : https://robertina.net/

13) Elsa Guillaume : https://elsaguillaume.com/

14) Noémie Sauve : https://noemiesauve.com/

15) Antoine Bertin : https://www.studioantoinebertin.com/

16) Exhibit « La Grand Expédition » Centquatre Paris : Nov.6th 2024 – March 2 nd 2025 https://www.104.fr/ficheevenement/la-grande-expedition.html

17) Exhibition catalogue/ book: "Les artistes révèlent l'Océan" The Eyes publishing

https://theeyes.eu/livres/tara-les-artistes-revelent-locean/

18) Exhibit « Becoming Ocean » at Villa Arson, Nice : May 8th – August 24th 2025 https://villaarson.fr/programmation/expositions/becoming-ocean-a-social-conversation-about-the-ocean/

19) Villa Arson, Art School and Art Center, Nice : https://villa-arson.fr/en/

20) TBA21–Thyssen-Bornemisza Art Contemporary : https://tba21.org/

21) UNOC / Third United Nation Ocean Conference, Nice June 9-13th, 2025 https://unocnice2025.org/en/

22) 6th Biennale des Arts et de l’Océan, Nice, May to October 2025 : https://anneedelamer.nice.fr/biennale-des-artset-de-locean/

23) Rachel Carson « The Sea Around Us / La Mer Autour de Nous » Wildprojects Edition (Marseille) :

https://wildproject.org/livres/la-mer-autour-de-nous

24) One Ocean Science Congress, Nice June 3-6th 2025 : https://one-ocean-science-2025.org/

25) Tara Polar Station : https://fondationtaraocean.org/en/schooner/tara-polar-station/

26) Tara Schooner : https://fondationtaraocean.org/en/schooner/tara-schooner/


Arts et sciences

2025, vol. 9, n° spécial UNOC 2025, 17-30 pages, DOI : 10.21494/ISTE.OP.2019.0420 (First published in 2019, vol. 3, n° 2) ISTE OpenScience

Ernst Haeckel's Radiolarians and Medusa:

The influence of his visits to Villefranche on his science

and his art

Les radiolaires et les méduses d'Ernst Haeckel : influence de ses visites à

Villefranche-sur-mer sur sa science et son art

John R. Dolan 1

1

Laboratoire d'Océanographie de Villefranche-sur-Mer

ABSTRACT. Early in his long career, Ernst Haeckel (1834 - 1919) twice visited Villefranche-sur-Mer. First, as a student,

in 1856 during a sampling trip to Nice, and again in 1864 when sent to Nice by his parents for a change of scenery

following the untimely death of his first wife. The two visits appear to have been key events in the development of

Haeckel's science and art as they are the beginnings of his studies, first on radiolarians, and then on medusa. During the

1856 visit he observed for the first time living radiolarians, the group of microscopic planktonic protists, the subject of his

first monographic work in 1862 that brought him fame at a young age. During the 1864 visit he resided in Villefranche-sur-

Mer. There, for the first time, he made detailed observations on the development and morphology of medusa. He

subsequently produced monumental monographs on both radiolaria and medusa, e.g., the Challenger Reports, which

remain today his major scientific contributions. Haeckel's artistic fame is largely from his Kunstformen der Natur. The

book relies heavily on illustrations of both radiolarians and medusa, more so than other groups of organisms, and

contains iconic images of medusa and radiolarians, suggesting a major importance in Haeckel's art for the two groups

linked closely with Haeckel's visits to Villefranche-sur-Mer.

RÉSUMÉ. Au début de sa longue carrière, Ernst Haeckel (1834 - 1919) s’est rendu deux fois à Villefranche-sur-mer.

D'abord en tant qu'étudiant, en 1856, lors d'un voyage d'échantillonnage à Nice, puis de nouveau en 1864, lorsque ses

parents l'envoyèrent à Nice pour un dépaysement total à la suite du décès prématuré de sa première femme. Les deux

visites semblent avoir été des événements clés dans le développement de la science et de l'art de Haeckel au début de

ses études, d'abord sur les radiolaires, puis sur les méduses. Lors de sa visite en 1856, il observa pour la première fois

des radiolaires vivants, groupe de protistes planctoniques microscopiques, sujet de son premier travail monographique

en 1862 qui le rendit célèbre à un jeune âge. Pendant la visite de 1864, il résida à Villefranche-sur-mer. Il y fit pour la

première fois des observations détaillées sur le développement et la morphologie des méduses. Il a par la suite produit

des monographies monumentales sur les radiolaires et les méduses, par exemple les rapports Challenger, qui demeurent

aujourd’hui ses principales contributions scientifiques. La renommée artistique de Haeckel provient en grande partie de

son livre « Kunstformen der Natur ». Le livre s'appuie beaucoup sur des illustrations de radiolaires et de méduses, plus

que d'autres groupes d'organismes, et contient des images emblématiques de méduses et de radiolaires, suggérant une

influence majeure dans l'art de Haeckel pour ces deux groupes, étroitement liée aux visites de Haeckel à Villefranchesur-mer.

KEYWORDS. history of science, plankton, scientific and artistic voyages, microscopy.

MOTS-CLÉS. histoire des sciences, plancton, voyages scientifiques et artistiques, microscopie.

Backstory/Introduction

This article grew out of a recent serendipitous discovery. I was searching for an obscure species

description by Ernst Haeckel, an important personality in the history of biology. He was a

contemporary and staunch advocate of Charles Darwin and is one of the best known and most read

zoologists (Egerton 2013). I came across another obscure species description by him, one that

surprised me. It was a description of a meduse (a jellyfish), and he called it "Carmarina", the Niçois


word for sea meat, a term used by the fisherman of the Nice region for gelatinous plankton caught in

their nets.

Haeckel's use of the word suggested a certain familiarity with both the medusa and the fishermen of

the region. Given the date, his striking illustration appeared to be his first published drawing of any

medusa. His illustrations of medusa are famous, even iconic. In fact, Haeckel is well known for his

detailed studies and magnificent illustrations, primarily of two groups of organisms: radiolarians,

microscopic creatures with fabulously intricate skeletons or shells, and the medusa (fig.1).

Fig 1. Examples of Haeckel's illustrations of radiolarians (left panel), plate in Haeckel 1862 and medusa

(right panel), plate 3 in Haeckel 1879. In actual size, the radiolarians shown are about 1/10 mm in diameter

and the medusa shown are about 3 cm long.

I was familiar with the fact that he first saw living radiolarians when he visited the Bay of

Villefranche in 1856 as a young medical student (fig. 2), the veritable start of his studies radiolarians.

However, the medusa description suggested his other major obsession, medusa, may have also begun

in the Bay of Villefranche with his 1864 stay. The stories presented here, of two relatively short visits,

aim to convince you dear reader, that his visits to southern France were key events in the development

of Haeckel's science and his art. I hope to convince you that his major scientific and artistic legacies,

both of which are considerable, are intimately linked to Villefranche.


Fig. 2. Ernst Haeckel and his microscopes, ca. 1856, when he was a medical student. His parents bought him

his first microscope, a Schiek barrel microscope (on the table at the left), in 1853 and he quickly became very

fond of it, calling it "my darling" and "my divine microscope"; he developed the unusual ability to look through

the eyepiece with one eye and with the other draw what he saw in the microscope (Otis 2007). Photo courtesy

of SLUB Dresden/Deutsch Fotothek.

Haeckel's 1856 Visit

Ernst Haeckel saw the Bay of Villefranche for the first time during a sampling trip to Nice in the

autumn of 1856, at age 22. He had been invited to participate in the expedition by his teacher, Albert

Kölliker. Based in Nice, they spent 4 weeks in the region. Many years later, Haeckel stated: "In

company with Heinrich Müller and K. Kupffer, we investigated especially the rich pelagic animal life

of the beautiful bay of Villafranca. There, for the first time I met those wonderful forms of the pelagic

fauna which belong to the classes of the siphonophores, pteropods, and heteropods. I also there first

saw living polycyttariia, acanthrometra, and polycystina, those phantasmic forms of radiolaria, in the

study of which I spent so many later years". (Haeckel 1893). According to a letter sent to his parents at

the time (Haeckel 1856), during their stay they met repeatedly with the Berlin Professor Johannes

Müller who was himself sampling along the French Mediterranean coast, in Villefranche, Nice, St.

Tropez and Cette (now Sete) for his study of Mediterranean radiolarians published posthumously in

1858 (Müller 1858).

Johannes Müller is credited by Haeckel as the one who initiated him to the study of planktonic

organisms, earlier, in 1854 during a trip to Helgoland. Haeckel (1893) states " When at Helgoland,

investigating the wonders of the plankton with the microscope, Johannes Müller, pleased with the care

and patience with which his zealous students tried to study the charming forms of medusa and

ctenophores, spoke to me the ever-memorable words, "There you can do much; and as soon as you

have entered into this pelagic wonderland you will see that you cannot leave it." According to Haeckel

(1893), in Nice in 1856, Müller stressed to him the particular interest of studying radiolaria as he "....

called my attention to the many and important questions which the natural history of these enigmatical

microscopical organisms present.".

Haeckel mentions, in that same letter to his parents (Haeckel 1856), that towards the end of their

stay, they used "Müller's fine nets" to collect organisms in the Bay of Villefranche noting that among

the catch were the most remarkable Thalassicolla (a group of radiolarians). It was perhaps this outing

from Nice that provided the sample containing the radiolarian Haeckel described in his 1862

monograph, Sphaerozoium italicum (Fig. 3), as from his stay in Nice in 1856. It appears to be the first

new species of radiolarian he collected. In recent years, the species was collected again from the Bay

of Villefranche and molecular data from it used to refine the phylogeny of radiolaria (Bass et al. 2005).


Fig 3. Plate 33 from Haeckel's 1862 monograph Die Radiolarien, showing the radiolarian species first

collected during his 1856 visit, Sphaerozoum italicum (figs. 1 & 2).

The diameter of the radiolarian is about 1/3 mm.

The 1862 'Die Radiolarien' was Haeckel's first monographic publication (excluding his thesis and

habilitation volumes) and he dedicated it to Johannes Müller. The study was largely the result of a long

stay in Messina devoted mostly to collecting and examining radiolarians. Haeckel credited Müller's

suggestion, made in Nice in 1856, as the motivation for his "spending an entire year in pelagic fishing"

(Haeckel 1893). The massive two-volume work earned Haeckel a promotion to "Extraordinary

Professor" and the Cothenius medal of the Leopold-Caroline Academy of German Naturalists in 1863.

Haeckel sent a copy to Darwin and it is said to have astonished him (Richards 2008).

In subsequent years, Haeckel turned his attention to other groups of organisms and topics. He

returned to studies of radiolarians only in the 1880's as he was asked to exploit the sample gathered

during Challenger Expedition. His resulting radiolarian "masterpieces" are the monographs

constituting the "Report on the Radiolaria" (Haeckel 1887a,b), totaling over 1700 pages of text and 140

plates. Although Haeckel's taxonomy has been revised, and many species have been found to be

synonyms, (e.g. Aita et al. 2009; Lazarus 2014), the work is still regularly cited today (e.g. Biard et al.

2017, Grattepanche et al. 2017, Dolan et al. 2019, Kachovich et al. 2019)

It is speculation to propose that Haeckel may not have studied radiolarians if Kölliker had not

invited Haeckel to join his sampling expedition. However, it is not speculation to state that a firm link

exists, not least of all in Haeckel's mind, between his 1856 visit to Villefranche and his subsequent

work on radiolarians.


Haeckel's 1864 Visit

As mentioned above, following the appearance of his monograph on radiolarians, Haeckel was

named "Extraordinary Professor" in Jena. This was in June 1862. The financial stability of the new

position allowed him to marry Anna Sethe (to whom he had been engaged since 1858) in August of

1862. Unfortunately, the marriage was short-lived. Anna Sethe Haeckel died suddenly in early

February of 1864, perhaps of appendicitis. Haeckel was completely devastated by the loss of his wife.

Apparently unable to end his grieving, his parents sent him to Nice for a six-week stay in March-April

1864 for a change of scenery (Richards 2008).

From a long letter to his parents (Haeckel 1864a), we know that Haeckel found Nice disagreeable.

He quickly moved to Villefranche, profiting from contacts made during his previous visit. He took

rooms in Casa Montolivo, describing his host, Abbé Montolivo, as a "marine doctor". Abbé Montolivo,

along with Jean Baptiste Vérany, were the Nice naturalists who had provided aide in sampling during

Haeckel's previous visit (Haeckel 1856). In Villefranche, he sought to distract his grief through work

(Haeckel 1864a). He apparently worked quite hard as his stay in Villefranche was remarkably

productive.

Haeckel's time in Villefranche appears to have marred by bad weather, responsible for many days of

poor sampling conditions in the bay (Haeckel 1864b). This, and the fact that he was residing in

Villefranche, he proudly declared himself to be the first naturalists to reside in Villefranche rather than

visit from Nice (Haeckel 1864a), perhaps explains why his work consisted of largely observing living

specimens rather than cataloguing new forms. Haeckel did not completely ignore radiolarians,

devoting some time to observing feeding behaviour (Haeckel 1865a). He also described new species

(Fig. 4). Notably, his description of one as "Protogenes primordialis" appears to be the beginning of

his now discredited theory of the primitive protist cell as a form without a distinct nucleus (Haeckel

1871). However, he must have spent most of his time observing medusa, especially following their

complicated developmental stages, given the large amount of data he gathered on medusa.

Fig. 4. The plate from Haeckel 1865a (Über den Sarcodekörper der Rhizopoden) illustrating the radiolarians

observed during his 1864 visit. Fig. 1 & 2: Protogenes primordialis 'feeding'; Fig. 3: Acanthodesmia

polybrocha; Fig. 4: Actinelius purpureus; Fig. 5: Cyrtidosphaera echinoides. The actual size of all are about 1/4

mm in longest dimension.


Haeckel's first published illustration of medusa, describing a new species (Haeckel 1864c) came

from his 1864 stay in Villefranche (Fig. 5). Remarkably, Haeckel named the new form in a sort of

tribute to the local fisherman, using their term "carmarina" (Niçois for sea meat) for gelatinous

plankton, as the name of the new genus, Carmarina. It would be featured later in his art book

Kunstformen der Natur. Haeckel also named a new medusa shown in Figure 5, found while walking

along the bay, for his deceased wife: Mitrocoma annae (Anna's headband) in a most poetic manner

(Richards 2008). The small medusa would be the first of 3 species named for Anna Sethe, the second

being the iconic medusa Desmonema annasethe of the Kunstformen der Natur (plate 8 below in fig. 6).

The third, and last species, he named for her was a radiolarian, Dictycodon annasethe also shown in a

plate in Kunstformen der Natur (plate 31 below in Fig. 8).

Fig. 5. Haeckel's first published illustration of a meduse: Plate 11 from Haeckel 1864c (Die Familie der

Rüsselquallen, Medusae Geryonidae) Carmarina hastata. Actual size of the meduse is about 10 cm long.


Fig. 6. One of the new medusa Haeckel found in Villefranche he named for deceased wife Anna Sethe

Haeckel, Mitrocoma annae. It was the first of 3 species he named for her. The illustration is from his 1879

'System der Medusan'. The actual size of the 'bell' portion is about 4 cm

Haeckel's publications based on his observations of medusa while in Villefranche (for less than 4

weeks!) number 4 (1864b,c; 1865b,c) with a 5th as a monograph combining 3 of the 4 papers (1865d).

He continued his studies of medusa later in Jena with descriptions of fossil medusa (1865e, 1866,

1869a, 1874) and extant medusa (1869b). His medusa studies culminated with his Das System der

Medusen (1879, 1880) and the deep sea medusa of the Challenger expedition (1881, 1882). In the Das

System der Medusen, in the 1879 atlas of illustrations, 9 of the 154 species shown are stated to have

been specimens from Villefranche, presumably collected or drawn during his 1864 visit. Through the

years, much of his taxonomy has been revised but other parts have been validated using molecular

methods (e.g. Bayha, et al. 2010).


Haeckel's interest in medusa, unlike the radiolarians, cannot be said to have begun in Villefranche.

His first 'field trip' devoted to plankton was to Helgoland (Germany) in 1854 in the company of

Johnannes Müller who introduced him to "plankton fishing" (Florey 1995). According Haeckel, it was

then that he discovered "the charming forms of medusa and ctenophores" (Haeckel 1893). Nonetheless,

Haeckel's first detailed studies of medusa were conducted in Villefranche and only after his stay in

Villefranche did he begin his ultimately very large number of studies on medusa (Fig. 7).

Fig. 7. Haeckel's output of publications concerning radiolarians and medusa from 1860 to 1890 based the

titles given in Way's Haeckel bibliography (Way 1909). Haeckel's many publications on medusa began

appearing shortly after his second visit to Villefranche. Note the apparent alternations of periods dominated by

publications on one group. Haeckel's total output from 1855 to 1900 numbered 107 titles. Radiolarians and

medusa accounted for 14 and 17 titles respectively, many more than any other individual groups. For example,

the third most represented group was sponges with 4 titles (not shown).

Radiolarians and Medusa in Haeckel's Art

Haeckel's artwork is known almost exclusively from his Kunstformen der Natur, although it can be

found in many of his monographic works such as the Challenger Reports (Williams et al. 2015).

Kunstsfomen der Natur was published in 10 installments of 10 plates each from 1899 to 1904. The 100

plates had a considerable impact on the Art Nouveau movement and continue to have an impact in the

fields of art and design. Furthermore, according to some, he is remembered today, even in the scientific

community, more for his artwork than for his contributions to science (Williams et al. 2015). Given

that Haeckel's work on radiolarians and medusa can be closely linked to his visits to Villefranche, one

might ask how did radiolarians and medusa feature in his artwork? How important were they?

The relative importance of radiolarians and medusa in Haeckel's art quickly becomes apparent in

surveying the 100 plates of the Kunstformen der Natur. Medusa and radiolaria feature very

prominently (Fig. 8), accounting for 22 of the 100 plates. Among the 10 installments, usually both

were represented. No other groups of organisms are so abundantly featured. However, a greater

prominence was given to medusa to which he devoted 12 plates compared to 10 for radiolarians. One

might speculate that Haeckel had a slight preference for medusa. In this regard it perhaps worth

recalling that Haeckel named his house in Jena "Villa Meduse" and named two medusa, and but one

radiolarian, for Anna Sethe Haeckel.


Fig. 8. The plates from Kunstformen der Natur featuring radiolarians or medusa numbered 22 out of the 100

plates. The numbers in the lower right corners denote the plate number. The book was published in

installments, sets of 10 plates. Thus, the first installment contained plates 1 to 10, the second 11 to 20, etc.,; it

can be seen then that most of the 10 installments contained both a medusa and a radiolarian plate. Plate 8

shows Desmonema annasethe, one of the two medusa species Haeckel named for his deceased wife. Plate

26 featured the medusa Carmarina hastata described from Villefranche. Plate 31 includes Dictyocodon

annasethe (bottom row center), the radiolarian he named for he named for deceased wife.


According to Richards (2009), Haeckel's illustrations in Kuntsformen der Nature have been

criticized by some as being more artistic than scientific, for example artificially adding symmetry to

radiolarians. In this regard it is interesting to compare an illustration from the original description to

that in the Kunstsformen plate. Figure 9 shows the illustration of Dictycocodon annasethe from the

original description in the Challenger Report with that in Plate 31 of the Kunstformen. Near perfect

symmetry is apparent in the Kunstformen version compared to the rather approximate symmetry

shown in the Challenger Report (Haeckel 1887b). Aesthetics trumped accuracy in the Kunstformen

illustrations of at least some organisms. Furthermore, Haeckel's own words concerning the use of

color in the Kunstformen (from Richards 2009, a translation of text from the Forward) are telling. It

would appear then the Kunstformen illustrations likely should be considered as primarily artistic works

rather than scientific illustrations.

I have been convinced that colored images (even of a mediocre production) are much more

valuable for a vivid intuitive awareness of nature than the photograph or the simple blackand-white

illustration. Indeed, a crude color sketch (if it conveys the landscape in a vivid

fashion) has a deeper and more stimulating effect than the best black-and-white illustration

or photographic representation. This distinction lies not only in the effect of color itselfsince

different individuals are sensitive in different measures- but also because the painter,

as thoughtful artist, reproduces in his subjective image the conceptually articulated

character of the landscape and emphasizes its essential features. The objective image of the

photograph, by contrast, reproduces equally all parts of the view, the interesting and the

mundane, the essential and the inessential. Thus the colored photograph, if it should be

brought to perfection, will indeed never be able to replace the individually conceived and

deeply felt image of the painter.

Fig. 9. Dictyocodon annasethe from Kunstformen der Natur in plate 31 (left) and from the plate 71 in the

Challenger Report (right). Note the near perfect symmetry in the Kunstformen version compared to the original

version. The actual size is about 1/5 mm long


It is perhaps noteworthy that the two volume tribute to Ernst Haeckel, published a few years before

his death (Schmidt 1914). and filled with laudatory texts from scientific notables was illustrated not

with any of his scientific illustrations but mainly with Haeckel's landscapes. Furthermore, the last

image of Haeckel in the book of remembrances is of him as an artist (Fig. 10).

Fig. 10. Ernst Haeckel in 1914 from the Heinrich Schmidt's 1914 tribute to Haeckel "Was wir Ernst Haeckel

Verdanken". Photo by Alfred Bischoff taken in 1914.

Regardless of any scientific view, for the general public, Haeckel's illustrations are artworks. For the

general public, what in Haeckel's art appears to be most popular? One manner of assessing popularity

is the price of the individual plates of Kunstformen der Natur plates. Dealers in old books and prints

offer for sale the individual prints. Presumably the plates were printed in about equal numbers so that

price differences among the plates should reflect differences in demand more than supply. One dealer

offering individual plates for many years now is Stefan Wulf of Berlin. His 2019 catalogue (Wulf

2019) includes all the plates.

In Wulf's catalogue, the prices of the Kunstformen plates vary considerably, from $25 to $250, with

the highest price asked for the plate 8 of the medusa Desmonema annasethe. One naturally assumes

that the multi-colored plates fetch a higher price than the mono-chromatic plates, regardless of subject,

and indeed the average prices are $91 and $49, respectively. Still, among the multi-colored plates, the

medusa plates average $91 and the radiolarian plates $61. Thus, it appears that, based on prices, the

medusa plates are more popular than the radiolarian plates. Are the medusa and radiolarian plates

overall more appreciated, based on price? The average price of the medusa and radiolarian plates is

$71 compared to $63 for the other subjects. It appears then that is a higher demand for plates with

medusa or radiolarians than for other subjects.


Conclusion: Villfranche & Haeckel

As a final note concerning the importance of Haeckel's visit to Villefranche, it does appear the

subjects linked to his stays in Villefranche, the radiolarians and medusa, were to the most popular with

Haeckel himself, among the many zoological topics he studied, given the number of publications he

ultimately devoted to them (Fig. 5). His medusa and radiolarian artwork, compared to his other

subjects, appear to be the most popular with today's general public based on the asking prices for

Haeckel's artworks. Thus, his major scientific and artistic legacies, both of which are considerable, are

intimately linked to Villefranche. Despite the apparent links, Haeckel did not return to Villefranche

until late in his life, just a few years before his death in 1919. He visited his former student Michael

Davidoff at the Russian Zoological Station in Villefranche while attending the 1910 opening of the

Oceanographic Museum (Davidoff 1914). One can only speculate that perhaps Villefranche was also

intimately linked to his grieving for Anna Sethe.

Acknowledgements

Encouragement was kindly provided by Robert J. Richards and Marie-Christine Maurel. The

comments of David Montagnes on earlier drafts resulted in considerable improvement. However, I

retain full responsibility for all errors of fact and interpretation.

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Arts et sciences


On the Works of Albanis Beaumont (1747-1810), a key

contribution to the establishment of the region of Nice

as a favorite destination

Sur les œuvres d'Albanis Beaumont (1747-1810), contribution majeure à

l'établissement de la région niçoise comme destination privilégiée

John R. Dolan 1

1

Sorbonne Université, CNRS, Laboratoire d'Océanographie de Villefranche-sur-Mer, Station Zoologique, 06230

Villefranche-sur-Mer, France, john.dolan@imev-mer.fr

ABSTRACT. Through his illustrated books featuring the region, Albanis Beaumont was as an early contributor

establishment of the region of Nice, now known as the Côte d'Azur, as a destination for the wealthy and the curious of

Great Britain. His were large-format works, presenting attractive and remarkably detailed scenes of the countryside and in

particular views of the coast. Beaumont's texts covered aspects of both ancient and natural history, topics of interest to

the aristocracy of the period. Although well known in his time, Beaumont is today a forgotten figure and his works known

to few, save collectors of rare books. Here first is presented a brief account of the unusual life of Albanis Beaumont and

the equally unusual life of his artistic collaborator, Cornelius Apostool, the engraver who created many prints from

Beaumont's drawings. The biographies are followed by an 'exhibition' of his works featuring views of the region including

notes on their reception at the time. These are the 1787 "Voyage Historique et Pittoresque du Comté de Nice", the 1794

"Select Views of the South of France with Topographical and Historical Descriptions the 1795 "Travels through the

Maritime Alps from Italy to Lyon across the Col de Tende, by the way of Nice, Provence, Languedoc, etc.", as well as the

unauthorized translation of the 1787 work, published anonymously as "An Historical and Picturesque Description of the

County of Nice" (1792).

RÉSUMÉ. Grâce à ses livres illustrés sur la région, Albanis Beaumont a été l'un des premiers à faire de la région de Nice,

aujourd'hui connue sous le nom de Côte d'Azur, une destination prisée des riches et curieux de Grande-Bretagne. Ses

œuvres de grand format présentaient des scènes de campagne attrayantes et remarquablement détaillées, notamment

des vues du littoral. Ses textes couvraient des aspects de l'histoire ancienne et naturelle, sujets d'intérêt pour l'aristocratie

de l'époque. Bien que célèbre à son époque, Beaumont est aujourd'hui un personnage oublié et ses œuvres sont peu

connues, hormis des collectionneurs de livres rares. L'ouvrage présente d'abord un bref récit de la vie atypique d'Albanis

Beaumont et de celle, tout aussi atypique, de son collaborateur artistique, Cornelius Apostool, graveur à l'origine de

nombreuses estampes réalisées à partir de ses dessins. Les biographies sont suivies d'une exposition de ses œuvres

présentant des vues de la région, accompagnées de notes sur leur réception à l'époque. Il s'agit du « Voyage historique

et pittoresque du comté de Nice » de 1787, des « Select Views of the South of France with Topographical and Historical

Descriptions» de 1794, du « Travels through the Maritime Alps from Italy to Lyon across the Col de Tende, by the way of

Nice, Provence, Languedoc, etc. » de 1795, ainsi que de la traduction non autorisée de l'ouvrage de 1787, publiée

anonymement sous le titre « "An Historical and Picturesque Description of the County of Nice» (1792).

KEYWORDS. Côte d'Azur, Maritime Alps, Travel Albums, Tourism, Geotourism, Cornelius Apostool.

MOTS-CLÉS. Côte d'Azur, Alpes Maritimes, Récits de Voyage, Tourisme, Géotourisme, Cornelius Apostool.

Introduction

The fame of the region of Nice as a favored destination is generally traced back to the 18th century

books of the Scotsman Tobias Smollett and the German Swiss Johann Georg Sulzer. Smollett was a

literary figure of considerable renown, and Sulzar was a major scientific and philosophical personality.

Their books were quite popular. Smollett's 1776 "Travels through France and Italy" went through

several editions, and Sulzer's 1780 "Tagebuch einer von Berlin nach den mittäglichen Ländern von

Europa in den Jahren 1775 und 1776 gethanen Reise und Rückreise" (Diary of a journey from Berlin to


the southern countries of Europe in the years 1775 and 1776 and return journey) was translated into

French (Sulzer 1781; 1789) and Italian (Amoretti & Salzer 1865). Both Smollett and Sulzer had

traveled to Nice for reasons of health. Their books included long sections describing their stays in Nice

and extolled both the winter climate and the scenery. However, the books did not contain any

illustrations of the scenery described, for example by Sulzer (1780) as of "incomparable beauty" or

"prettier than be imagined". It was not until several years later that Albanis Beaumont published the

first album of scenes of Nice (Mézin 2005) in his 1787 "Voyage Historique et Pittoresque du Compte

de Nice", and the first illustrated travel guide to the region (Boyer 2002) in his 1795 "Travels through

the Maritime Alps from Italy to Lyon across the Col de Tende, by the way of Nice, Provence,

Languedoc, etc.". The works of Beaumont have then long been recognized as playing a significant rôle

in attracting visitors, especially the English, to the region of Nice (e.g. Bosio 1925).

Beaumont's books should likely be classified as art books rather than travel books. They are large

format works (typically 50 x 30 cm) in which the prints were a principal, if not the main component,

and were directed towards an audience of means, those who might undertake overseas voyages.

Nonetheless, his works were reviewed, for the most part positively, in the popular literary press of the

day and thus were likely widely known, not only among the wealthy travelers but also those interested

in natural history. This was because Beaumont's books contained many descriptions of geological

formations, deposits, caves and fossils. His were among the earliest geological observation of the

region of Nice (Barale 2016). Beaumont's depictions and descriptions of geological formations as

points of interest, future tourist attractions, made him one of the founders of 'geotourism' (Cayla et al.

2015).

Today Beaumont's books are quite expensive, held in very few libraries, and best known by dealers

in rare books. Surprisingly and sadly, most of Beaumont's book are not available online. Consequently,

the artwork in Beaumont's books that attracted people to the region of Nice is unknown to most of us.

The goal of this essay is two-fold. The first goal to shine a light on the interesting lives of the

exceptional artists who created the illustrations in Beaumont's books. This includes both Albanis

Beaumont , the author of the texts and primary artist, and Cornelius Apostool, the producer of the

many aquatint plates from Beaumont's drawings. Thus, first will be given short biographical sketches

of the unusual careers of Beaumont and Apostool. The second goal is to present examples of their

striking illustrations, those that depicted the region of Nice for the first time and that no doubt attracted

travelers on a "grand tour", and those seeking to escape the winters of northern Europe.

1. Biographies of the artists

1.1. The Curious Life of Albanis Beaumont

Beaumont made no mention of his life in his books, save for two footnotes in volumes of his last

major work, "Descriptions des Alpes Grecques et Cottiennes" (Beamont 1802a,b; 1806a,b). In one note

he states that he was born in Chambéry (at the time under the house of Savoy, now in the French

department Savoi), and the king Victor Amédée of Sardinia sent him to Nice to be trained in

hydrological works under the chief engineer of the county of Nice. The king subsequently sent him to

study in Turin to obtain a Sardinian diploma as his previous studies had been in Chambèry and Paris

(Beaumont, 1802b). In the other note he stated that while in Nice, the Duke of Gloucester (younger

brother of the British King George 3rd) asked that Beaumont be permitted to be employed as a tutor

for his son, William, and that during a long stay in England he published works on the Alps based in

part on his travels with the Duke and his family (Beaumont 1802a). Fortunately, more is known about

Beaumont.

There are several more or less complete accounts of his life (Nagler 1835, Chapperon 1842, Rabut

1872, Mettrier 1911, Brondel 2005a,b, 2007). While the accounts differ in some details, they largely

agree on the sequence of his career as first an engineer, and then in turn, a tutor to the children of a

prince, a landscape artist, a travel writer, a sheep farmer, and finally a mine developer. Except where


otherwise noted, the summary of Beaumont's life given below relies largely on the articles by Brondel

that are the most detailed and documented accounts.

Beaumont was born on the 19th of May 1753, the only child of Bernard and Claudine Beaumont in

Bissy, near Chambèrey. His father was the cook and groundskeeper of a lawyer. Despite his own

statements, there are apparently no traces of any of Beaumont's studies in Paris or Chambèry. With

some certainty Beaumont can be placed in Nice in 1781, sent by the king of Sardinia to work under

Jean-François Michaud (1734-1809), chief engineer of the county of Nice, then working on the

construction of the port of Nice. In 1783 the king had Beaumont pursue a diploma in architecture in

Turin, granted with project of a church. On his return to Nice, still in the service of the Sardinian king,

he met William Duke of Gloucester who was passing the winter in Nice. The Duke had been banished

from the court of his older brother King George 3 for having married outside the nobility, and

undertook lengthy travels about Europe with his wife, children, and a considerable entourage. The

Duke asked Beaumont to seek the consent of the king of Sardinia to become a tutor to his seven year

old son, and the request was granted. No details are available with regard to the conditions of

Beaumont's employ. He remained with the Duke's family throughout their subsequent travels, up to

and including, the return to England in 1787. Shortly before his departure for England, Beaumont

published, in Geneva, his first album of landscapes, "Voyage Historique et Pittoresque du Comté de

Nice" (Beaumùont 1787a) dedicated to the Duke of Gloucester, who apparently suggested the project

to Beaumont. It was followed in short order by an album of alpine scenes, "Voyages Pittorseque aux

Alpes Pennines, etc." (Beaumont 1787b), dedicated to the Duke's eldest daughter Sophia.

Once in England, Beaumont became a productive author of travel albums in English. He published a

series of books on the south of France and the Alps, all but the last with prints of surprising detail

engraved by Corneluis Apostool from drawings Beaumont has done during his travels. The first was

"Travels through the Rhaetian Alps the Year 1886 from Italy to Germany, Through Tyrol" (Beaumont

1792) with the mention of 'ten large aqua-tinta engravings' prominently mentioned on the title page.

Beaumont dedicated the work to his young charge, Prince William Frederick. The book made

Beaumont's mark in Britain as the title pages of all the subsequent books in English would include the

notation "by the author of Travels through the Rhaetian Alps". The next title was "Select views of the

South of France with Topographical and Historical Descriptions" (Beaumont 1794), again with the

aqua-tint plates by Apostool, the 15 plates included views of Antibes, Toulon and Marseille. "Travels

through the Maritime Alps from Italy, to Lyons across the Col de Tende, by the way of Nice, Provence,

Languedoc, etc." appeared in 1795. It was dedicated to the Duchess of Gloucester, and contained 15

aqua-tint plates by Apostool, including views of Views of Nice, Villefranche, Monaco, and

Vingtimille. Beaumont's last book in English was "Travels through the Lepontine Alps from Lyon to

Turin by the way of the Pays-de-Vaud, the Vallais, the monts Great St. Bérnard, Simplon, and St.

Gothard" (Beaumont 1800). It contained 27 plates, noted only with his initials A.B. in the lower left

corners. The engraver was not indicated. However, it was unlikely to be Apostools's work as he had

returned to Amsterdam in 1796.

For reasons that are unclear, Beaumont left England to return to Savoy to begin a new career to raise

sheep in his native Savoy in late 1796. It was perhaps because his pension from the King of Sardinia

was uncertain following the death of Victor Amédée or simply a desire to return to a rural landscape.

Beaumont had married while in England, in 1870, to Louise Poignand. The Beaumont's voyage and in

particular, her impressions of post-revolutionary France, were recorded in a series of letters published

in 1798 as "A Sketch of Modern France in a series of Letters to a Lady of Fashion Written in the Years

1796 and 1797 during a Tour through France" (Moody 1798). The author of the letters was

anonymous until fairly recently when it was shown to be by Louise Beaumont (Wellington 2003). It

was a remarkably successful book, extensively reviewed or excerpted in the British press (Anon.

1798a,b,c,d,e; Anon. 1799), and translated into French (Babeau 1888). It appears likely that Louise

Beaumont's book sold far better than any of husband's works!


Once in Savoy, Beaumont began his project of raising the Spanish race of Merino sheep for their

wool on a farm near Geneva. By 1803 he had a herd of 500 sheep but encountered difficulty in selling

wool and lambs. By 1809 he gave up on raising sheep, and acted on plans to build an iron mine and

foundry in the region. It would be Beaumont's last, and only completely unsuccessful project. He

encountered unexpected difficulties such as water in the deposits, the quality of the ore extracted, and

disputes with the partners who financed the project. Beaumont died on November 27, 1811, bankrupt,

leaving his wife a pauper. Beaumont's passing was briefly noted in the British press as having been

"celebrated for his splendid Travels in the Rhaetian, Maritime and Lepontine Alps" (Anon. 1812a,b).

1.2. The Remarkable Career of Cornelius Apostool

The account given here of Apostool's career are based on information in the articles by Murray

(1937), Jonker (1977), and Hinterding & Horbatsch (2016). Cornelius Apostool was born on August

6th 1767, the 11th of twelve children into a family of modest means in Amsterdam. As a youngster he

was apprenticed to a gold and silver merchant. At age 17, he began attending classes in the

Stadstekenacademie ("City Drawing Academy") as a pupil of the landscape painter and draughtsman

Hendrik Meyer. In 1786, Apostool traveled to London with Meyer. It is unclear exactly when and how

he came to be employed as an engraver, creating aquatints from drawings of landscapes. His first

documented work was creating plates for Samuel Ireland's "Picturesque Tour through Holland,

Braband and part of France" published in 1790. Apostool is assumed to have done the plates for

Ireland's subsequent pictorial works in the genre of 'Picturesque Travels' such as his 1794 "Picturesque

Views on the River Thames".

Apostool's first works with Beaumont were the 10 plates for "Travels through the Rhaetian Alps,

etc". published in 1792 followed by "Select Views of the South of France, etc." with the 15 plates by

Apostool which appeared in 1794. Apostool's most important work with regard to exposing the scenery

of the region of Nice to a wide audience was his 18 plates in Beaumont's 1795 "Travels through the

Maritime Alps". Apostool's overall production during his time in London was estimated to be about 80

plates, including his own album of 15 aquatint prints, reproductions of the landscapes of Dutch

Masters in his "The Beauties of The Dutch School; selected from Interesting Pictures of Admired

Landscape Painters." (Apostool 1792-1793). Thus, while not a majority, a substantial part of his work,

while in London, was for the albums of Beaumont.

Apostool returned to Amsterdam in 1796 and largely abandoned working as an artist. He obtained a

series of professional appointments or commercial jobs in which he honed skills of making friends and

conducting negotiations. In 1807 he was appointed secretary to the Dutch ambassador in Naples.

Shortly after, at age 46, he was named director of the Royal Museum in Amsterdam. With fall of

Napoleon Bonaparte, Apostool gained fame with his role in recovering artwork taken from Holland by

Napoleon. This included the 10,243 prints taken from the Royal Library in the Hague, considered a

national heritage. The recovered prints were transferred to the Royal Museum in Amsterdam under

Apostool's direction. Little is known of his personal life. He never married and had no children. He

died in Trippenhuis on 10 February 1844 at the age of 77.

2. The Exhibition of Beaumont's Scenes

In following section, Examples of Beaumont's illustrations of the Region of Nice as well Provence

and the Maritime Alps are presented, grouped by publication. It is important to note the size of an

image as it appeared in the work (given in the legends) is likely considerably larger that the size on the

page being viewed. The legend texts given are adapted from the publication.


2.1. Voyage Historique et Pittoresque du Comté de Nice

Figure 1. The cover of Beaumont's first work "Voyage Historique et Pittoresque du Comté de Nice" and the

first plate, his map of the region of Nice.

This first work of Beaumont in 1787 was an album consisting of a map, 12 landscapes, and only 6

pages of text. In the preface, dedicating the work to the Duke of Gloucester, Beaumont states "It is

relying entirely on your indulgence, that I dare take the liberty to place before you this first attempt

with my weak talents". It is said the Duke of Gloucester financed the production of the album (Potron

2005). One can assume the work was quite successful as an unauthorized translation, without an author

named, was published in London in 1792 (Anon. 1792). It was identical in format, and composition,

with the plates colored by hand as was the 1787 original. The possibility that Beaumont's illustration

might prompt visits to Nice were overtly mentioned in a review of the 'pirated' version in a London

serial, The Monthly Review (Anon. 1793): "The present volume is not only an elegant but a splendid

production. The plates consist of twelve views, which are very neatly engraved, and remarkably well

coloured and the romantic and pleasant appearances which they afford are admirably adapted to

tempt the beholder to visit those scenes themselves, ....".

The plates were not signed so that while the original drawings were no doubt those of Beaumont,

the engraver is anonymous. Some have assumed that Beaumont engraved the plates and that the

watercolor is the work of Gabriel Lory l'Ancien (e.g., auction house descriptions). However, there

appears to be no documentary evidence that Beaumont ever did any engravings, nor is there any

documentary evidence of the identity of the colorist. The colorist should be credited with some

creativity, adding items not apparent in the engraving. For example, close inspection of the plates by

Joëlle Defaÿ, former scientific illustrator, Archivist and Librarian of the Museum d'Histoire Naturelle

de Nice, of original copies of the work held in the Bibliothéque de Cessole in Nice, revealed that in

plate 9 (shown here as Fig. 5), the sailboat, and its reflection in the water, in lower center of the plate,

is not in the engraving; it was added by the colorist.


Figure 2. Plate 1. This drawing shows the gulf of St. Hospice, the site of the annual tuna fishery. The little

tower on the right is kind of casement where there cannons to defend the coast. This place is half a league

from Ville Franca, its peninsular situation is very agreeable. Actual size of image 14.0 x 23.7 cm.

Figure 3. Plate 6. This shows the cavern of St. André, about a mile distant from the castle. The road to it is

very difficult, although many English ladies have passed it. This place is so lonely, notwithstanding its vicinity

to the town, that one could suppose oneself transported among the most solitary mountains of the Alps. In the

summertime, you can pass under the arch and penetrate into the other side of the cavern where you find

yourself in a very lonely valley. Actual size of image 14.0 x 23.7 cm.


Figure 4. Plate 8. A view of the church of St. Pons and also the convent of Cimiez, on the left of the drawing.

You see also the course of the Paillon which washes the walls of Nice, and winds through the bottom of the

valley on the side of to the Great Turin Road. This is a most delightful walk, from the goodness of the road and

the shade which may be enjoyed. Actual size of image: 25.8 x 45.2 cm

Figure 5. Plate 9. This drawing shows the entrance to the port of Lympia, and the two extremities of the town

behind the castle which was built on the mountainside that covers the port, as well as the different views of the

country which are on the border of the sea half-way to the Var. The ruins seen in the foreground of the picture

are the remains of a Lazaret, which the sea has almost destroyed. Actual size of image: 25.8 x 45.2 cm


Figure 6. Plate 11. This drawing represents a great part of the coast of Nice, the town and the surrounding

mountains, with the entrance to the port of Ville Franca and its lighthouse. This view is from above the valley of

Magnan, on the Var road. I have given in the drawing an idea of the manner in which the grounds are

cultivated. By this view may be seen how much of country is covered with olive trees.

Actual size of image: 25.8 x 45.2 cm.

Figure 7. Plate 12. The port of Villa Franca, and the back of the castle of mount Alban. By this drawing, a

judgement may be formed of the size of the road. The town and the castle are also shown. This port was

formerly, according to the opinion of some learned men, the port of Hercules.

Actual size of image: 25.8 x 45.2 cm.


2.2. Select Views of the Antiquities and Harbours in the South of France; with Topographical

and Historical Descriptions.

Figure 8. The cover of Beaumont's 1794 "Select views of the South of France, etc. and the first plate,

"Harbour of Antibes". The plate legend reads: This view taken from a height north west of the city, contiguous

to the road that leads to the village of Biot. Towards the east in the background, and on the right of the drawing

lies the quadrangular fort which protects the town and the port. Still farther to the right lies the city of Nice,

which from its distance, is but imperfectly seen, so are the range of mountains of Turbia and Monaco. On the

west are the city and harbor of Antibes. Actual size of the image is 20 x 30 cm.

Beaumont introduced the album with the following long sentence: "The South of France preſents a

large and extensive field for obſervation, abounds in rich scenery, and contains many beautiful and

magnificent remains of antiquity; in exploring which, the author of this work frequently employed his

pencil: and, flattering himself that his labors may, in part at least, be acceptable to the English nation,

he has undertaken a selection of such Views as appeared to him most interesting; in which number he

includes those of its three principal harbours, viz. Toulon, Marseilles, and Antibes"

The work consists of 15 aquatint engravings of landscapes by Apostool, after the drawings of

Beaumont, and 53 pages of text. Aquatint engravings differed from standard engravings in that,

instead of engraving directly a metal plate, usually copper, the plate was first treated to add a thin film

of resin. Engraving the surface produced lines as series of very small points, yielding a printed image

that resembled a painting. Apostool's prints were monochromatic using a sepia-colored ink. Such prints

could also be hand colored using water colors. The "Select Views ..." work was published in two

editions, one monochromatic and one with colored plates. The later version was enthusiastically

reviewed in The Monthly Review (Anon. 1794), stating "Those who have seen his Travels over the

Rhaetian Alps and his other works, will not doubt the elegance of the execution of the plates now

before us, ...."


Figure 9. Plate 3. This view is from an eminence which lies between Fort Pharon and Fort Rouge, or St.

Antoine, northwest of the city. La Croix des Signaux is the most distant mountain, the extremity of which forms

Cape Cepet. Fort Malgue is situated on an elevation towards the east, and on the right side. At the foot of this

hill stands Grosse Tour. On the left are seen, through the trees, the forts Balaguier and L'aiguillette. Between

forts and Grosse Tour lies the city of Toulon and its harbors, which are both in the inner road. Actual size of

the image is 20 x 30 cm.

Figure 10. Plate 6. Harbor and City of Marseille. This view represents the entrance of the harbor, as also its

forts, and is taken from the extremity of the Island of Rotoneau, south west of the city. On the north, or to the

left of the drawing, stands the fort of Notre Dame de la Garde, and on the right, Fort St. John. The stone piers,

which are seen in the water, serve to support a large chain, which shuts the harbor every night. These piers

are fixed at about 18 fathoms and a half from each other. Actual size of the image is 20 x 30 cm.


2.3. Travels through the Maritime Alps, from Italy to Lyons, Across the Col de Tende, by the

way of Nice, Provence, Languedoc, etc. with Topographical and Historical Descriptions to

which are added Some Philosophical Observations on the Various Appearances in

Mineralogy, etc. found in those countries.

Figure 11. The cover of Beaumont's 1795 "Travels through the Maritime Alps etc.," plate 14 Villa Franca. In

the drawing, no. 14, I have given a view of the beautiful bay and surrounding mountains, taken from between

the eastern battery and the column which marks the freedom of the port.

Actual size of the image is 20.2 x 30.4 cm

The "Travels through the Maritime Alps, etc." contains not only 18 plates, but also a considerable

amount of text, 127 pages, distinguishing it from Beaumont's earlier works. It was designated by Boyer

(2002) as the first travel guide to the region. It appears to be an apt description as it contains not only

landscapes but also descriptions and travel advice. For example, the text concerning Villefranche

(Villa-Franca) begins with the paragraph: "It is usual for strangers, during their residence at Nice, to

go in parties to Villa- Franca by sea, and return by land; but, owing to the badness of the road, which

is not passable for carriages (although supposed to have formed a part of the ancient Via Aureliana),

and the extreme steepness of Mont Alban, this little excursion must be effected on horses or mules;

and, as the latter are by far the safest, I would particularly recommend them, in preference to the first.

Never shall I forget with what pleasure I made my last trip to Villa-Franca, and the beautiful and

enchanting view I enjoyed from the environs of the fortress of Mont Alban, which stands on the

mountain contiguous to the road: in fact, I know of none to equal it, either for extent or variety of

objects (all fantastically beautiful), except the one seen from the Chartreuse at Naples."

The Travels through the Maritime Alps was at first very favorably reviewed in the serial The British

Critic (Anon. 1795). The long review with excerpts began with "This volume claims our attention, and

that of the public, on more accounts than one. It contains all the splendid perfections of the art of

printing, and exhibits scenes of nature hitherto but little known, from the difficulty with which they can

be explored." A later review (Anon. 1798f), was much less laudatory, and ended with the declaration

that the plates "... form the principal recommendation of the work".


Figure 12. Plate 16, Monaco. The situation of the city is extremely picturesque and romantic, seated, as it

were, at the edge of a rock which projects considerably into the sea. An accurate idea of it is given, which

likewise includes that of the Turbia, or Trophæum Augusti, which ſtands on the summit of the mountain northwest

of the city. This view was taken in the neighbourhood of Cape St. Martin.

Actual size of the image is 20.2 x 30.4 cm

Figure 13. Vintimiglia. The town of Vintimiglia, a view of which, taken from the banks of the Nerva, is most

agreeably situated at the mouth of the Roia, a river already mentioned in the course of the present work. It is

likewiſe seated at the foot of a stupendous rock, on which stands the fort which protects the town and coast.

This town, which takes its present name from its distance to Nice, which is exactly twenty miles, was anciently

called Albintemelium. It does not appear to have been a place of particular note, or at least there are no

vestiges remaining to indicate the ſtate of its former splendour. Actual size of the image is 20.2 x 30.4 cm


Conclusion

With this essay, hopefully the remarkable work of Albanis Beaumont and Cornelius Apostool will

become known to wider public, beyond collectors of old books and historians of the Côte d'Azur.

While many early writers such as Smollett (1766), Sulzer (1780), Davis (1807) and Millin (1816)

wrote of the beauty of the region of Nice and encouraged traveler to visit, it was Beaumont and

Apostool who actually showed the beauty of the region of Nice. Thus they too should share in the

credit (or blame, depending on your point of view!) for bringing people to the Côte d'Azur.

Acknowledgements

Sincere thanks go to Joëlle Defaÿ for providing basic instruction in the graphic arts, and introducing

me to the incredibly helpful and patient Elise Carbou-Hansson and Benjamin Person of the

Bibliothèque du Chevalier de Cessole (City of Nice). All of the images of Beaumont's "Voyage

Pittoresque..." were kindly provided by the Bibliothèque du Chevalier de Cessole.

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Arts et sciences


Two centuries of arts and science in Nice

and Villefranche sur Mer: 1) Pioneers: 1800 to 1900

Deux siècles d’arts et de sciences à Nice et Villefranche sur Mer :

1) Les anciens : de 1800 à 1900

Christian Sardet 1

1

Sorbonne University, CNRS, Laboratoire de Biologie du Développement, Institut de la Mer de Villefranche sur Mer

(IMEV), 06230, France, christian.sardet@imev-mer.fr

ABSTRACT. We tell the story of exploration of the flora and fauna in the Nice region, and especially pelagic organisms,

the work of some twenty biologists associated with painters. In 1800, Antoine Risso, Jean Gabriel Prêtre, François Péron

and Alexandre Lesueur described and painted flowers, fish and some marine invertebrates. In the middle of the century,

Jean Baptiste Vérany, a naturalist from Nice, welcomed prestigious German and Swiss biologists - Johannes Müller,

Rudolf Leuckart, Ernst Haeckel and Carl Vogt - who described andstudied little known organisms such as siphonophores

and radiolarians. And in the 1880s, Hermann Fol, Jules Barrois and Alexis Korotneff set up a marine station in

Villefranche sur Mer, recognized as an exceptional site for the study of plankton. During this period, the Natural History

Museum of Nice was enriched by exceptional collections of flowers, fish and funghi assembled by Risso, Vérany and

Jean Baptiste Barla, who employed Vincent Fossat for his talent as painter / illustrator. By the end of the century, the Nice

region attracted foreign royalty and aristocrats, visiting biologists and impressionists painters. In a companion article

(Sardet 2025 / 2 Modern era: 1970 to 2024) we show that organisms studied in the 19 th century are still the subject of

research at the Institut de la Mer de Villefranche (IMEV).

RÉSUMÉ. Nous racontons l’histoire de l’évolution des connaissances de la flore et la faune dans la région niçoise et en

particulier celle de la faune pélagique. Dans les années 1800, Antoine Risso, Jean Gabriel Prêtre, François Péron et

Alexandre Lesueur décrivent et peignent les fleurs, les poissons et des invertébrés marins. Au milieu du siècle, le

naturaliste niçois, Jean Baptiste Vérany accueille des savants allemands et suisses

- Johannes Müller, Rudolf Leuckart, Ernst Haeckel et Carl Vogt - qui influencent les recherches et le destin de la biologie

dans la région par leurs descriptions illustrées d’organismes jusqu’alors ignorés comme les siphonophores et les

radiolaires. Et dans les années 1880, Hermann Fol, Jules Barrois et Alexis Korotneff créent une station marine

accueillante à Villefranche sur Mer, reconnue depuis comme un site exceptionel pour l’étude du plancton. A partir du

milieu du 19 ème , le Muséum d’Histoire Naturelle de Nice s’enrichit des collections de Risso, Vérany et Jean Baptiste Barla

épaulés par Vincent Fossat, un peintre / illustrateur talentueux. A la fin du siècle, la région niçoise attire les souverains et

aristocrates anglais et russes, des biologistes et des peintres impressionistes. Dans un article compagnon (Sardet 2024

/2 Les modernes – de 1970 à 2024), nous montrons que les organismes explorés au 19 ème siècle sont toujours l’objet de

recherches à l’Institut de la Mer de Villefranche (IMEV).

KEYWORDS. Nice, Villefranche sur Mer, plankton, protists, Antoine Risso, François Péron, Alexandre Lesueur, Jean

Baptiste Vérany, Jean Baptiste Barla, Vincent Fossat, Ernst Haeckel, Johannes Müller, Carl Vogt, Hermann Fol, Jules

Barrois, Alexis Korotneff.

MOTS-CLÉS. Nice, Villefranche sur Mer, plankton, protists, Antoine Risso, François Péron, Alexandre Lesueur, Jean

Baptiste Vérany, Jean Baptiste Barla, Vincent Fossat, Ernst Haeckel, Johannes Müller, Carl Vogt, Hermann Fol, Jules

Barrois, Alexis Korotneff.


Introduction

In the early 1800s, Nice was a provincial town of 40,000 inhabitants, accessible only by stagecoach

or boat. In 1860, under Napoleon III, the county of Nice, which previously belonged to the kingdom of

Piedmont-Sardinia, became part of France. Four years later, Nice was accessible by rail. Starting in the

middle of the century, visitors flocked to the Côte d'Azur, drawn by the presence of Russian, English

and Belgian royalties and artistocrats who came with their courts to enjoy the mild winter climate.

Artists and biologists were attracted by the diversity of landscapes, flora and marine fauna.

Villefranche sur Mer, situated 7 km east from Nice, was a free port when it was founded, and in the

1800s the bay and city of Villefranche became a place of refuge, supply and rest for ships and their

crews. Complicated political agreements brought France and the Kingdom of Piedmont-Sardinia closer

to the Russian empire. As a result the bay of Villefranche harbored a large number of Russian, English,

French, Italian and even American navies. Today, the bay of Villefranche welcomes cruise ships that

flood the French riviera with their passengers (Braconnot & al. 2004).

The Nice region (Figure 1) boasts a remarkable seafront, bordered to the west by the sedimentary

delta at the mouth of the Var river, and to the east by the bay of Villefranche sur Mer. This bay opens

onto a seabed hundreds of meters deep, through which the Ligurian current flows carrying a variety of

planktonic organisms - from bacteria and fish larva to jellyfish - which drift together. Depending on the

season and winds, some of these plankton find themselves trapped in the natural receptacle of the bay

of Villefranche sur Mer and its deep waters. Here, pelagic organisms, usually found only on the high

seas, come close to shore and are accessible with small boats.

From the early 19 th century, naturalists from Nice started exploring the local flora and fauna. They

befriended and helped European colleagues explore the coast and collect plankton. During this period,

anatomical descriptions were illustrated with drawings by professional artists. These illustrations were

transformed into engravings for publication in the few scientific journals of the time, or in

monographs.

The 19 th century was a time when illustrated books and newspapers became popular. Travel was

increasingly facilitated by coal-powered ships and trains. In the second half of the 19 th century,

photography and illustrated publications took off. Biologists developed their networks of universities,

academies and publishers across Europe (Jessus & Laudet, 2022). Researchers helped each other,

collecting and studying the extraordinary diversity of marine organisms, stimulated by new ideas on

classification and evolution and the emerging cell theory. European and Russian professors and

students explored the Mediterranean coasts in the 1850s, staying in the Nice region to study and

illustrate jellyfish, siphonophores, appendicularians, radiolarians and more. At the end of the 19 th

century, thanks to young Swiss, French and Russian biologists, set up a marine station in Villefranche

sur Mer which permitted scientific research and in-depth exploration of local plankton, a tradition

continuing to this day ( Anon. 2010, 2024, Trégouboff 1983, Dolan 2014, 2024).

In this article we describes a century-long progression. In a second article, we show how Côte

d'Azur organisms described 100 to 200 years ago still inspire the research and pictorial works of

researchers at the Villefranche sur Mer marine station (IMEV) in the 21 st century (see companion

article : Sardet 2025 / 2 Modern era: 1970 to 2024).


Fig. 1. Nice region: A - The coast of Nice in the 18th century. B - The bay of Villefranche sur Mer in the 18th

century. On the right, buildings on the site of today's port de la Darse.

C - The port of La Darse in Villefranche sur Mer at the end of the 19th century (postcard)

D - The bay of Villefranche sur Mer at the end of the 19th century (postcard).

E & F - Current views of the bay of Villefranche sur Mer.

G - The port de la Darse (spelled DARCE here) and the Lazaret in 1748

H - Topography of the seabed in the Nice area


1800 - 1900: Naturalists and painters explore Nice's biodiversity

Naturalist began exploring the region in the early 1800s notably Antoine RISSO, author of

remarkable publications on the local flora and fauna, illustrated by Jean Gabriel PRÊTRE. In 1809

RISSO welcomed and helped zoologist François PÉRON and watercolorist Charles-Alexandre

LESUEUR, visiting Nice, to describe and illustrate gelatinous plankton organisms, jellyfish, mollusks

and ctenophores. In the mid-1800s, Jean Baptiste VERANY, along with another Jean Baptiste,

BARLA, created the Nice Natural History Museum. With the help of a talented local painter, Vincent

FOSSAT, they contributed extraordinary collections. In the 1850s, VÉRANY welcomed and

befriended scientists from Germany and Switzerland, attracted by the rich planktonic fauna abundant

in the bay of Villefranche sur mer. These visitors - Johannes MÜLLER, Rudolf LEUCKART, Carl

VOGT and Ernst HAECKEL - were brilliant and influential biologists who described and artistically

illustrated little known pelagic organisms such as siphonophores and radiolarians. Research into

marine biodiversity gained momentum with the creation in 1881 of the first laboratory in Villefranche

sur Mer by Hermann FOL and Jules BARROIS. In 1886, Alexis KOROTNEFF took over and

established the Station Russe de Zoologie in Villefranche sur Mer, the forerunner of IMEV, the Institut

de la Mer de Villefranche, today's marine station. All these men, whose dates of birth and death are

indicated, succeeded one another over the course of the century (see chronological chart below and

Figure 2).

1770 1780 1790 1800 1810 1820 1830 1840 1850 1860 1870

Jean Gabriel PRÊTRE (1768-1849)

Painter-illustrator attached to the Muséum d'Histoire Naturelle, Paris

François PÉRON (1775 - 1810)

Zoologist aboard the Baudin expedition exploring the Austral Lands (1800 -1803)

Antoine RISSO (1777 - 1845)

Pharmacist and botanist from Nice, author of monographs, some illustrated by Prêtre

Charles-Alexandre LESUEUR ( 1778 - 1846)

Painter and illustrator from Le Havre, who joined Péron for the Baudin expedition

Jean Baptiste VÉRANY (1800 – 1875)

Naturalist from Nice, author of artistic monographs on cephalopods

Johannes MÜLLER (1801 - 1858)

German zoologist and protistologist, Berlin professor and mentor to Haeckel

Jean Baptiste BARLA (1817 - 1890)

Naturalist from Nice who built up collections of mushrooms and fish with Fossat

Carl VOGT (1817 - 1895)

German / Swiss zoologist, professor in Geneva, evolutionist and socialist activist

Vincent FOSSAT (1822 - 1897)

Painter from Nice, employed by Barla to illustrate extraordinary collections

German zoologist, friend of Vérany. Professor in Giessen, then in Leipzig

Rudolf LEUCKART (1822 - 1898)

(1934- 1919) Ernst HAECKEL

German zoologist, professor and rector at the University of Jena. Traveller and artist

(1845-1892) Hermann FOL

Swiss zoologist, professor in Geneva, founded a laboratory in Villefranche with Barrois

(1851 - 1915) Alexis KOROTNEFF

Russian zoologist, professor in Kiev, founded the Villefranche Russian Zoological Station

(1852 - 1943) Jules BARROIS

A zoologist and professor in Lille, he founded the first laboratory in Villefranche with Fol


Fig. 2. The characters in this story and some of their publications:

A - Antoine RISSO and the first volume his 5-volumes "Histoire Naturelle de Nice et des Alpes Maritimes"

(1826)

B - François PÉRON, Alexandre LESUEUR and the book dedicated to theirsojurn on the Côte d'Azur (G.

Baglione & J. Goy, 2009)

C - Jean Baptiste VÉRANY and his book of chromolithographs of cephalopods (1851)

D - Jean Baptiste BARLA and Vincent FOSSAT, whose watercolor illustration of a fish illustrates the cover of

a recently published book

E - Rudolf LEUCKART and Johannes MÜLLER and their books translated into English

F - Ernst HAECKEL and Carl Vogt and their best-known publications

G - Hermann FOL and his monograph on fertilization (1879)

H - Jules BARROIS and the cover of his thesis (1878) and prtrait of Alexis KOROTNEFF


1800 - 1830: Risso and Prêtre describe and illustrate the flora and fauna of Nice

The history of the naturalist quest in the Nice region began in the early 19 th century with Antoine

Risso (Gasiglia 1970, Dolan 2023a). Born in Nice in 1777 into a modest family, Risso was orphaned at

an early age. He was raised by an uncle who directed him to be an apprentice to a pharmacist and

botanist starting at the age 12. Risso managed to open his own pharmacy at age 26 and was appointed

assistant curator of the departmental botanical garden. After describing local plants such as olive and

citrus trees, Risso turned his attention to the animals that local fishermen brought back in their nets

(Risso 1810, 1813). He not only characterized and identified fish, but also the gelatinous catch that

fishermen call "carminaria" - literally, sea-meat, a tangle of jellyfish, mollusks and other soft,

transparent organisms(Figure 3). Risso revived them in aquariums, carefully observing and describing

them. He also caught the organisms from small boats, taking temperature readings at the same time. In

1810, he published "Ichtyologie de Nice", a natural history of fish in the Alpes Maritimes, with 11

unsigned plates.

In 1826, after selling his pharmacy, Risso published an ambitious 5-volume "Histoire Naturelle".

The talented illustrator, Jean Gabriel Prêtre produced 44 plates, which were reproduced by half a dozen

engravers (Risso 1826). The fifth volume contained descriptions and plates devoted to marine

invertebrates (Figure 3). Unfortunately, little is known about the precise dates and conditions under

which these first representations of the marine fauna of Nice were produced, and we don't know

how Risso and Prêtre worked together or with the publishers. The 5-volume edition must have cost a

fortune! From a family of Genevan artists, Prêtre was about ten years older than Risso, and by the time

they worked together, Prêtre had already produced some remarkable illustrations of birds and other

animals for the books by George Cuvier and his colleagues at the Muséum d'Histoire Naturelle in Paris

(Cuvier 1816). Over 700 illustrations by Prêtre are known, an impressive body of work (Dolan 2023a).

Some of the organisms immortalized by Risso and Prêtre, such as jellyfish and sea urchins, were

further explored by their successors starting in the 1850s. First by German and Swiss scientists such as

Ernst Haeckel and Carl Vogt, who explored Mediterranean coasts in search of planktonic organisms,

including unicellular protists (Dolan 2019). This research was amplified in the 1880s, when a

laboratory was set up in Villefranche sur Mer (Anon 2010, 2024, Trégouboff 1983). This laboratory

was designed to host visiting European researchers, as had already been done at the Roscoff and

Banyuls sur Mer marine stations and in Naples (Fischer 2002, Debaz 2005, Groeben C. 2020, Jessus

& al. 2021)

In fact, the pioneering research carried out in the 19 th century on the anatomy, physiology and

reproduction of terrestrial and marine organisms forms the basis of a body of knowledge that continues

to this day. Teachers, researchers and members of the region's academic and associative communities

continue to study and introduce the public to the rich local marine flora and fauna. Four study centers -

the Université de Nice Sophia Antipolis (UNSA), the Institut National de Recherche Agronomique

(INRA d'Antibes / Sophia Antipolis), the Centre Scientifique de Monaco and the Institut de la Mer de

Villefranche sur Mer (IMEV) - are doing advanced research work in these fields. Some examples of

research carried out at IMEV over the last 50 years are described in the companion article (Sardet

2025).


Fig. 3. Illustrations by painter Jean Gabriel Prêtre for naturalist Antoine Risso

A - Molluscs in volume 4 of Risso's « Histoire Naturelle » (1826)

B - Fish in volume 3 (plate 16) of the « Histoire Naturelle »

C - Brittle stars and sea anemones in volume 5 (plate 7) of « Histoire Naturelle »

D - The Equorea Rissoana jellyfish and sea urchin tests in volume 5 (plate 7)

QR code: other illustrations available on the Aquaparadox website (John Dolan, IMEV)


1809-1910: Péron and Lesueur discover the Nice area and its marine fauna

Risso's work on the fish of the Alpes-Maritimes region was remarkable. François Péron, a young

zoologist, and his painter/illustrator friend Alexandre Lesueur wanted to meet Risso and observe the

pelagic fauna they had already observed and described during the Baudin expedition in the Southern

Ocean (1800-1804, Baudin 2001). On their return, they lived and worked together in Paris, where they

wrote and illustrated a monograph entitled "Voyage de découvertes aux terres australes". This

expedition, commissioned by Bonaparte and organized by Nicolas Baudin, took Péron and Lesueur as

far as Australia and Tasmania, collecting over 100,000 samples, including 2,500 previously unknown

species. The pelagic fauna of the Atlantic and Pacific - jellyfish, molluscs, ctenophores, salps,

pyrosomes - was described and illustrated for the first time with accuracy and precision by Péron and

Lesueur, who was also an accomplished engraver (Péron & Lesueur 1809).

Lesueur was born into a middle-class family in Le Havre in 1778. Enlisted at the age of 21 as

assistant gunner, he quickly established himself as an illustrator for the Baudin expedition, and

befriended Péron. Born in Allier in 1775, Péron had a difficult youth. He lost his right eye and was

taken prisoner during the French Revolution in 1792. After studying medicine, Péron was hired as a

zoologist/anthropologist by the Baudin expedition. This 2-ship voyage cost the lives of many

explorers, including Captain Baudin himself, who died of tuberculosis in Mauritius in 1803. Péron also

contracted tuberculosis, and on his return to France, his doctor advised him to spend the winter in the

south of France for his health.

In 1809, Péron and Lesueur traveled to Nice by stagecoach and boat on the Rhône river, then

crossed Provence (Goy & Baglione 2009, Baglione 2024). After a 3-week journey, Péron and Lesueur

arrived and then stayed in Nice for 6 months. To collect pelagic organisms, they benefited from the

help of Risso and the local authorities. Lesueur recounts: "The Commissioner of Nice provided us with

a boat, skipper and four strong sailors chosen from the deserters who were in prison. They came on the

day indicated to took us on board, and then we began our experiments on the Mediterranean". Péron

and Lesueur collected, then observed some gelatinous organisms in aquariums. They described in

particular a remarkable ctenophore, Cestus veneris or Venus belt an organism that can still be observed

every spring in the bay of Villefranche sur Mer(Figure 4B). They also measured temperatures and

noted that gale force winds caused changes in the pelagic fauna, a first step towards oceanography and

ecology.

The winter of 1809 was harsh and cold, but Péron and Lesueur managed to get out in the boat and

collect, describe and illustrate pelagic fauna and some benthic organisms such as ascidians and mollucs

(Figure 4A, 4C). Unfortunately, Péron's health deteriorated rapidly. Lesueur accompanied him to Paris,

then to his native Massif Central, where Péron died in 1810, in the hope that Lesueur would publish

their observations. Lesueur did so, before leaving for the USA, where he took part in several

expeditions exploring and illustrating the fauna, flora, native peoples and landscapes until 1837 (Dolan

2020). Lesueur returned to Le Havre in 1845 to create the Natural History Museum where he was the

first director and curator, shortly before his death in 1846. Bequeathed by his family to the Museum

after his death, Lesueur's works were forgotten. Rediscovered, his remarkable drawings and

watercolors have been the subject of several exhibitions (Baglione & Crémière 2009).


Fig. 4. Alexandre Lesueur's watercolours on vellum of the marine fauna of Nice

A - Ciones (Ciona intestinalis), ascidians measuring a few centimeters.

Inv. MHNH 75017, Size of work 24.4x40.8 cm

B - Belt of Venus (Cestus veneris), a ctenophore that can measure up to 80 cm.

Inv. MHNH 67 050, Size of work: 28.0x43.2 cm

C -Firoles (Pterotrachea sp.), planktonic gastropod mollusks belonging to the heteropods. They measure a few

centimeters. Inv. MHNH 72010, Size of work: 29.8x43.9

These works are kept at the Natural History Museum of Le Havre (MHNH).

QR code: These and other works can be seenas part of a video interview with MHNH

curator Gabrielle Baglione.


Barla, Vérany, Fossat: a new generation completes the biodiversity inventory

Antoine Risso's impetus to study the biodiversity of local flora and fauna was amplified by a new

generation of naturalists from Nice. Jean Baptiste Vérany - 23 years younger than Risso, like Risso,

was a pharmacist and botanist. He briefly ran the family pharmacy, helped Professor Franco Bonelli

build collections for Turin's Natural History Museum, and guided visitors and tourists interested in

local flora and fauna. Vérany developed a passion for molluscs, particularly cephalopods. In 1851, he

published a monograph whose delicately colored and transparent chromolithographs are considered

works of art (Verany 1851, 1862, Dolan 2022a). Vérany’s collections formed the basis of the Musée

Municipal de la Ville de Nice created in the old town in 1846. Ten years later, under the impetus of

another Jean Baptiste, the naturalist Jean Baptiste Barla, a new more ambitious museum was born : the

Muséum d'Histoire Naturelle de la Ville de Nice.

Jean Baptiste Barla was from a family which came in Nice from Piedmont and had made their

fortune trading in cod liver oil and hemp. He learned botany from his uncle, studied, played the violin,

traveled and met naturalists. Barla developed a passion for the region's flora, especially orchids, as well

as fish and mushrooms, whose diversity is exceptional in the mountainous back country of Nice

(Trimbach 1996). Barla was assisted in his task by a virtuoso gouache and watercolor artist, Vincent

Fossat (Defaÿ 1998, Dolan 2022b). As Barla was himself an accomplished artist, it is sometimes

difficult to distinguish between Fossat and Barla’s paintings. From 1851 onwards, Barla was able to

live on his income, since he inherited a substantial fortune from his grandfather. Barla employed Fossat

as a painter for 3 to 5 francs a day from 1853 until his death in 1891.

Fossat, born Del Fossat, was the seventh of twelve children in a family that had come to Nice from

the Imperia region in the 16 th century. Orphaned at the age of 13, with little education, Fossat made a

living from various day jobs - fisherman, gardener, coachman, servant. He began to paint Nice and the

surrounding area on commission, sometimes under assumed names. Fossat's artistic career took on a

new dimension with the naturalistic works he produced for Barla, and also for Vérany. These included

thousands of watercolors and gouaches of plants, fish and mushrooms. In 1855, they also published a

number of portraits of marine invertebrates, sea urchins, starfish and brittle stars (Figure 5).

Barla and Fossat's best known works are the painted cast/models of mushrooms and cardboard

filled, then gouache imprints of fish - the exsiccata - which constitute a kind of fish herbarium (Barla

1892). The painted mushrooms and the 1,500 cardboard fish imprints are a treasure trove preserved at

the Muséum d'Histoire Naturelle de la Ville de Nice, a center of intense naturalist activity in the 19 th

century (Chamagne-Rollier & Defaÿ 2013).

Financed by Barla and supervised by Vérany, the Museum was created in 1865 and directed by

Barla. Barla transferred to the City of Nice the imposing building he had built at his own expense. This

fine institution regularly exhibits its marvellous collections of works by Risso, Prêtre, Verany, Barla

and Fossat and their successors. The Museum has recently entered a new phase: the digitization,

restoration and genetic exploration of its conserved works. It also organizes cycles of scholarly

conferences paying tribute to its glorious past.


Fig 5. Moulded and painted works by Jean Baptiste Barla and Vincent Fossat

A - Painted cast/models of mushrooms (Amanita muscaria), photo @Jean Marc Alpesse

B - Exsiccata (fish skin stretched over cardboard and repainted) of a red scorpion fish

(Scorpena scrofa or Capoun in Niçois)

C, D - Eel (Anguila vulgaris, Anghilla in Niçois), watercolor (close-up) by Fossat. Inventory no. 2005.0.730; 71

(plate no.) POP platform, French Ministry of Culture

E - Starfish (Palimpus membranaceus), watercolor by Fossat. Inventory no. 2005.0.1016; 371 (plate no.) POP

platform, French Ministry of Culture

F - deep sea urchin ( Diadema europea ), watercolor by Fossat. Inventory number 2005.0.1011 ;371

(plate no.), POP platform, French Ministry of Culture

G - brittle stars (Ophiomyta pentagona), watercolor by Fossat, size 27.0x 35.2 cm. Inventory no. 2005.0.1025;

380 (plate no.) POP platform, French Ministry of Culture

QR code: gives access to John Dolan's illustrated article on Vincent Fossat (Dolan 2022b)


1850: Visiting foreign scientists and beginnings of the marine station in Villefranche

Our understanding of the principles of life and evolution was revolutionized in the 19 th century.

From the 1830s onwards, Theodor Schwann, Matthias Schleiden and their microscopist colleagues

developed the foundations of a cell theory. It states that animals, plants and all living things are made

up of cells that reproduce by division (Sardet 2023). In the 1850s, the identification of natural selection

as the driving force behind evolution by Charles Darwin and Alfred Russel Wallace revealed that

organisms evolve and are linked by common ancestors. Naturalists, zoologists and botanists began to

compare the anatomy and physiology of the organs, tissues, gametes and embryos of animals and

plants, as well as unicellular eukaryotes (the type of cells which have a nucleus) known as protists

(Jessus & Laudet 2020). Leading figures such as Henri Lacaze-Duthier ( Jessus & al. 2021) and Anton

Dohrn ( Groeben 2020) set up major marine stations (Roscoff in 1872, Naples in 1874) in addition to

those already in existence (Concarneau 1859, Arcachon, 1863) or considered such as Messina where

Dohrn worked in 1868 before he set up the Naples station. These marine stations gave researchers

access to the great diversity of marine flora and fauna.

A new era dawned with the visits to Nice of German and Swiss teachers in the 1850s. One

particularly important visitor was Johannes Müller - an influential professor in Berlin and mentor to a

whole new generation of cell biologists (Müller 1843). Like his colleague Rudolf Leuckart, a professor

in Giessen (Germany), Müller traveled the coasts of Europe with his students, observing marine

creatures through the microscopes they carried with them.

Aided by his friend Verany, Rudolf Leuckart stayed in Nice in 1853 to study marine invertebrates -

worms, jellyfish and crustaceans. Then, in 1856, Müller and his family visited Nice joined by Swiss

biologist Albert Kolliker and students, including Ernst Haeckel, who was 22 years old at the time.

They collected plankton in the bay of Villefranche sur Mer, describing and drawing protists and

gelatinous organisms (Dolan 2019). In 1864, shortly after the death of his young wife, Haeckel

returned to Villefranche for a few months. He produced his first "art and science" drawings of jellyfish

and radiolarians ( Figure 6). By the end of the 19 th century, Haeckel was a celebrated scientist, artist

and philosopher, spreading the theory of evolution throughout Europe with his elder compatriot Carl

Vogt, one of the most influential biologists of his time (Vogt 1848). After studying medicine in

Germany and France, Vogt first drew attention to the phenomenon of programmed cell death in 1842.

He stayed in Nice and Villefranche in 1851 - 52 and published extraordinary descriptions and

representations of siphonophores in his monograph entitled "Recherches sur les animaux inférieurs de

la Méditerranée" (Vogt 1853). Vogt was also a political figure. Nicknamed "the revolutionary

scientist" for his socialist ideas and causes, Vogt took refuge in Switzerland. In the early 1850s, he

became a professor at the University of Geneva, where he later served as rector.

Praising the merits of plankton in the bay of Villefranche-sur-Mer in 1876, Vogt was the first to

advocate the necessary creation of a "marine station" in Villefranche like those already existing

elsewhere (Vogt 1876). A few years later, two young biologists - Jules Barrois, from Lille, and

Hermann Fol, a zoologist from Geneva inspired by Vogt - set up their first laboratory in Villefranche

sur Mer (Anon. 2010, 2024, Dolan 2024).


Fig 6. Plankton works by Johannes Müller, Rudolf Leuckart, Ernst Haeckel and Carl Vogt

A - Radiolarians discovered in Nice and Villefranche by Müller in 1856

B - Radiolarians observed in Nice and Villefranche in 1864, published by Haeckel (Dolan 2019)

C - QR codes: access to the Aquaparadox website (John Dolan, IMEV)

Left: Species discovered in Villefranche (Dolan 2014). Right: Leuckart poster (Dolan 2023c)

D - Radiolarians, wall poster 29 by Leuckart. Drawing by Otto Bütschli, University of Vienna

E - Drawing of marine fauna by Vogt for his book "Ocean und Mittelmeer (Vogt 1848)"


1880: Fol and Barrois first, then Korotneff - birth of the Villefranche marine station

Five years after Vogt wished for a marine laboratory to be set up in Villefranche sur mer, Jules

Barrois, a young PhD from Lille University, and Swiss zoologist Hermann Fol granted this wish. Fol

was certainly inspired by his mentor Vogt. In 1878 Fol became Vogt’s colleague as a professor at the

University of Geneva. Six years earlier, Fol had already spent several months in Nice to take care of

his health, while collecting and observing planktonic organisms – mainly appendicularians and

pteropods (Dolan 2024).Barrois was only 30 years old when he set up his first laboratory with Fol. We

don’t know why Barrois was originally attracted to the Nice region, but he seems to have arrived as

early as 1879 (Fokin 2008). Barrois had written his thesis on the development of bryozoans and

nemerteans at the University of Lille and at the Wimereux marine station created in 1774 (Barrois

1877). In Villefranche, Barrois mainly studied metamorphosis in several species of echinoderms.

Hermann Fol was a brilliant character with a reputation for being quarrelsome (Dolan 2024, Mahé

& Sardet 2009). Born in 1845 in the Paris region into a family of Suiss bankers, Fol was sent at an

early age to study in Geneva with Edouard Claparède, a zoologist famous for his work on protists and

invertebrates. He was also a pupil of Müller and became a colleague and friend of Haeckel. Claparède

encouraged Fol to study medicine with Haeckel in Jena, and in 1869 Fol defended his medical thesis

working in Zurich and Berlin. The thesis concerned the anatomy and development of ctenophores,

considered the most ancestral planktonic animals (Sardet 2013, 2023, Dolan 2024).

When Fol returned to the Nice region in 1879, he was 36 years old. He had already made a major

discovery, being the first to describe and illustrate the penetration of spermatozoa into oocytes during

fertilization (Fol 1878, Buscaglia & Duboule 2002). Fol made his discoveries in Messina in the 1870s

using gametes from starfish, sea urchins and chaetognates. In 1878, he transferred his laboratory from

Messina to Villefranche. In extensive correspondence with his other mentor, Henri Lacaze-Duthier,

creator and director of the Roscoff and Banyuls marine stations, Fol mentioned the presence in 1879 of

other young researchers on the Côte d'Azur as well as a visit from Vogt(Jessus & Laudet 2022). From

1881 onwards, Fol commuted between the Villefranche-sur-Mer laboratory and Geneva, where he

taught embryology and carried out various activities - photography, monitoring microbial

contamination of drinking water, creation of a scientific journal, etc. (Dolan 2024). Based on his

research in Villefranche, Fol published articles on ciliates (tintinnids) and the curious rhizopod protist

Sticholonche zanclea (Fol 1883). His pioneering work was followed by research carried out over a

century later in Villefranche sur Mer (see companion article Sardet 2025 / 2 Modern era: 1970 to

2024).

In 1881-82, Barrois and Fol, encouraged by Charles Darwin and Henri de Lacaze-Duthier, set up the

first laboratory in Villefranche sur mer on the site of the former 18 th century galley slave prison in the

port of Villefranche (port de la Darse : Anon 2010, 2024). Barrois and Fol first occupied a tower in the

« lazaretto » (quarantine building) with equipment provided by Fol. Later Barrois and Fol set up the

Laboratoire de Zoologie Marine – under the aegis of the Ecole Pratique des Hautes Études – in the

larger galley slave prison which also served as a hospital. Fol and Barrois welcomed numerous

colleagues: some forty visitors and their research topics are listed year by year from 1881 to 1889

(Barrois & Fol 81-89). One of the visitors, the Russian zoologist Alexis Korotneff, also dreamed of

establishing a laboratory in the same building, but under the aegis of the Russian tsarist empire. The

Russian navy had for a time docked in the bay of Villefranche and used the premises as a coal depot.


Fig. 7. Works by Hermann Fol, Jules Barrois, Alexis Korotneff and Elie Metchnikoff

A – Fol's drawings of sperm penetrating into the oocyte (Fol 1878)

B – Barrois’s drawings of the development of the briozoan Pedicellina sp. for his thesis

C – Fol’s drawings of tintinnid ciliates in their lorica (see Dolan 2024)

D – Metchnikof’s drawing of gastrulation in a jellyfish (Metschnikoff 1886)

E – Korotneff’s drawings of the protist Sticholonche zanclea and its intracellular parasites


Two visitors to the Villefranche marine station – Elie Metchnikoff and Alexandre Kowalevski – are

famous for their scientific contributions. In 1886, the Russian zoologist Elie Metchnikoff was

welcomed to the Villefranche station where he described the formation of the gastrula in jellyfish. His

work that would be extended 150 years later at the marine station ( see companion article, Sardet

2025). For his research into the mechanisms of immunity, Metchnikoff shared the 1908 Nobel Prize for

Medicine with Paul Erlich. The other famous Russian visitor – Kowalevski – is considered the father

of evolutionary embryology. In particular, he demonstrated that ascidians are chordates, and as such

they were the ancestors of vertebrates. He also demonstarted that all animals undergo a gastrulation

phase during embryogenesis (Kowalevsky 1866, Jessus & Laudet 2024). He had already visited the

Nice region in 1878 - 79, and was welcomed at the Villefranche marine station in 1882 to study

embryogenesis and gastrulation in chitons (molluscs, see visitor lists: Barrois & Fol 1881-1889).

Curiously, Korotneff only appears as one of the visitors in the Barrois and Fol lists from 1882

onwards, although he was probably on site before that. In any case, in 1884, Korotneff founded the

« Station Russe de Zoologie » with the support of the Russian government's Ministry of the Navy,

which had the use and responsability of the galley slave building which was no longer needed by the

Russian war fleet. It was dubbed la " Maison Russe" by the locals ( Anon. 2010, 2024, Trégouboff

1983). Korotneff, who had studied zoology in Moscow, was appointed professor in Kiev in 1886. A

frequent traveler, he was only present in Villefranche from time to time. He was assisted on site by

Michael Davidoff, who succeeded Korotneff in 1915. At the marine station, Korotneff studied the

embryology of salps and ascidians. He was also interested in cnidarians and the protist Sticholonche,

about which Fol had already published (Figure 7, Fol 1883, Korotneff 1891).

Within the Maison Russe, relations between Barrois and Fol on the one hand, and Korotneff on the

other were cordial at first, but deteriorated after Korotneff's absence. Korotneff obtained the right to

occupy the premises from the French government (a Russian ally) and Fol and Barrois were expelled

from the building by the police in 1887 (Trégouboff 1983).

Russian influence and the Villefranche sur Mer marine station

The marine station occupied a large, 70 meter-long building originally constructed in 1769, to serve

as a prison/hospital for convicts, and prisoners of war (Figure 8, Anon 2020). It became known as la

« Maison Russe » in the mid-19 th century at the time of the establishment of a Russian colony in Nice

(Braconnot et al. 2004). In 1856, the Treaty of Paris ended the Crimean War, during which France,

England and the Kingdom of Piedmont-Sardinia (to which the County of Nice belonged at the time)

clashed with Tsarist Russia. In 1856, and again in 1859, the Empress Alexandra Feodorovna, widow of

Tsar Nicholas 1 er , landed in Villefranche and spent several months in Nice. She met the King of

Sardinia, who facilitated negotiations for the galley slave building to become the « Maison Russe ».

The building served as a coal depot for the Russian navy, which cruised the Mediterranean and had

been based in the bay of Villefranche since the 1770s. The presence of this military base and the

empress and her court attracted a Russian colony of over 150 wealthy Russian families to Nice. They

built sumptuous villas and an imposing Orthodox church. In 1860, the county of Nice and Savoie

became part of France, but the « Maison Russe », although owned by the French state, continued to be

used by the Russian navy, until it was banned from the Mediterranean in 1878 by the United Kingdom,

France and the Austrian-Hungarian empire at the Congress of Nations in Berlin.


Fig 8. The Marine Station of Villefranche sur Mer

A - The lazaret tower in which Fol and Barrois set up their first laboratory in 1880

B - Villefranche Bay was home to navies from the 18th century to the 20th century

C - Villefranche sur Mer zoological station and collecting boats in the 20th century

D, E - Institut de la mer de Villefranche (IMEV) on the port de la Darse

QR code: website (IMEV : Institut de la Mer de Villefranche) ( Anon. 2010, 2020).


The galley slave building remained at the disposal of the Russians biologists for a while even after

the Bolshevik revolution. Finally, in 1931, the marine station known as la « Station Zoologique » was

officially attached to the French Ministry of Education and placed under the authority of the director of

the Arago laboratory (the marine station of Banyuls).

The director of the Villefranche sur Mer's « Station Zoologique » was the Russian planktonologist

Grégoire Trégouboff, successor of Michael Davidoff. Trégouboff directed the institution until 1956

under the aegis of the University of Paris ( Anon. 2010, 2024, Trégouboff 1983, Dolan 2014).

Surprisingly, the Station Russe de Zoologie somehow continued to exist until 1930 with a few Russian

scientists and a modest Villefranchois staff. After the Bolshevik revolution of 1917, despite financial

difficulties, the marine station continued to exist thanks to the support of members of Russian

Academy of Sciences in exile. Support also came from members of the large Russian aristocracy and

intelligentsia fleeing the Bolshevik revolution to live in Europe and the Côte d'Azur(Trégouboff 1983).

From the naturalist painters of Nice to the visiting impressionist painters

In the 19 th century, some twenty well-known painters made a living from their art in the Nice region,

painting urban, village scenes, seascapes and landscapes, in watercolor, gouache and oil in modest

formats (Giraudy 1998). The people of Nice are familiar with some of these painters: Roassal (1781 -

1850), Fricero (1807 - 1870), Carlone (1812-1873),Trachel (1820-1872), Fossat (1822-1891), Costa

(1833-1921), Comba (1834-1872), Mossa (1844-1926), Besset (1861 -1902), since many of the city's

main streets, avenues or boulevards are named after them. In connection with our story, two painters,

Vincent Fossat and Joseph Fricero, deserve special mention. Fossat, worked with the naturalist Barla

and Vérany. He also also painted landscapes and the work of the fishermen he frequented. He made

precise paintings of fish as soon as boats arrived (Defaÿ 1998, Dolan 2022). The other painter, Joseph

Fricero was a traveler and adventurer born into a large family of winemakers and merchants in the

Nice region (Giraudy 1998). A childhood friend of Guiseppe Garibaldi, Fricero began drawing in his

youth and studied under the Roman painter Christian Borghese at the Barberi School of Drawing in

Nice. What connects Fricero to our story are his extensive travel in Italy, Sicily, Spain, Tunisia,

Turkey, Sweden, etc. and his encounters with foreigners. That led him to Saint Petersburg, where he

befriended Prince Gagarin, a member of the Russian aristocracy. Adopted as a drawing teacher by the

family of Tsar Nicholas 1 er , Fricero married Josephine, known as Yousia, the Tsar's natural daughter.

Protégés of Empress Alexandra Feodorovna, who stayed in Nice with her court in 1856 and 1858 after

the death of Nicholas 1 er , Yousia and her husband Joseph Fricero and 4 children, were close to the

Russian sovereigns. Fricero and his family settled in Nice on a vast estate where he painted Nice

tirelessly. His production slowed down in the 1860s, when Nice became French, and the wintering

population became less fond of his academic paintings. Fricero died in 1870 in relative isolation. His

paintings were rediscovered and appreciated from the 1920s onwards.

The 1860s-70s were a time of great change in painting, epitomized by the first Impressionist

exhibition at the Atelier Nadar in Paris in 1874. The first Impressionist painter on the Côte d’Azur was

Berthe Morisot who started painting in Nice and its harbor in 1882 (Lindskog & Mathieu, 2024, Anon.

2024b). She confided to her sister Edma, "I don't understand why this country doesn't serve as a great

studio for all the young landscape painters. In addition to its beauty, you enjoy a fixity in time and

good weather that allows for more conscientious research" (Giraudy 1998, Anon. 2024b).


Fig 9. Impressionist artists paint in Nice and Villefranche sur Mer


In early 1888, Claude Monet (1840-1926), inspired by the stories of Guy de Maupassant, set sail for

the Côte d'Azur, which he had visited 4 years earlier, and stayed in Antibes to paint. He attracted his

friend Auguste Renoir ( 1841-1919), who settled permanently in Cagnes in 1902. Another

impressionist from Le Havre, Eugène Boudin (1824 - 1898), came to paint the bay of Villefranche,

crowded with warships, every summer from 1892 until his death (Giraudy 1998). Other famous

painters followed - Chagall, Soutine, Modigliani, Matisse, Picasso and others - who appreciated the

beauty, light and climate of the region.

Thanks

Many thanks to John Dolan and Elisabeth Christian for encouraging me to write this article. John

generously shared his knowledge of the subject and the characters, and provided invaluable help with

the history, illustration and referencing. Gabrielle Baglione of the Muséum d'Histoire Naturelle in Le

Havre and Joëlle Defaÿ of the Muséum d'Histoire Naturelle de la Ville de Nice were wonderful hosts,

allowing me access to and reproduction of works by Lesueur et Péron, Risso, Vérany, Barla and

Fossat. I would like to thank them for commenting on and correcting my first manuscript. Catherine

Jessus also shared her knowledge of the characters and provided insightful comments.

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Vogt C. 1876 Les laboratoires de zoologie maritime. Rev. Sci. Yr 5, n°49 : 539-543.


Arts et sciences


Two centuries of arts and science in Nice and

Villefranche sur Mer: 2) Modern era: 1960 to 2024

Deux Siècles d’Arts et de Sciences à Nice et Villefranche-sur-Mer :

2) Les Modernes : 1960 à 2024

Christian Sardet 1

1

Sorbonne University, CNRS, Laboratoire de Biologie du Développement (LBDV), Institut de la Mer de Villefranche sur

Mer (IMEV), 06230, France, christian.sardet@imev-mer.fr

ABSTRACT. In a first, companion article (Sardet 2025 / 1 Pioneers: 1800 to 1900) we tell the story of the exploration of

biodiversity in the Nice region, and in particular of pelagic organisms. In this second article we examine how, more than a

century later, research in cell and developmental biology and in physiology evolved at the marine station of Villefranche

sur Mer. While research in the biology and ecology of plankton remained predominant at the site, and gradually led to the

growth of a large multidisciplinary laboratory (Laboratoire d’Océanographie de Villefranche: LOV), physiology and cell

biology were introduced in the 1960s as research teams on the physiology of fish and biology of protists joined the marine

station. In the 1980s a new research group was created by the CNRS which has grown to the present Laboratoire de

Biologie du Developpement (LBDV). We describe how imaging and molecular biology techniques were used to

understand fertilization and development in sea urchins, tunicates, cnidarians, and many other marine organisms

previously studied by the founders and visitors of the marine station in the 19 th century. We also detail the development of

new model organisms – the ascidian Phallusia, the appendicularian Oikopleura and the hydrozoan medusa Clytia. Finally

we discuss the promotion of scientific discoveries via aesthetic photographs, drawings, exhibits and web sites.

RÉSUMÉ. Dans un article complémentaire (Sardet 2024/ 1 Les anciens : de 1800 à 1900), nous avons relaté l'histoire de

l'exploration de la faune de la région niçoise, et en particulier des organismes pélagiques. Dans cet article, nous

examinons comment, plus d'un siècle plus tard, la recherche scientifique en biologie et physiologie cellulaire et

moléculaire du développement a évoluée à la station marine de Villefranche sur Mer. Alors que la biologie et l'écologie du

plancton sont prédominants sur le site et ont progressivement conduit à la croissance d’un grand laboratoire

d'Océanographie de Villefranche (LOV), à partir des années 1960 de nouvelles équipes de recherche sur la physiologie

des poissons et des protistes ont été accueillies. Et dans les années 1980, une équipe de recherche créée par le CNRS a

évoluée graduellement en l'actuel Laboratoire de Biologie du Développement (LBDV). Nous décrivons comment les

techniques d'imagerie et de biologie cellulaire moléculaire ont permis d’analyser l’ovogénèse, la fécondation et le

développement chez les oursins, tuniciers, cténophores, cnidaires et d'autres organismes marins dont certains étaient

déjà étudiés par les fondateurs et les visiteurs de la station marine au 19 ème siècle. Nous soulignons que de nouveaux

modèles – l'ascidie Phallusia, l'appendiculaire Oikopleura et la méduse hydrozoaire Clytia – se sont développés sur le

site. Nous détaillons aussi les efforts des chercheurs pour promouvoir leurs découvertes par le biais de photographies, de

dessins, d’expositions et sites internet esthétiques.

KEYWORDS. Villefranche sur Mer, plankton, protists, sea urchins, ascidians, appendicularians, ctenophores, cnidarians,

chaetognats, siphonophores, tintinnids, Paracentrotus Phallusia, Oikopleura, Clytia.

MOTS-CLÉS. Villefranche sur Mer, plancton, protistes, oursins, ascidies, appendiculaires, ctenophores, cnidaires,

chaetognates, siphonophores, tintinnides, Paracentrotus Phallusia, Oikopleura, Clytia.

1. Introduction

From the 1960s-1970s, the Villefranche sur Mer marine station evolved into a multidisciplinary

research and teaching center(Anon. 2010, 2024). The zoological research initiated 150 years ago by

Fol, Barrois, Korotneff and their visitors (see companion article: Sardet 2025 / 1 Pioneers: 1800 to

1900) expanded under the aegis of the Université Pierre & Marie Curie (UPMC, now Sorbonne

Université) and the Centre National de la Recherche Scientifique (CNRS). Other research teams joined

the site in the 1960s, supported by the Commissariat à l'Énergie Atomique (CEA team led by Jean


Maetz on fish physiology) and the Université de Nice Sophia Antipolis (UNSA, protistology team led

by Jean Cachon). At the same time, other research and teaching activities - involving departments of

Paris University

(UPMC) - developed on the Villefranche site from the 1960s onwards. These focused on geological

and oceanography studies and the physical chemistry of the oceans. Research, teaching and the hosting

of visitors were carried out in the former galley slave building and now extend to other buildings along

the Darse harbor of Villefranche – a former rope factory as well as 2 buildings dating from recent

decades (see Anon. 2010 document on the 125 year anniversary celebrating the creation of the marine

station).

In the 1960s, under the direction of Paul Bougis, the Station Zoologique de Villefranche sur Mer

became officially independent of the marine station of Banyuls sur Mer (laboratoire Arago), consisting

of 3 laboratories associated with the university of Paris (UPMC: Zoology & Ecology of Plankton/

Marine Physical-chemistry/ Geodynamics) and the CEA and UNSA teams mentioned above. The

creation and installation of a new CNRS team (Marine Cell Biology headed by Roger Lallier &

Christian Sardet) in the early 1980s added new research topics in cellular and molecular biology of

development.

Over the last few decades, under the aegis of UPMC and CNRS, the Villefranche, Banyuls and

Roscoff marine stations gradually acquired the status of “Observatoire Océanologique”. The Station

Zoologique de Villefranche successively became the CEROV (Centre d'Études et de Recherche

Océanographiques) in 1983, then the OOV (Observatoire Océanologique de Villefranche) in 1989, and

in 2019, the IMEV (Institut de la Mer de Villefranche). During these administrative transformations

driven by UPMC and CNRS, the physiological (CEA) and geological (UPMC) research teams left the

Villefranche site to join larger laboratories (Unités Mixtes de Recherches: UMR) in Nice and Sophia

Antipolis under the aegis of UNSA (Université de Nice Sophia Antipolis).

Our aim in this article is to highlight some biological research that is in remarkable continuity with

the pioneering studies carried out more than a century earlier by naturalists from Nice and Villefranche

and some of the visitors they welcomed (see companion article: Sardet 2025 / 1 Pioneers: 1800 to

1900). This continuity is due in part to the availability of benthic organisms (echinoderms, ascidians,

etc.) and of planktonic organisms (protists, cnidarians, ctenophores, tunicates, etc.) which are easily

and rapidly collected near the marine station with boats that go out to sea every day. The organisms are

kept and/or raised in aquariums by experienced staffs that have perfected culture techniques.

We also emphasize the arts & sciences dimension of research linked to the intensive and creative

use of various imaging techniques in biology. Many researchers are keen to draw attention to their

discoveries through the aesthetics of cover photos, drawings, exhibitions, conference presentations,

websites or posts on social networks.

Jean Maetz and his passion for fish and fish physiology

We begin this overview with a tribute to the naturalist school of Nice in the 19 th century, and in

particular Antoine Risso, Jean Baptiste Barla and Vincent Fossat, who were passionate students of

local fish and invertebrates (Sardet 2025 / 1 Pioneers: 1800 to 1900). A century later, Jean Maetz

inherited that passion for fish. For the staff and visitors to the Villefranche sur Mer marine station, Jean

Maetz is the name of a research and teaching building constructed in 1983 by the Commissariat à

l'Énergie Atomique (CEA) and Sorbonne Université (UPMC at the time). The building was named in

memory of physiologist Jean Maetz, who died in 1977 at the age of 54 in a car accident in Scotland. In

1964, Maetz had moved his laboratory to Villefranche, a satellite of the CEA's Laboratoire de Biologie

in Saclay, to study with René Motais euryhaline fish such as eels and trout, which are able to adapt

from fresh water to sea water by excreting salt. As a young researcher, I was welcomed to Villefranche


in 1976 by Jean Maetz and his CEA team, to take part in research on the cells - chloride cells -

responsible for ion exchange in the gills. These cells excrete salt by amplifying the pump proteins of

their dense network of internal membranes, while modifying their junctions with neighboring cells

(Fig.1). The work on these cells, renamed the MRC (Mitochondria Rich Cells) was published in cell

biology and physiology journals. It still underpins current theories of osmoregulation in euryhaline fish

(Sardet et al. 1979, Evans et al. 2005).

Fig. 1. Jean Maetz and the physiology of osmoregulation in euryhaline fishes

Top left, Jean Maetz in his white lab coat

A – Eel: watercolor painting by Vincent Fossat (Nice Natural History Museum collection)

B – Watercolor painting of gill filaments with vessels filled with red cells (red)

and chloride cells (green) at the base (C Sardet)

C – (1a) Thin section electron microscopy of gill filaments (R) showing chloride cells (C)

(1b) Scanning electron microscopy of gill surface with crypt of chloride cells (arrow) from (Sardet et al. 1979

D – Watercolor painting of chloride cells (green) at the base of gill filaments. A complex of several adjacent

chloride cells forms in response to sea water adaptation (C Sardet)

The sudden death of Jean Maetz led to the relocation of the physiology laboratory from CEA at the

marine station in Villefranche to the University of Nice Sophia Antipolis under the direction of René

Motais. It changed my own destiny, giving me the opportunity to research fertilization at the marine

station by creating, with colleagues from Villefranche, Paris and Nice, a new research team among

those encouraged by the CNRS life sciences department in the early 1980s, when François Mitterrand

had become the new French president.

Hermann Fol's legacy - from fertilization to protists

Hermann Fol, the founder with Barrois of a first laboratory in Villefranche sur Mer in 1881, was an

extraordinary personality who disappeared at sea in 1892. His genius and uncompromising, difficult


character shine through in his publications and abundant correspondence with his 2 main mentors,

Henri de Lacaze-Duthiers and Carl Vogt (Jessus & Laudet 2022, Dolan 2024, Sardet 2025).

Fol is without doubt the biologist who left in Villefranche the most interesting legacy from a

scientific point of view, if we consider the scope of his discoveries and interests. Over a period of

twenty years (1869-1889), Fol published pioneering research on protists (rhizaria, tintinnaria),

appendicularians, echinoderms, chaetognaths, planktonic molluscs and human embryos (Bedot 1894,

Dolan 2024). When he transferred his personal laboratory by boat from Messina to Villefranche in

1878, Fol had just published his discovery of fertilization in echinoderms and chaetognaths (Fol 1878),

and his research on the development of planktonic molluscs (Fol 1875). Previously, he had described

and remarkably illustrated his work on the anatomy and development of ctenophores (his thesis) and

appendicularians (Fol 1972, Dolan 2024). In fact, all the organisms mentioned have subsequently been

the subject of in-depth research at the Villefranche marine station. I myself worked on the fertilization

and development of ctenophores, echinoderms and chaetognaths, and realized that these subjects had

been pioneered a century earlier by Hermann Fol! What was striking about Fol was his ability to

discover, describe and illustrate new phenomena accurately and aesthetically. Almost always

publishing alone in various magazines, he was also very particular about the engravers he chose

himself (Dolan 2024).

Sticholonche zanclea - a rowing protist

When examining the contents of a plankton net towed at depth in the Bay of Villefranche, the eye is

inevitably drawn to Sticholonche zanclea, an atypical rhizarian measuring 0.2 mm that moves with the

aid of oar-like extensions called axopods. Fol, and then Alexis Korotneff, published their research on

this protist in the 1880-90s (Fol 1883, Korotneff 1891). A century later, Jean and Monique Cachon

turned their attention to Sticholonche, spurred on by Lewis Tilney, professor at the University of

Pennsylvania, a visitor to Villefranche on sabbatical. Tilney was famous for his contributions to the

understanding of the cytoskeleton, the support network and dynamic musculature of cell motility. Jean

and Monique Cachon had introduced microcinematography and electron microscopy techniques to

Villefranche in the 1960s. They were renowned for their work on the structure of heliozoans,

acantharia and other protists, carried out with Jean and Colette Febvre, members of the Nice university

protistology laboratory hosted in Villefranche. In the case of Sticholonche, the Cachon couple and

Tilney wondered what mechanisms enabled the rows of oars - axopods made up of microtubule

bundles - to move. They showed that these oars/axopods were anchored at the base by ball-and-socket

joints that pivoted in depressions in the nuclear membrane, reminiscent of a hip joint (Fig.2). They

observed contractile filaments that appeared to be involved in calcium-controlled movement. This

work was published in the Journal of Cell Biology, the best journal in the field at the time (Cachon et

al. 1977). As a young researcher newly arrived in Villefranche, I have fond memories of discovering

protists thanks to Jean and Monique Cachon and Jean and Colette Febvre (Febvre-Chevallier & Febvre

1994). Jean Cachon sorted and prepared the specimens, Jean and Monique examined them together

under the microscope, and Monique meticulously drew the pictures. And I learned a lot from Lewis

Tilney, a master at asking the right question to the appropriate organism.

Taking advantage of the European network of marine stations hosting facilities (see EMBRC: Anon.

2024c), a new generation of cell and molecular biology researchers work at the marine station of

Villefranche to collect and study radiolarians and rhizaria, and particularly their genes and symbiotic

relationships with microalgae (Decelle et al. 2012).


Fig. 2. Monique & Jean Cachon and the heliozoan Sticholonche zanclea

A - Monique & Jean with their Japanese microscopist friend Hidemi Sato

B - Scanning electron microscope photos of siliceous skeletons of polycystine radiolarians (M & J Cachon:

page 84 of Plankton-Wonders of the Drifting World, Sardet 201, Ulmer

C - Sticholonche zanclea: drawing by Hermann Fol (Fol 1883)

D - Drawing by Monique Cachon showing a cross-section through Sticholonche.

E & F - Electron microscope thin section of the base of an oar/axopod anchored

in a depression in the nuclear membrane of Sticholonche

(E) and corresponding drawing by Monique Cachon (F), see (Cachon et al. 1977).

QR Code in the left-hand corner. By photographing it with your phone, you can watch Sticholonche rowing

(film made by a Japanese colleague).

Tintinnids - decorated ciliates

Tintinnid ciliates are barely visible to the naked eye, but they are among the most interesting microorganisms

in plankton, as they are constantly on the move and have a shell decorated with other

protists - called a lorica - in which the ciliated cell contracts or stretches. Protruding out of the lorica,

the ciliate captures and feeds on the smallest algae in the plankton. Tintinnids, of which over 500

species have been described, are thus part of a functional group called microzooplankton (Dolan et al.

2012, Dolan 2013). Hermann Fol studied tintinnid ciliates of Villefranche in 1879 and 1880 after

Haeckel had made earlier less precise descriptions (Fig. 3, Fol 1881, Dolan 2024).

Tintinnids are frequently and easily collected with a fine mesh net in the bay of Villefranche, which

probably explains why Fol studied these ciliates. He was also the first to attempt to determine the

chemical nature of tintinnid loricas. A century later several researchers worked on tintinnids in

Villefranche, starting in the late 1950s with the Argentinian Ernesto Balech, who after a stay of several

months in Villefranche published a landmark taxonomic monograph (Dolan 2017). In the 1960s and

1970s in Villefranche, Michelle Laval-Peuto and Fereidoun Rassoulzadegan became interested in the

cytology and ecology of these ciliates (Rassoulzadegan et al. 1988). More recently, John Dolan

published articles on the diversity of tintinnid species assemblages in the bay of Villefranche and in

deep waters offshore (Dolan 2012, 2013). Nonetheless, many aspects of tintinnid biology and ecology


remain to be elucidated and Villefranche is a perfect location for researchers interested in the

taxonomy, ecology, parasitism and symbioses of tintinnids (Vincent et al. 2018).

Fig. 3. Tintinnid ciliates in the bay of Villefranche sur mer

A - Drawings by Ernst Haeckel (see Dolan 2024)

B - Drawings by Hermann Fol (Fol 1881, see Dolan 2024)

C - Cover of the book on tintinnid ciliates edited by John Dolan (Dolan et al. 2012)

D - The tintinnid Codonellopsis (photo J. Dolan) whose lorica is decorated with calcareous scales of

coccolithophores. Scanning microscope by I Machour & C Bachy, CNRS photo library (Plankton – Wonders of

the Drifting World, Sardet 2015, Univ. of Chicago Press).

The larvacean Oikopleura dioica - a research model developed in Villefranche

Fol had just published his observations of larvaceans (also called appendicularians) done in

Messina when he first stayed in Nice for a few months (Fol 1872). In a letter to his future wife Emma

Bourrit, he describes these tadpole-like urochordates: "I went out yesterday morning. The sea was

calm, the weather splendid and a small boat was docked near the beach. I couldn't resist the

temptation. I grabbed a jar and headed out to sea. After a quarter of an hour, my jar was full and I

returned to examine it at my leisure. It contained about twenty larvaceans, microscopic long-tailed

creatures that swam in all directions, as transparent as crystal and as agile as little fish. But here's one

that stops. It's in the process of forming an envelope serving as a net, and soon it's on the move again

in its crystalline envelope 20 times its size. It's well protected now. Let's capture it in a glass tube and

put it under the microscope. Just as it feels it's been caught, a flick of the tail and the larvacean is off,

leaving its empty envelope as its only booty".

Fol's pioneering work on appendicularians was taken up again at Villefranche in the 1950s by

Robert Fenaux and his successors, including Gabriel Gorsky who, with Fabien Lombard, succeeded in

mastering the cultivation of the species Oikopleura dioica to analyze their reproduction and physiology

(Fig.4, Fenaux 1963, Gorsky et al. 1987, Lombard et al. 2009). In the late 1990s, the technical

expertise developed at Villefranche was transferred to the Michael Sars Center in Bergen, enabling

Daniel Chourrout, Eric Thompson and their collaborators to establish Oikopleura dioica as the

reference experimental model for appendicularians (Seo et al. 2001, Marti-Solans et al. 2015)). Since

then, half a dozen laboratories in Europe, Japan and the United States have adopted O. dioica and

helped elucidate and manipulate its genome - the smallest known genome in the chordates (Nishida

2008). The appendicularian model is very attractive. The adult animal consists of fewer than 5,000

cells, with perfectly established lineage and differentiation into a half dozen tissue types. The larvacean

O. dioica can reproduce in just a few days, and using its “house”/net and a sucking technique it feeds

on nearby bacteria, micro-algae and particles. The appendicularian secretes and deploys several


“houses”/nets (called logettes) per day thanks to highly polyploid epithelial cells that secrete the net’s

fibers in a precise pattern (see fig 4 and Thompson et al. 2001).

Fig. 4. The larvaceans (also called appendicularians)

A – Drawings of the anterior part of the appendicularian Oikopleura dioica (Fol 1872)

B – Domains of polyploid cells secreting the “house”/net (Thompson et al. 2001)

C – Appendicularian “house”/net (Fenaux 1986)

D – Book on the ecological impact of appendicularians (Gorsky et al. 2005)

QR Code: use your mobile phone to watch a film about larvaceans

Due to their abundance and proliferation in all oceans, and their ability to filter particles and

microorganisms, appendicularians and their “houses”/nets play an important role in the constitution of

marine snow particles, contributing to carbon sequestration as they sink to the abyss. This is the

direction that recent research has been taking in Villefranche (Fig. 4, Gorsky et al. 1984, Guidi et al.

2009).

Fertilization and development - a new team

In the 1980s, a CNRS research team (ER250 Biologie Cellulaire Marine), of which I was one of the

instigators, was established at the marine station, reviving early research on the development of marine

organisms in Villefranche by Barrois, Fol, Metchnikoff, Kowalski and others a century earlier (see

companion article: Sardet 2025 / 1 Pioneers: 1800 to 1900). In fact, this research never ceased in

Villefranche, as local zoologists continued to explore the characteristics and development of marine

organisms - salps, appendicularians, pteropods, cnidarians, fish, etc. They feature prominently in the

reference book on Mediterranean plankton (Manuel de Planctonologie Méditerranéenne from

Trégouboff & Rose 1957).

The ER 250 research team was first headed by CNRS embryologist Roger Lallier, who had been at

the Villefranche site since the 1960s doing research on the development of the sea urchin


Paracentrotus lividus (Lallier 1975). The oocytes and embryos of this species are characterized by a

sub equatorial pigment band, making it possible to study the expression of vegetative/animal polarity

during development. Lallier modified the development of sea urchin embryos by using a variety of

chemical compounds that either animalized the embryos (enriching them with ectodermal tissue) or

vegetalized them (enriching them with endodermal and mesodermal tissue). In addition to resident

investigators (Roger Lallier, Danièle Carré, Christian Sardet), the new group, soon attracted CNRS

researchers from Paris (Jacky and Marie Paule Cosson), Nice (Christian Gache) and the first PhD

students (Richard Christen, Thierry Lepage). Moving from the CEA to the CNRS, I succeeded Roger

Lallier as head of the CNRS research unit in 1985, and we welcomed other CNRS and INSERM

researchers, as well as post-docs and students working on fertilization, development and motility. We

also incorporated Jean Cachon's protistology team into a new Unité Mixte de Recherche (UMR 671

Biologie Cellulaire Marine) with around thirty members. In 2000, the laboratory became UMR 7009

Biologie du Développement under the direction of Christian Gache (2000-2008). It was later headed by

Evelyn Houliston (2009 - 2018) and Alex Mc Dougall (2019 - present), two CNRS researchers who

had first joined our laboratory as post-docs.

Fertilization and cell activation – the importance of ionic signals

Exchanges with Roger Lallier on sea urchin fertilization and development, and the discovery at that

time of calcium signals triggered by the fertilizing spermatozoa in fish and sea urchins (see Sardet

2023) prompted us to study ion fluxes in embryos with professors and researchers at the university of

Nice, who had been my colleagues at the Villefranche CEA laboratory (Christen et al. 1979, Girard et

al. 1982, Sardet et al. 1984). These research subjects, new to me, corresponded to the birth of my first

son Noé and the meeting of visiting biologists interested in developmental biology with whom I

worked in Villefranche (Marko Zalokar and Lewis Tilney) and in the USA (David Epel, Dan Mazia as

well as Lionel Jaffe, Mark Terasaki and Shinya Inoue, pioneers of calcium signals and the microscopic

imaging revolution).

In the 1990s, our Villefranche laboratory became one of the best equipped in France for light and

electron microscopy sample preparations and observations largely thanks to the efforts of two

inventive CNRS engineers/researchers - Christian Rouvière and Patrick Chang (Sardet & Chang 1985,

Rouvière et al. 1994, Sardet et al. 1998). Our research group developed a new experimental model

inherited from Marco Zalokar – the ascidian Pallusia mammillata – a solitary ascidian species whose

oocytes and embryos are more abundant and far more transparent than those of the reference species –

Ciona intestinalis (Zalokar & Sardet 1984). This allowed us to analyze in detail calcium signals at

fertilization and the reorganization of the cortex and cytoplasm in fertilized oocytes and the

consequences for embryonic development (Fig. 5). It was already known that in mouse oocytes,

fertilization triggered oscillations in intracellular calcium concentration lasting for hours. Thanks to

experiments on several species of ascidians initiated with our colleagues at Woods Hole, and pursued

with Alex McDougall and Rémi Dumollard at Villefranche, we were able to show that the periodic

bursts of calcium were in fact propagating calcium waves emitted by a localized "pacemaker". Alex

McDougall demonstrated for the first time that these periodic calcium waves were strictly necessary to

complete the meiotic cell cycle (Speksnijder et al. 1990, McDougall & Sardet 1995, Dumollard et al.

2002).

Research was pursued on Phallusia with Janet Chenevert, Philippe Dru, Fabrice Roegiers, François

Prodon, Alexandre Paix and Nang Le Nguyen. We focused on cortical structures and determinant

macromolecules during early development, and in particular on cortical determinant RNAs, which play

a key role in embryonic polarity (Prodon et al. 2005, Paix et al. 2011). During this latter period, we

benefited from intense exchanges with our Japanese colleagues Hiroki Nishida, Lixy Yamada and

Kazuo Inaba (Prodon et al. 2010). This research continues in Villefranche and other laboratories in

Europe have now adopted this experimental model.


Fig. 5. Calcium signals in ascidians oocytes following fertilization

A & B – Alexandre Lesueur and Ciona intestinalis he drew in Nice in 1804

C – The ascidian Phallusia mammillata photographed by Christian Rouvière

D – Calcium waves (yellow to red) repeatedly crossing ae fertilized oocyte. In the center, a schematized

spermatozoon (graphic by Mohamed Khamla)

E – Left: calcium signals triggered by fertilization (F) and during the first meiotic division (MI Phase).

Right: calcium oscillations during the second meiotic division (MII Phase).

F – Cover inspired by one of our publications on calcium signals (Dumollard et al. 2002)

Asking the right question to the right organism - chaetognaths and ctenophores

Hermann Fol's ghost must have suggested that we take an interest in ctenophores – Fol had

written his thesis on their development – and also in chaetognaths, which Fol used to extend his

observations on fertilization (Fig. 6, Fol 1979, see Sardet 2025 / 1 Pioneers: 1800 to 1900). Inspired by

early work and the expertise of our zoologist colleague Danielle Carré, we began asking these

planktonic animals two pertinent questions in the 1980s.

Chaetognats

With regard to the hermaphroditic chaetognaths(arrowworms) Sagitta and Spadella, whose

reproductive cycles and tissue morphology had been figured out by Danielle Carré, we asked how an

exceptionally large germinal granule was formed and how it enabled the cells that inherited it to

become the germ cells at the origin of male and female gonads. It was in fact in chaetognaths that these

germline-determining granules were discovered in the 1900s (Wilson 1925). We were able to observe

how this very large germinal granule (it measures twenty microns), forms at the moment of mitosis at

one pole of the fertilized egg. This granule and its descendants remain visible throughout the embryo's

divisions (Fig. 6, Carré et al. 2002). This makes it possible to analyze and manipulate germ cell

development.

We now know from genetic and biochemical analyses in the fly Drosophila and the nematode worm

Caenorhabditis that these granules contain aggregates of RNA molecules and associated proteins in the

form of biomolecular condensates (Sardet 2023). These germ granules condensates direct the

differentiation of the cells that inherit them into germ cells. An algerian colleague, Chakib Djediat ,

successfully focused on this project (Carré et al. 2002), which deserve to be continued in the light of

recent advances about the differentiation of germ cells and the discovery of biomolecular condensates.


Fig. 6. Chaetognaths

A - Drawing by Hermann Fol of the meeting of the male and female nuclei in the fertilized egg of the Sagitta

chaetognath (Fol 1978)

B - Illustration taken from our publication on chaetognaths (Carré et al. 2002).

C - Development cycle of chaetognaths and their germ cells (in green) from Carré et al. 2002.

Ctenophores

We also explored basic embryological questions using the oocytes and embryos of the ctenophore

(comb jelly) Beroe ovata. In ctenophores, fertilization plays a major role in the acquisition of the

unique oral-aboral axis in embryos and adults. As its name suggests, Beroe ovata is characterized by

oocytes with a large diameter (1.5 mm) much larger than those of other ctenophores. These oocytes

and embryos are particularly well suited for microscopic imaging due to their extraordinary

transparency and the fact that all events - from fertilization to the successive divisions of the embryo -

take place within a thin cortical layer some ten microns thick beneath the surface.

Every spring during the 1990s, we - Danielle Carré, Evelyn Houliston, visiting collaborators,

fishermen and myself - would watch for the arrival of these iridescent animals in the bay. Early in the

morning, we collected animals aboard boats or a pneumatic zodiac in the Bay of Villefranche, a sort of

treasure hunt. Sometimes we'd bring dozens of animals back to the laboratory, sometimes we'd return

empty-handed, but happy with these mornings spent scanning the surface of the meandering currents.

In any case, thanks to imaging techniques coupled with frame-by-frame video recording (at first, we

were using bank surveillance equipment!), we observed that the female nucleus in the oocyte would

often travel to explore several male nuclei introduced by different spermatozoa that had entered the

oocyte as fertilization in Beroe is polyspermic (Carré & Sardet 1984, Rouvière et al. 1984). The female

nucleus would then fuse with the chosen male nucleus to participate to the first mitotic division

defining the site of the first unipolar division, thus determining Beroe's unique oral-aboral embryonic

axis (Fig. 7). When we first published them, these observations of the behavior of a cell nucleus

stimulated the imagination of many biologists calling for further experimentations (Carré et al. 1991).

We also demonstrated the potential of this experimental model for understanding the role of cell cycle

factors in the establishment of the embryonic axis (Houliston et al. 1993). Unfortunately, to our

knowledge, work on the biology and development of this species seems to have ceased for the

moment.


Fig. 7. The ctenophore Beroe ovata

A - Three images from a film of the fusion of the spermatozoon with the oocyte

B - Three images from a film showing the exceptional clarity of first mitosis

C - Cover illustration inspired by one of our publication (Houliston et al. 1993)

D -Drawing of a fertilized oocyte in which the trajectories of the female nucleus to successively explore several

male nuclei were filmed for 1 hour (Carré & Sardet 1984)

QR Code: use your phone to watch a film about ctenophores

Exploring cnidarians - a century after Vogt and Metchnikoff

In 1853, Carl Vogt – known as the revolutionary scientist – (Fig.8) and in 1886, Elie

Metchnikoff,the co-discoverer of immunity, collected and made pioneering observations on cnidarians

(siphonophores, jellyfish, corals and anemones) in Villefranche. Cnidarians are ancestral animals

sharing the common characteristic of possessing and using stinging cells - called cnidocytes - to

paralyze prey. Carl Vogt described and beautifully illustrated the siphonophores collected in the bay in

his monograph "Recherches sur les animaux inférieurs de la Méditerranée" (Vogt 1854, Sardet 2025).

Siphonophores live as colonies of organs – floating, propulsive, reproductive, and digestive - arranged

along a filament called a stolon. Some of the 175 known species extend their colonies over tens of

meters fishing small organism including fish with stinging filaments. For this reason siphonophores are

the longest animals in the world.

In the 1980s, our zoologist colleague Danielle Carré showed us that the oocytes of siphonophores

released by reproductive colonies - the gonophores – remarkably attract sperm in a species-specific

way. Each oocyte is capped by a hemispherical cupule attracting spermatozoa, at one pole of the

oocyte corresponding to the fertilization site (Fig. 8). By dissecting and solubilizing cupules, we were

able to show that the attraction was due to a molecule which, in an electrophoretic micro-gel, attracted

sperm of the right species in the form of a band (Carré & Sardet 1981 and drawing F in Fig. 8).


Fig. 8. Siphonophores

A – Plate 19 in "Recherches sur les animaux inférieurs de la Méditerranée" (Vogt 1854)

B - Carl Vogt at the age (40) when he made his observations in Villefranche sur Mer

C - A calycophore siphonophore of the genus Chelophiès (page 116, Sardet 2013)

D- Eudoxia (reproductive medusa) of a siphonophore carrying male and female gametes

(page 121, Sardet 2013)

E - A siphonophore oocyte attracting spermatozoa of the same species to one pole.

F - Drawings illustrating the attraction of spermatozoa (red) to the cupule (yellow).

QR Code: use your phone to watch a film about siphonophores

However, the analytical methods used at the time did not make it possible to identify the molecular

nature of the attractant, which would probably be possible now.

Our colleagues Marie-Paule and Jacky Cosson went on to show that attraction was in fact linked to a

change in sperm behavior (Cosson et al. 1983). In the vicinity of the cupule emitting the attractant

molecules, the spermatozoa switched from a rectilinear swimming trajectory to swimming in small

circles, keeping them close to the attracting cupule capping the oocyte fertilization site. This swimming

change is mediated by the concentration of calcium ions (Cosson et al. 1984).

Further research into the development of siphonophores is currently underway at Villefranche

(Mańko et al. 2023). Incidentally, our original publications on sperm attraction (Carré & Sardet 1981,

Cosson et al. 1983) are unfortunately not listed in the PubMed bibliographic databases.

A new experimental model - the micro medusa Clytia hemisphaerica

We have already described experimental models - the ascidian Phallusia, the appendicular

Oikopleura - developed at the Villefranche marine station and how these models have been adopted by

other laboratories(see previous chapters). The most recent and remarkable example of the development

of an experimental model concerns a small medusa - Clytia hemisphaerica. This hydrozoan jellyfish is

the subject of a large-scale project initiated in the mid-2000s by Evelyn Houliston with Tsuyoshi

Momose (Momose & Houliston 2007).


Since then, some twenty research colleagues, teachers, post-docs, students and visitors have been

involved in elucidating the cellular and molecular mechanisms at work in oogenesis and in spawning,

in the establishment of embryonic axes, in tissue differentiation, regeneration and the ecology of this

jellyfish. This research is described in some thirty publications (see reviews by Houliston et al. 2010,

and Houliston et al. 2022).

Fig. 9. Clytia hemisphaerica an attractive jellyfish experimental model

A - Drawing of the polyp and jellyfish life cycle (Houliston 2022)

B -Cover photo of Clytia in Trends in Genetics (Houliston & Momose 2010)

C - Cover photo showing a section of a Clytia embryo section using fluorescence microscopy: the cell nuclei

are blue, the cilia are green (Momose & Houliston 2007)

QR Code: use your phone to watch the film Clytia, a laboratory favorite

The judicious choice of Clytia as a model was originally based on observations made in

Villefranche by Danielle and Claude Carré (Carré & Carré 2000). They showed that it was possible to

cultivate this jellyfish measuring a few millimeters in the form of colonies of "immortal" polyps that

propagate vegetatively, providing offspring by constant reproduction via budding. Since then, cultures

and techniques, including those for imaging, visualization, gene manipulation and transgenesis, have

been optimized and published (Lechable et al 2020, Weissbourd et al. 2021, Houliston 2022). As a

result, Clytia has become the reference experimental model for hydrozoans adopted by other

laboratories around the world. Although the genomes of some model cnidarians - the anthozoan

Nematostella and the hydrozoan Hydra - are known, the sequencing of the genome of the hydrozoan

Clytia, spearheaded by Lucas Leclère and Richard Copley (Leclère et al. 2019), allows investigations

into the mechanisms at work during the complete life cycle of cnidarians in all their complexity. As a

typical hydrozoan, Clytia lives and reproduces as fixed budding polyps and at the same time as

swimming male and female jellyfish. Yet, all these very different life forms share the same genome.

Remarkably, the Clytia model lends itself equally well to work in neuroscience and ecology (Vogt

2022, Houliston et al. 2022), and to model gastrulation (Kraus et al. 2020). This brings us back to the

origins, when in 1886, the Russian zoologist Elie Metchnikoff came as a visitor to the Villefranche

marine station and described the formation of the gastrula in Phialidium, since renamed Clytia

(Metschnikoff 1886).

Genes in action – the development of sea urchins and ascidians

Since the 1970s, the development of animals and plants has been analyzed in terms of the

expression in time and space of key gene networks, some of which are universally shared. Specific

genes are transcribed and expressed in the form of proteins, in different regions the oocyte (so-called

maternal genes) and/or embryo (so-called zygotic genes) at crucial moments after fertilization, the first


cell divisions, gastrulation or metamorphosis. Choosing to work on the Paracentrotus lividus sea

urchin model already worked on at Villefranche by Roger Lallier, our colleague Christian Gache left

the Biochemistry Department at the University of Nice to bring his knowledge of molecular biology

and gene expression to our Villefranche laboratory from the outset.

Fig. 10. Development of the sea urchin Paracentrotus lividus

A - Cover photo showing a normal sea urchin embryo at the pluteus larva stage (left) and an animalized

embryo (right) (Croce et al. 2006)

B - Life cycle of the sea urchin Paracentrotus lividus (Formery et al. 2021)

C - Confocal microscopy of a young adult(juvenile)Paracentrotus lividus. Muscles are in blue, the nervous

system in yellow, and the nuclei in white (Formery et al. 2021).

QR Code: use your phone to watch a film about sea urchins

The research initiated by Christian Gache has been continued by Thierry Lepage, Jenifer Croce,

Christian Ghiglione, Guy Lhomond, and many other researchers, technical staff, students and visiting

researchers and professors such as David McClay (Duke Univ. USA), right up to the present day.

Gache and Lepage first isolated the enzyme that enables sea urchin larvae to hatch (break their

envelope) and showed that its gene is expressed in a polarized fashion (Lepage & Gache 1989, Lepage

et al. 1992). Then Gache, his team and their successors - the teams of Thierry Lepage and then Jenifer

Croce and their collaborators - explored the roles of key genes in the differentiation of embryonic

tissues (ectoderm, mesoderm, endoderm). They also investigated how the pluteus larva acquires its

skeleton made of calcium carbonate semi-crystals (Croce et al. 2006, Robert et al. 2014). Collectively,

these research teams provided the research community with remarkable genomic (Lepage et al. 2004,

Marletaz et al. 2023) and morphological (Formery et al. 2021) observations as well as essential tools.

Solitary and colonial ascidians, tunicates known as sea squirts (Ciona intestinalis, Phallusia

mammillata, Botryllus schlosseri) have also been used extensively in Villefranche sur Mer to

understand gene expression. Having joined the laboratory some twenty years ago, Clare Hudson and

Hitoyoshi Yasuo and their research team defined some of the rules of tissue differentiation in embryos

of Ciona intestinalis, the most widely used experimental model for ascidians. In a single day, ascidian

embryos develop into a motile tadpole consisting of only 6 tissues made up of less than 3,000 cells,

with all cell lineages perfectly known. The experimental strategy employed by Hudson and Yasuo

combines micromanipulation and ablation of some specific cells in embryo at early stages, followed by

analysis and modeling of the expression of essential genes and proteins using imaging. These

approaches enabled them to understand how a small number of cells involved early on (at the16-64 cell

stages) differentiate in just a few hours to form the muscle and nerve tissues of the tadpole. They

compared gene expression in this simpler ascidian model with the mechanisms at work in the more

complex vertebrate embryos (Yasuo et al. 2007, Hudson et al. 2011, 2021).


Fig. 11. The development of the ascidian Ciona intestinalis

A - Cover photo showing that in a normal embryo (top), 10 neural chord precursor cells express the Brachyury

gene. Its expression is altered in embryos (middle and bottom) following micromanipulations of cells

(Hudson & Yasuo 2006)

B - Drawing showing which cells in the early embryo specify different parts of the ascidian tadpole's nervous

system (Hudson et al. 2011)

C - Cover photo showing different motor neurons expressing fluorescent genes (in green), motor neurons

injected with a lipophilic molecule (in magenta) and cells in which nuclei are revealed using a DNA fluorescent

dye (in white). From Hudson et al. 2011.

The other experimental model developed at Villefranche - Phallusia mammillata - is being used by

Alex McDougall, Rémi Dumollard, Janet Chenevert and their collaborators to analyze how the

orientation of cell divisions and the de-synchronizations of cell cycles determine the positioning and

differentiation of cells (stages 16 to 128 cells) whose fates are rapidly fixed (Dumollard et al. 2013,

McDougall et al 2019, Chenevert et al. 2020).

The Villefranche sur Mer development biology laboratory, considered to be one of the world's

leading laboratories in the field of comparative development biology has shared widely its expertise

and methodologies and organized several meetings of the cell and developmental biology research

community in Villefranche (Sardet et al. 2008, 2011, Yasuo & McDougall 2018, Dumollard et al.

2017).

Finally, to close this chapter, I'm sorry I do not have the space to do justice to our colleagues from

Villefranche sur Mer who are successfully pursuing comparative molecular and cellular approaches on

a variety of other marine organisms (Amphioxus, Botryllus, Clytia, Mytilus, Salpa, etc.).

Tara oceans – a human and scientific adventure

Villefranche has been world-famous for its plankton ever since pioneering naturalists from Nice,

Germany and Switzerland revealed the biodiversity of pelagic organisms that drift and dwell in the bay

ever changing with the seasons and the weather conditions.

Following in the footsteps of the pioneers, Grégoire Trégouboff and the Villefranche zoologists and

their successors at the LOV laboratory popularized the history of plankton biodiversity and ecology

(Trégouboff & Rose 1957, Trégouboff 1983, Anon. 2024a).

In 2008, with Eric Karsenti and a few colleagues I came up with the idea of a global expedition to

study plankton in all oceans on board the schooner Tara, the Villefranche marine station emerged as an

essential partner in the project (Karsenti & Di Meo 2012). On a table corner in the Villefranchge

harbor – le port de la Darse - Eric Karsenti, Gaby Gorsky and I began to sketch out the broad outlines

of the Tara oceans expedition a global exploration of plankton in all oceans. We convinced colleagues


at marine stations - Villefranche, Roscoff, Banyuls, Naples - and at a dozen other research laboratories

in Europe and USA to work together and share their indispensable oceanographic and biological

expertise. The schooner Tara came in the bay of Villefranche for an initial planning conference, and

after a year of planning, the expedition left Lorient for the Mediterranean in September 2009 (see our

films of the expedition's departure: Anon. 2019). Gaby Gorsky, Marc Picheral, Lars Stemann and

colleagues from Villefranche and colleagues from the marine stations of Roscoff, Banyuls and Naples

began equipping and adapting the schooner and guiding its course for a global exploration of plankton.

Eric Karsenti (EMBL, Heidelberg) and Etienne Bourgois (agnèsb) led the first expedition with some

twenty scientific coordinators as part of a consortium of half a dozen major institutions (CNRS, CEA,

EMBL, Sorbonne University, etc.).

Aware that the main strategy of the expedition - the analysis of the planktonic ecosystem using

genomics - would not be an effective way to popularize plankton, we decided to tell the stories of

organisms in plankton via multimedia documents of the plankton chronicles project (Sardet 2017). I'm

very grateful to fellow zoologists from Villefranche, and in particular Claude Carré, who taught me

and told me many stories about planktonic creatures. Véronique Kleiner (CNRS Images), Noé Sardet

and Sharif Mirshak (Parafilms, Montreal) made it possible to photograph and to produce short films

and the well used Plankton Chronicles multilingual website on the beauty and diversity of planktonic

creatures. A book on plankton for a wider audience was published in French (Sardet 2013), and

translated in English (Plankton - Wonders of the Drifting World (Univ. Chicago Press 2015), Japanese

(2014), Germany(2018) and Chinese (2019).

Fig. 12. Plankton – an arts and science perspective

A - Cover of "Manuel de Planctonologie Méditerranéenne " (Trégouboff & Rose 1957)

B – A page from Volume 1 of the Manuel with illustrations of Sticholonche and Zoothamnium

(Chapter XV, plate 55)

C - Cover of the May 22, 2015 issue of Science magazine announcing the 5 articles detailing the first scientific

results of the Tara oceans expedition (photos of planktonic organisms from the book "Plankton - aux origines

du vivant" C. Sardet, Ulmer 2013).

D - The video episodes on the "Chroniques du plancton" website in French, English and Spanish have been

visited by 200 to 1,000 people a day since 2013.

QR Codes: use your phone to view Chroniques du plancton (films, news, etc.) and the different expeditions the

Tara oceans expedition


The first Tara oceans expedition ran from 2009 to 2012 (Anon. 2019), then the expedition set off

again under the direction of Chris Bowler (ENS, Paris) around the Arctic in 2014 and has continued in

various forms ever since (Anon. 2024b). Over 150 publications, most of them in major international

journals, attest to the impact and success, of the expedition and its scientific discoveries which we will

not detail here (Anon. 2024b).

The project was atypical in the sense that an ad-hoc group of biologists, geneticists,

bioinformaticians and oceanographers from various institutions (CNRS, EMBL, CEA Genoscope, etc.)

joined forces with a private organization (Fondation Tara Océans / agnèsb, owner and manager of the

schooner Tara) with a simple idea in mind: to collect and analyze the entire planktonic ecosystem

together with many physical-chemical parameters in hundreds of well-chosen sites worldwide. This

ambitious project relied on massive gene analysis by the Genoscope (metagenomics) and the use of

interpretation tools (AI, correlation analyses etc.). Automated image acquisition and recognition was

also key to the expedition success, a field in which the Villefranche marine station played a leading

role thanks to the involvement of Gaby Gorsky, Fabien Lombard, Lionel Guidi, Marc Picheral and

their colleagues (Gorsky et al. 2019, Lombard et al. 2019, Picheral 2022). We set off on our expedition

in September 2009 with a great deal of enthusiasm and sympathy, but also with a lot of criticism and

skepticism. As a consequence we did not get much of grant money that might have made things easier.

It was a gamble!

From the outset, I was keen to assemble on board Tara a library of planktonic works, and above all a

photocopy and digitized version of the "Trégouboff", the plankton bible: "Manuel de Planctonologie

Méditerranéenne" published in the 1950s by Grégoire Trégouboff, then director of the Villefranche sur

Mer zoological station (Trégouboff & Rose 1957). The book is in two volumes, one volume containing

plates of organisms compiled from many zoologists' drawings, the other volume a dense text with

digressions and references. Although somewhat difficult and confusing to use, the "Trégouboff", is a

must-have reference for plankton and a source of pride for Villefranche.

In addition to its scientific aspects, the Tara oceans expedition aimed to raise awareness about the

planktonic ecosystem among the general public and young people in schools. Like the scientific

component of the expedition, the communication impact went beyond our expectations, thanks to the

generosity of the participating scientists as well as the communication and education team set up by the

Tara Oceans Foundation (Anon. 2024b). Acting as liaison between the Foundation's communications

team and the scientific team, I began photographing and filming the organisms from the outset to tell

their stories. Many other educational and artistic projects (exhibitions, applications, publications,

games) have been created and continue to be created around the Tara oceans expeditions such as the

“La grande expédition“ the art and science exhibit organized recently by the Tara Océans Foundation

at the CentQuatre in Paris (Anon. 2024b, d). This desire to share information is in the air of the times,

and scientific communication and mediation efforts in Villefranche have multiplied in recent years, in

particular through the "Ocean Culture" outreach project.

Conclusion

This overview of research over the last 60 years represents my personal view of events, and I

apologize in advance to anyone I may have overlooked or forgotten in recounting this saga. This story

is also a way of paying tribute to the work of some of the Villefranche biologist colleagues whom I

frequented and mentioned in this article and who are unfortunately no longer with us: Jean Maetz,

Lucienne Fenaux, Jean Cachon, Marie Paule Cosson, Jean Febvre, Monique Cachon, Maurice Fenaux,

Roger Lallier, René Motais, Richard Christen.

To find out more about current research into the biology and development of organisms, plankton

and oceanography, visit Institut de la Mer de Villefranche (IMEV) website, the Tara oceans website

(Anon. 2024b) and Traversing EuropeanCoastlines (TREC ) the last expedition of the Tara schooner.


Thanks

Many thanks to John Dolan and Elisabeth Christian and Marie Christiane Maurel for encouraging

me to write and translate this article. John generously shared his knowledge. I would also like to thank

Evelyn Houliston, Hitoyoshi Yasuo, Luca Leclère, Janet Chenevert, Gaby Gorsky and Jenifer Croce

for their comments.

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Arts et sciences


The Université Internationale de la Mer, Arts and

Sciences

L’Université Internationale de la Mer, les arts et les sciences

Jean-Eric Aubert 1

1

President of the Université Internationale de la Mer, jean-eric.aubert@univ-mer.org

ABSTRACT. This article discusses the importance of combining both art and science approaches for raising awareness

on ocean issues and developing a marine and maritime culture. This is illustrated by the experience of the Université

Internationale de la Mer, based on the French Côte d’Azur. The article describes different ways by which arts and sciences

help in ocean acculturation and exploration, as well as the need of combining artistic and scientific mindsets for ocean and

coastline foresight exercises. Both acculturation and foresight efforts will be key for coping with long term ocean challenges

highlighted at UNOC3.

RÉSUMÉ. Cet article souligne l’importance de combiner des approches à la fois scientifiques et artistiques pour développer

la conscience des problématiques océaniques et développer une culture marine et maritime. L’expérience de l’Université

Internationale de la Mer, basée sur la Côte d’Azur, en donne l’illustration. L’article esquisse différentes manières par

lesquelles la combinaison des arts et les sciences contribue à l’acculturation à l’océan et à son étude. Il montre également

le rôle d'une double démarche scientifique et artistique dans la prospective des mers et des littoraux. Acculturation et

prospective sont essentielles pour relever les défis posés à long terme par les transformations de l’océan, comme le met

en évidence UNOC3.

KEYWORDS. Ocean transformations, Sea level rise, Marine biodiversity change, Acculturation, Operational Foresight,

International University of the Sea.

MOTS-CLÉS. Dérèglements de l’océan, Montée du niveau des mers, Changement de la biodiversité marine, Acculturation,

Prospective opérationnelle, Université Internationale de la Mer.

The Université Internationale de la Mer : origins and activities

The Université Internationale de la Mer (UIM -- International University of the Sea) was founded in

1986 to provide foreign students coming to France with basic knowledge of marine life and maritime

activities. It was the result of a partnership between the Collége International de Cannes (CIC --

International College of Cannes) and the Centre d’Études et de Recherches de Biologie et

d’Océanographie Médicale (CERBOM -- Center for Studies and Research in Biology and Medical

Oceanography) based in Nice. The former had been initiated by Paul Valéry in the 1930s to familiarize

foreign students with the French language and literature. The latter, a unit of the Institut National pour

la Santé et la Recherche Médicale (INSERM -- National Institute for Health and Medical Research), had

been founded in 1960 by a doctor (neurosurgeon), Maurice Aubert, who wanted to combine his expertise

with his passion for the sea 1 . Thus, in its early days, the UIM raised awareness of marine and maritime

1

For a history of CERBOM and a biography of its founder, see the foreword to the book Systèmes d'information des microorganismes

marins, published by ISTE (2021)


issues among several classes of dozens of foreign students, particularly Americans. The premature death

of its founder at the Collège International de Cannes then put the project on hold.

The CERBOM, for its part, continued its work linking health and the sea: fundamental research,

particularly on the biological balance of the seas and the antibiotic properties of the marine environment;

applied research on sea pollution prevention and treatment, thalassotherapy, etc.; and various inventories

on the state of the seas surrounding France and bathing the Mediterranean coasts (hydrological,

bacteriological, and chemical aspects) 2 .

Following the closure of CERBOM in the mid-1990s, the UIM and its founder were welcomed by the

municipality of Cagnes sur mer, in the port of Cros de Cagnes, in the fishermen's guild building, which

had then been renovated. The UIM, while inheriting the scientific work of the CERBOM, then developed

as an educational structure for a wide range of audiences: welcoming students in partnership with

departments of the University of Nice-Sophia Antipolis (marine biology, marine geography), training

maritime professionals (captains and seamen, aquaculture technicians), and familiarizing local

schoolchildren with the sea.

Figure 1. View of the sea from the beach at Cros de Cagnes. In the background on the left side is the landing

track of Nice airport. The sea is blue-green due to the inflow of water from the Var River. The sea becomes

darker blue on the horizon line. Photo credit: Fabienne Goux-Baudiment

2

All of CERBOM's publications, supplemented by those of the UIM, are included in a book entitled “Un demi-siècle d'étude du

milieu marin appliquée à l'homme” (Half a century of marine research applied to humans, 1958-2008). The book (190 pages) is

organized as an Index that lists and summarizes 585 articles and papers, 10 books, and 25 inventories resulting from oceanographic

campaigns. It is available on the website of the Université internationale de la Mer. https://www.univ-mer.org


Figure 2. The UIM is located on the northern coastline of the Baie des Anges, nearby the Var river, positioned

on the upper right side in this excerpt from a Cassini map of Provence. Cassini maps —also known as

Académie maps—were created in the 18th century and covered the whole of France. They provided

remarkable detail of the coastlines and relief thanks to geodetic triangulation methods that were highly

innovative for the time. Note that the Nice coastline is not included in the map,

as the city joined France only in 1860. Photo credit: Fabienne Goux-Baudiment

Following the retirement of its founder in 2008, the UIM gradually reduced its activities to training

seamen and captains only. It was relaunched in 2018 at the request of the municipality, consolidating

these maritime training courses and developing distance learning courses on “blue economy” for Frenchspeaking

Africa, in partnership with Senghor University, an operator of the Francophonie, as well as

organizing various activities to promote ocean awareness: conferences, presentations in middle schools,

seminars for elected officials, etc. 3

On the occasion of the third United Nations Ocean Summit (UNOC3, Nice, June 2-13, 2025), one

month before its opening, the UIM organized a conference entitled “Ocean 2100” focusing on the longterm

challenges and risks facing the ocean 4 . Some 20 high-level experts focused in particular on the

adaptation of coastal communities to sea level rise, the protection of biodiversity and marine resources,

and the mobilization of stakeholders to address these issues. This conference, while contributing to

preparatory discussions for the Summit, should open up new activities for the IUM, particularly in the

areas of ocean acculturation and operational foresight for the seas and coastlines.

The relationship between the ocean, the arts, and science has played and will continue to play an

important role in the UIM's activities.

3

See the website https://www.univ-mer.org

4

https://www.univ-mer.org/conference-ocean-2100-enjeux-et-risques-mercredi-7-mai-2025/


Art, science, and ocean acculturation

Art has always played an essential role in stimulating human societies' interest in the ocean. Think of

the poets (Homer and the Odyssey, Virgil and the Aeneid), or closer to home, painters (Turner and his

seascapes, Boudin and his views of the Normandy coast), and musicians (Debussy and La Mer). Today,

artists are sailing aboard the schooner Tara, accompanying it on its voyages around the world to show

and convey the most remote places visited during its expeditions 5 .

But we can go further by merging art and science in the representation of marine life: the Nice-based

naturalists and painters Risso, Barla and Fossat perfectly depicted fish, shells, as well as plants, land

animals, and more 6 . This alliance between art and science was reflected on the walls of CERBOM in

black and white photographs of plankton adorning the laboratory walls, and a large mosaic in the

entrance to the building depicting stylized planktonic species. Advances in photographic techniques and

methods of observing the marine environment now make it possible to produce works, such as those of

Christian Sardet, of exceptional beauty and evocative power 7 .

In its acculturation activities, the UIM attaches great importance to the quality of representations of

the ocean in its interventions, particularly through the images used in meetings with young people or in

MOOCs for distance learning. However, it has placed particular emphasis on videography. Since The

Silent World, Commander Cousteau's pioneering work, cinema has become the primary tool for

engaging all audiences in the sea, its wonders and its secrets. The UIM has therefore set up a video

library on its website containing more than 300 works (three quarters of which are in English) covering

a wide range of subjects, from marine biology to oceanography and geopolitics, and covering different

regions of the world. This video library is freely accessible 8 .

The beauty of the ocean – whatever the way you experience it – is, in fact, the main reason why we

are interested in it, the force that draws you to it, the power that pulls you into its depths... or simply onto

the beach near the waves. This reality is well evoked by Isabelle Autissier in a radio program (in 2016)

devoted specifically to “the beauty of the ocean” 9 . She points out the psychological benefits that this

empathetic relationship with the sea brings to individuals, benefits that have been confirmed by

neurological observations. It is through emotion and the senses that acculturation to the ocean begins, as

with all things.

By developing an intimate relationship with the ocean, anyone can be drawn into a genuine scientific

activity. This is the principle of participatory science. In this regard, the UIM has been a pioneer in

organizing the Delphis operation from 1997 to 2007, bringing together every year on mid-August, a

hundred recreational boaters willing to observe cetaceans between Corsica and France. Each boater was

allocated a square mile for his/her observations, which were accompanied by wind and current readings,

temperature measurements, and water and plankton samples (using a small ad hoc net). The scientific

contributions of these campaigns helped lay the foundations for the PELAGOS marine sanctuary, one

of the first marine protected areas in the Mediterranean and in the world.

5

https://www.104.fr/fiche-evenement/la-grande-expedition.html

6

See Christian Sardet's article at https://www.openscience.fr/No-Spe-Villefranche

7

As part of the Becoming Ocean exhibition at the Villa Arson in Nice: https://villa-arson.fr/programmation/expositions/becomingocean-a-social-conversation-about-the-ocean/

8

https://www.univ-mer.org/videotheque-video-library-oceans-mers-et-littoraux/

9

https://www.radiofrance.fr/franceinter/podcasts/les-recits-d-isabelle-autissier/la-beaute-de-l-ocean-8080486


Science and the arts in the face of ocean issues

Beyond acculturation, the question is how individual and collective behavior can evolve in the face

of the major challenges and risks evoked above, resulting from human activities: rising sea levels,

declining marine biodiversity, increasing pollution, etc. This requires field surveys and research to

identify and measure the various factors that influence these behaviors: sensory and emotional contact,

as we have seen, but also the places where people live, with their topology, history and maritime identity,

social and cultural norms, ecological awareness and membership of groups committed to nature

protection, experiential learning (participatory science) 10 .

These surveys, in which the UIM should be involved, lead to recommendations on measures to be

taken or supported by public authorities where art in its various forms plays an important role: scenarios

and narratives about the future, creation of narratives 11 and symbols, storytelling, artistic innovation,

virtual and augmented reality, etc. Clearly, these analyses highlight significant differences between

cultures in terms of their relationship with the ocean and the role that the arts can play in changing these

relationships in a desirable direction.

Figure 3. This model of an Azores sailing fishing boat was carved over fifty years ago by a local artist from a

whale bone. He would probably not be able to use this material today, as hunting large cetaceans has been

banned to protect them. This illustrates the pressing issues facing the global community in the face of

declining marine biodiversity. Photo credit: Fabienne Goux-Baudiment

On a more general level, the reintroduction of art alongside science in the approach to the ocean is

part of a movement challenging the Anthropocene, seen as the destruction by humans of their biotope,

and the institutions, behaviors, and mentalities that have accompanied this behavior:

10

Stoll-Kleemann, S., Nicolai, S. (2024): Climate-Just Behavior: Foundations and Transformational Approaches. Routledge Focus.

See also Professor Susanne Stoll-Kleeman's contribution to the UIM Ocean 2100 conference on May 7, 2025, Acculturation to the

Ocean and Behavioral Change, publication in preparation.

11

Including comics, such as those developed by the French agency in charge of alerting on sea level rise (CEREMA) and presented at

the UIM conference Ocean 2100.


compartmentalization of disciplines, the unchallenged domination of capitalism, colonialism, etc. This

was the subject of a symposium held at the Museum National d’Histoire Naturelle on February 12, 2023,

whose contributions were the subject of “reflections on the links between the ocean, the arts, and

science” 12 .

This movement has given rise to a wide range of transdisciplinary initiatives and research, bringing

together the life and environmental sciences, the social and behavioral sciences (anthropology,

sociology, ethology), and of course the arts, from the most traditional (dance, song, etc.) to the most

contemporary (image and video generation using artificial intelligence). Some of this work is leading to

new methods and hypotheses in our understanding of the marine environment, for example on the

biomineralization of shells or the bioluminescence of organisms in the deep sea 13 .

Ocean and coastline foresight between art and science

Finally, it is worth mentioning an activity that combines the spirit of science and the spirit of art.In

the face of long-term issues, foresight is essential. Even if the ocean is one and must be approached as

such, in order to be operational, foresight must be conducted at the territorial level, including at the most

granular level, that of the municipality. French institutional mechanisms should facilitate such exercises,

even though they were designed for “standard” territories rather than for sea-based territories as such 14 .

Foresight, as a method to depict the future of human societies, is based on careful and in-depth

observation of reality through monitoring activities to identify major trends and structural factors, as

well as weak signals that are decisive for the future of the societies in question. It then proceeds to

exercise the imagination to think through various possible developments—scenarios, or even a single

narrative, if the future is sufficiently clear or if one is bold enough. Finally, it leads to policy

recommendations to be implemented to achieve the desired futures. Thus, foresight consists of three

stages: understanding, anticipating, and proposing. The first stage is a kind of scientific process. The

second stage is more like an artistic exercise, drawing on imagination, intuition, and a form of empathy

with the societies being observed to compensate for the limitations of reality. Finally, the third stage is

political in the noble sense of the term, as it requires the populations and authorities to take ownership

of their future in some way. Foresight thus appears at the crossroads of science, art, and politics, as

several of its initiators agree 15 .

The UIM intends to promote this operational foresight at different territorial levels, including that of

entire seas, such as the Mediterranean, by collaborating with established scientific, foresight, political,

12

https://stm.cairn.info/revue-natures-sciences-societes-2024-1-page-69?lang=fr

13

See the section “Scientific inspirations and mediations” in the above-mentioned article published in Cairn.

14

Ten-year territorial foresight studies are required in the preparation of regional master plans (“SRADDET”), and the

Development Councils (Conseils de Développement) set up at the level of urban communities are specifically responsible for

forward thinking (in an advisory capacity). In order to clearly put the focus on the sea-coastline interface, the term

“merritoires”, as opposed to “territoires” which relates to the land (terre), is proposed by Fabienne Goux-Baudiment

(contribution to the UIM Ocean 2100 conference, May 7, 2025).

15

Notably Thierry Gaudin, Qu'est-ce que la Prospective, Que Sais-je, 2013; and

Bertrand de Jouvenel http://www.laprospective.fr/dyn/francais/memoire/texte_fondamentaux/lart-de-la-conjecture-b-dejouvenel.pdf


and other institutions 16 . This foresight must be combined with acculturation efforts, in particular with

MOOCs incorporating relevant information on the countries and regions concerned.

Figure 4. This map of the western Mediterranean, dating from the Renaissance, is surprising because the

north is located on the right side of the frame and not at the top, as it became standard a few centuries later.

Nevertheless, the overall cartography—including the shapes of the coasts, bays, and capes and their

positioning—is relatively close to reality. Photo credit: Fabienne Goux-Baudiment

Conclusion

Ultimately, for human societies to take back control of their future in the face of ocean movements,

they will need to take steps that are both scientific and artistic, drawing on both sides of the brain, so to

speak—the left side, which is the seat of reason, and the right side, which is the seat of emotion. This

takeover also requires mobilization at all levels of granularity around which societies are organized, from

the individual to the global community.

16

Collaborations are envisaged with the IPOS (International Panel for Ocean Sustainability) platform, the Plan Bleu pour la

Méditerranée, and the 2100 Foundation.


N° Spé : UNOC 2025

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