François Alphonse FOREL and the oceanography of lakes · 2013-06-06 · In 1892, F.A. F OREL (FAF)...

| François Alphonse FOREL and the oceanography of lakes | 51 |

| ARCHIVES DES SCIENCES | Arch.Sci. (2012) 65 :51-64 |

François Alphonse FORELand the oceanography of lakes

Warwick F. VINCENT1* and Carinne BERTOLA2

Ms. submitted 25th July 2012, accepted 1st September 2012

1 Centre d’études nordiques (CEN : Centre for Northern Studies) & Département de biologie, Université Laval, Québec City, Québec,Canada. E-mail : [email protected]

2 Musée du Léman, Nyon, Vaud, Switzerland. E-mail : [email protected]

* Corresponding author : Musée du Léman, Nyon, Vaud, Switzerland. E-mail : [email protected]

❚AbstractIn 1892, F.A. FOREL (FAF) created and defined the new science of Limnology as “the oceanography of lakes”. His aim was toestablish an integrative discipline for the aquatic sciences in which diverse types of lake studies, from physics, chemistry andbiology, to anthropology and economics, would complement and inform each other to produce a meaningful synthesis. FAFgrew up in Morges, Switzerland, on the shores of Lake Geneva (lac Léman), and then left for 11 years to universities in Franceand Germany to study the natural sciences and medicine. Shortly after his return to Switzerland, he was appointed Professorof Anatomy and Physiology at the Academy of Lausanne. In the first years back at the lake, he made important discoveriesabout deep-living benthic animals and surface seiches that reinforced his early decision to adopt Lake Geneva as his “labo-ratory and aquarium” and to devote his career to limnological research. FAF’s remarkable breadth of interests and expertiseculminated in 288 reports and publications, including the first text book in general limnology (published 1901), and the sem-inal, three-volume monograph on the Limnology of Lake Geneva (1892, 1895 and 1904). FAF’s success was the result of hispassion for lakes and lake science, an ability to pose and critically evaluate insightful questions about the natural world, aflair for observation and for new observing technologies, a rigorous, encyclopedic ability to collect and synthesize all avail-able information, and a natural talent for networking, collaboration and knowledge transfer. His view of lakes as coupledphysical-chemical-biotic-human systems is particularly relevant to facing the challenges of global change, and the associatedrapid shifts in ecosystem services at a planetary scale.Keywords : Ecosystem services, Forel, Global change, History of Science, Lake Geneva, lac Léman, Limnology

❚1. Introduction

In the preface to his three volume treatise on LakeGeneva (known in Switzerland by its French name,Lac Léman), Prof. F.A. FOREL (Fig. 1) struggledwith the question of what he should call his integra-tive approach to lake science. His research encom-passed all aspects of the lake, from its morphometry,sediments, underwater light and mixing patterns, toits microbial, plant and animal ecology. He observedthat the environmental characteristics of the lakewere all closely interrelated, and he saw the lake as asystem that was intimately tied to its surroundingcatchment. As a doctor of medicine and the son of alocal jurist and historian, he also identified the lakeas a precious resource for the health and well beingof the many people who lived around it in the pastand present, and he quantified the lake fishing econ-omy in a way that foreshadowed the modern conceptof ‘ecosystem services’.

The broad sweep of F.A. FOREL’s research interestson the lake was simply too large to classify within anyspecialised subject area, and there were problems try-ing to insert it into any existing integrative discipline,as he explained in his preface (Forel 1892; p.VI; Fig.2) : I wanted to achieve a generalisation, an over-view of all the detailed facts, where each speciali-sed study would be supported by the data fromother studies. The theme of my description beingpartly terrestrial, this subject might be consideredGeography. But the geography of waters is calledOceanography ; I could therefore call the disciplineFreshwater Oceanography. But a lake, no matterhow large it might be, is not an ocean ; the limitedexpanse gives each lake its own proper characterthat is very different from the unlimited expanseof the vast ocean. F.A. FOREL went on to explain thatthis lack of fit to any existing discipline forced him to

| 52 | Warwick F. VINCENT, Carinne BERTOLA François Alphonse FOREL and the oceanography of lakes |


create a new word for a new discipline : ... it is neces-sary to forge the word limnology. Limnology isthus the oceanography of lakes.With 100 years since the passing away of F.A. FORELon 8 August 1912, it is an occasion to celebrate andtake stock of his remarkable accomplishments. Fromthe early 20th century onwards, F.A. FOREL has beenrecognized internationally as the founding authorityof lake science. In what may be the first English lan-guage textbook on limnology, J.G. Needham(Professor of Limnology at Cornell University) andhis co-author J.T. Lloyd (1916, p.76) noted the pro-found influence of the Swiss Master, F.A. Forel,who is often called the ‘Father of Limnology’. Hewas the first to study lakes intensively aftermodern methods. He made the Swiss lakes the bestknown of any in the world. His greatest work “LeLéman”, a monograph on Lake Geneva, is a mas-terpiece of limnological literature. It was he whofirst developed a comprehensive plan for thestudy of the life of lakes and all its environingconditions.Needham and Lloyd’s tribute describes an impressiveset of achievements, but F.A. FOREL’s life work andinspiration extend even beyond this list. His lesserknown contributions include seminal insights intolakes as microcosms, hydrologic optics and lakewater color, the nature of pelagic and littoral commu-nities, the structure and function of aquatic food websincluding microbial decomposition processes and

organic carbon cycles (acknowl-edged and praised by Raymond Lin-deman some 40 years later), obser-vations on the ecology of invasivespecies, notes on the intimate con-nection between human society andits freshwater resources, and the roleof Homo sapiens as a powerful,biotic component of lake ecosys-tems. This F.A. FOREL perspectiveon Lake Geneva and its surroundingsas a coupled physical-chemical-biotic-human sys tem seems particu-larly relevant today as we face theenormous challenges of globalchange, and the associated rapidshifts in ecosystem services at a plan-etary scale (Williamson et al. 2009).

❚2. F.A. FOREL’s early life

François Alphonse FOREL (FAF) was born on 2February 1841 in the Swiss town of Morges, where theForel family traces its ancestry back to Claude Forel, aparish officer and councillor who became establishedthere in 1589. Morges lies 16 km from Lausanne and 50km from the city of Geneva, but most importantly forthe young FAF, the family home was on the shores ofLake Geneva that provided clean water, boat trans-port, productive fisheries, spectacular scenery, andmany opportunities for exploration. His father,François Marie Etienne Forel (1765-1865) was arespected jurist-and historian, and FAF grew up sur-rounded by family members and friends who haddiverse scholarly interests. His cousins included AlexisForel (1852-1922), a chemist who became a well-known engraver, and Auguste Forel (1848-1931), whoworked in the disparate fields of psychiatry and antbiology. FAF notes that one of his earliest recollectionson the lake was at age 13, working with archaeologicalfriends of his father to search for bronze-age artefactsin the submerged littoral zone. As in many Europeanlakes, Lake Geneva contained the remnant pilings of‘stilt villages’ that dated back several thousand yearsbefore the present. Using a modified grab from hiscanoe, FAF was able to snare three ancient braceletsfrom the lake floor, much to the surprise and perhaps

Fig. 1. Water color portrait of F.A. FOREL

in the room he used as his study and labo-

ratory, at his home in Morges on the

shores of Lake Geneva (Lac Léman),

Switzerland. In 1892, F.A. FOREL founded

the new science of Limnology, defining it

as “the oceanography of lakes”. (Undated

painting by Ernest Biéler, reproduced by

permission of the Forel family).

| François Alphonse FOREL and the oceanography of lakes Warwick F. VINCENT, Carinne BERTOLA | 53 |


chagrin of the professional archaeolo-gists on board their adjacent boat. FAF’s initial schooling was in hishome town at Morges (Collège deMorges), and then later in Geneva atthe western end of the lake, wherehe attended secondary school(Gymnase de Genève) followed byuniversity studies at Geneva(Académie de Genève; Bachelor ofLetters and Bachelor of Physical andNatural Sciences). At the age of 18,he left Switzerland for France tostudy medicine and natural history atthe Académie de Montpellier for 2years. He continued his medicalstudies in Paris, where his activitiesincluded training visits to hospitalsas well as courses and seminars atthe Museum of Natural History. Hethen moved to the University ofWürzburg, one of the oldest and mosttraditional universities in Germany,to complete a final year of medical studies, graduatingDoctor of Medicine at the age of 24, in July 1865.During this period, he developed a great interest incomparative animal anatomy (zootomy), and for thenext three years he helped teach this subject at theuniversity. Although he was encouraged to continuehis career in Germany, he had been away from Morgesfor 11 years and now felt it time to return to look afterhis parents, whose two daughters had died at a youngage. In 1867, he returned home and began to under-take studies on Lake Geneva. He was engaged as a lec-turer in the Faculty of Sciences at the Academy ofLausanne in 1869, initially to teach microscopy andhistology, and the next year was appointed to the posi-tion of Professor of physiology and anatomy in thisfaculty (Bertola 1999).

❚3. Early career discoveries andnetworking

Already a professor at the age of 29, FAF had a fairlylight teaching load at the Academy, with 3 hours oflectures per week. Plenty of time was thus availablefor research, and he had the academic freedom to

choose his subject. FAF acknowledged that settingup a science program in anatomy, histology or physi-ology was the most logical set of options given histeaching duties in these subjects. But instead hemade the decision that his laboratory would be LakeGeneva, and that the subject of his research wouldbe all aspects of the physics and ecology of the lake.His former professor at Würzburg, Albert Kölliker,greeted this decision with some concern, and duringhis visit to Lausanne in 1871 he suggested that theyoung man focus on just one single, zoological topic.Fortunately, FAF was not to be dissuaded. His firstyears of research had already generated some veryexciting results that likely reinforced his plans for acareer in lake science. From the earliest days withhis father onwards, he had a passion for working onthe lake and unravelling its secrets, so much so thatwhen later in life he was invited by an editor to writea textbook in limnology he initially refused, sayingthat he could not remain calm and impartial aboutthis subject. He notes in the preface of the textbookthat eventually emerged from this discussion : Myrelation to limnology is much too personal andsubjective for me to be able to give an objectivepresentation of the facts (Forel 1901). The preface

Fig. 2. The page of the original handwrit-

ten manuscript on which FAF defines

‘Limnology’ for the first time. This and

other FAF manuscripts are archived in

the Forel Documentation Center at the

Musée du Léman (Lake Geneva Museum)

in Nyon, Switzerland

(www.museeduleman.ch by WF. Vincent).



then goes on to explain the evident,that the editor was not persuadedby this objection, and the projectwent ahead.One of FAF’s early successes was theaccidental discovery of a deep-livingnematode in the lake. In April 1869,two years after his return from Germany, he was mak-ing observations on the deposition and rippling ofsediments on the lake floor, and with a metal sam-pling plate he brought up some sediment, from adepth of 40 m offshore from Morges. Placing a sub-sample under the microscope to examine the natureof this silty material, he was startled to see a nema-tode swimming about, a species identified as Mermisaquatilis. He was immediately overcome by an excit-ing revelation : that the lake floor of Lake Geneva wasnot a sterile desert as previously thought, but in factcould be the habitat for a specialized community ofdeep-living benthic animals. The very next day heconstructed a dredge and returned to the lake tosample it at many depths, confirming that diversespecies of animals occurred in the sediments, even to300 m, the deepest abyssal region of the lake. Thiswas the start of a long term project that achievedenormous success, in part because of FAF’s earlydecision to work collaboratively with specialists ondifferent animal phyla rather than keeping the sam-ples and discoveries to himself. From 1874 to 1879,this project produced a series of 49 published reportswith 19 collaborators (Fig. 3) and later culminated inFAF’s 1885 publication La faune profonde des lacssuisses (The Deep Fauna of Swiss Lakes). This vol-ume in fact is broader than suggested by the title,with sections on physical and chemical limnology,and descriptions of several large lakes of the world.The ecological historian Frank Egerton comments :The Société Helvétique des Sciences Naturelles[Swiss Society for the Natural Sciences] hadrequested the work and awarded a prize for it. Itmust have been the best general work on freshwa-ter biology available at the time (Egerton 1962).

In those first years back at Lake Geneva, FAF madeanother discovery that also defined his career overthe subsequent decades. The boat harbor at Morgeshad two exits to the lake, one for the boats and a sec-ond much smaller one only 2 m across to allow waterexchange. From 7 May 1869 to 11 May 1870 he took aseries of careful measurements of the flow of waterthrough this smaller aperture, discovering that thecurrents continuously oscillated in direction, some-times flowing into the harbor and then a few minuteslater reversing to flow back out into the lake. Twoinsightful revelations came to him as he observed thiscurious pattern of oscillating flow. Firstly he sur-mised that it was somehow connected to the periodicrise and fall of lake level that had been noted aroundthe lake and especially at the city of Geneva for hun-dreds of years, and that was referred to in the localSwiss French dialect as a ‘seiche’, meaning rockingbackwards and forwards. Secondly and most impor-tantly, he concluded that these phenomena were theresult of standing waves propagating throughout thelake. His first paper on this subject was publishedshortly thereafter (Forel 1873), and he continued toamass data from his own observations and from othersources around the lake. This interest in physical lim-nology was likely stimulated and encouraged by hisformer professor of mathematics and physics, andlater research and teaching colleague, Dr. CharlesDufour, whose broad research interests includedmeteorology, glaciology and geology (Egerton 1962).In 1876, FAF constructed the first of a series of lim-nographs, an automated instrument that couldobtain a continuous record of lake level. This devicefollowed the design of tide gauges used by oceanog-raphers, but unlike the marine versions at the time, it

Fig. 3. One of the deep-living

invertebrates found in the sediments of

Lake Geneva. FAF’s zoobenthos project

resulted in 49 scientific reports in collabo-

ration with many taxonomic specialists

from within and outside Switzerland, and

stemmed from his discovery that lakes

contain a distinct abyssal fauna adapted

to the cold dark bottom environment. This

water mite was described by the distin-

guished medical pathologist Hermann

Lebert as the new species Campognatha

foreli, now Hygrobates foreli. From F.A.

FOREL (1879). (Image by W.F. Vincent).



could resolve mm-level fluctuations in water level,and even picked up the vibrations of steam shipspassing many km away. Over the subsequent 20 years, FAF worked withmany colleagues at Lake Geneva, Lake Constanceand elsewhere to define the properties of surfaceseiches, and he corresponded on this subject withscientists around the world, including specialists inFrance, Germany, England, Australia, Japan and theUSA. He was frustrated by his lack of formal trainingin differential calculus, but through his network ofcolleagues he found the expertise that he needed todevelop a quantitative theory of seiches. One of hiscontacts at the University of Leipzig passed on to hima treatise published in 1828 by a family member, amathematician by the name of J-R Merian who hadapplied Lagrangian analysis to the problem of watermovement in a rectangular tank. FAF transformedand applied this equation to lake basins, and foundthat he could estimate the average depth of the lakebased on the period (t) of the seiche. However, hewas still unsatisfied with the complexity of the equa-tion, and continued to discuss this in his meetingsand correspondence with other researchers. It was finally a distinguished physicist at theUniversity of Glasgow who provided the aid that FAFwas seeking. In a letter to FAF dated 6 October 1876,Professor William Thomson, better known as LordKelvin, took the Merian-Forel formula, simplified itfor the condition that lake depth (h) is very small relative to lake length (L), and deleted the minorterms to produce the elegant approximation :

budget that totalled 15000 Swiss francs. Perhaps forthis latter reason, the plan was not implemented, how-ever it stands as the first comprehensive environmen-tal research proposal for a lake, and its formulation andsigning 140 years ago might be considered the birth oflimnology (Bertola, 1999). At the very least, it set theresearch agenda for FAF over the next few decades,which ultimately resulted in his three volume treatiseon the limnology of Lake Geneva, covering all of thetopics described in this 1872 report.

❚4. Correspondence and monographs

Throughout his 25 years of tenure at the Academy ofLausanne, FAF maintained in addition to his teachingduties, a continuous output of scientific publications,frequent presentations at scientific meetings, andprolific correspondence with colleagues in Switzer-land and abroad. He thrived on collaborations, forexample with the renowned French limnologist AndréDelebecque (author of the classic monograph onFrench lakes; Delebecque 1898) to produce the firstbathymetric maps of Lake Geneva. The dozens ofletters archived at the Center for Documentation(Fig. 2) include an exchange with John Le Conte, Professor of physics and former president of the Uni-versity of California at Berkeley, and the first limnolo-gist to work on Lake Tahoe, California-Nevada.Among his remarkable diversity of research topics,FAF had been fascinated by the interplay betweensunlight and lakes, both at the lake surface and in thepropagation of light through the water column. Heestablished a standard protocol for the use of the Sec-chi disk, a white circular plate that had been inventedby Priest Angelo Secchi to measure the transparencyof the Mediterranean Sea. It was extensively tested byFAF in Lake Geneva as early as 1874 (Forel 1895 ; p.410), only a few years after its publication (Secchi1866). FAF then applied his standardized Secchi diskmethod to describe the seasonal dynamics of lakewater transparency, showing the variations fromaround 15 m in winter to around 5 m in summer. Healso developed a number of novel approaches inhydrologic optics, including an in situ method basedon photographic plates to measure photochemicallyactive radiation, and a color scale based on mixturesof colored reagents to quantify lake color (Fig. 4). In a letter dated April 9th 1884, Le Conte responds toFAF’s query as to why Tahoe was apparently moretransparent than Lake Geneva, and suggests that it isthe result of an absence of active glaciers in the Tahoebasin. Le Conte also expresses his interest in installingone of FAF’s high-tech limnographs in Lake Tahoe tomeasure seiches, but he mentions the difficulty offinding people who are attracted to science ratherthan profit : … it is difficult to find an investigator.Surrounded as you are by all the adjuncts of an

In a detailed discussion and application of this equa-tion, Forel (1892) noted that this was appropriateonly to basins of simple morphometry, and that formore complex basins, other formulations were re-quired, such as that developed by the French engi-neer Paul du Boys, another colleague whom FAF hadrecruited into his seiche discussions.Over the period 1871-2, FAF participated in two meet-ings at Nyon to define a broad of spectrum of researchon Lake Geneva. These meetings had been formallycommissioned by two scientific societies, and broughttogether distinguished scholars from Geneva andLausanne. Their work culminated in a report preparedand signed by FAF as secretary of the commission, on3 June 1872, that outlined 14 research topics thatneeded to be addressed, including geomorphology,bathymetry, sedimentology, lake transparency, cur-rents, water chemistry, aquatic flora and fauna, andthe archeological features of the lake. The documentwas published that year in No. 401 of the Bulletin ofthe Vaud Society of Natural Sciences (Vaud is the can-ton or governing district along the northern shore ofLake Geneva), and it described the limnological meth-ods that would be applied, as well as a proposed

WVincent

Sticky Note

This should read: t = L / √(gh)



old civilization, it is difficult for you to appreciatethe conditions of our new and undeveloped civili-zation. The overwhelming influence of the com-mercial spirit almost paralyzes the genuine see-ker of truth. The worship of gold inspires all clas-ses of society; and the true scientific sprit iscorrespondingly stifled…Reciprocating yourdesire to have closer scientific relations, I remainwith sentiments of the highest regard and esteem,Yours Most Sincerely, John Le Conte.FAF’s work over the next two decades resulted indozens of reports, notes and scientific articles, butthroughout this time he was working towards the big-ger synthesis. His many articles on benthic fauna inLake Geneva and other lakes were brought together ina collected volume in 1885, and a year later he revisedand published a little guidebook to Lake Geneva thatbriefly touched on many of its limnological features(Forel 1886). All of this work combined with his ency-clopedic collection of facts about the lake was to feedinto the ambitious overview that he had embarked ontwo decades earlier. On 2 February 1891, perhaps tomark his 50th birthday that day, FAF submitted thecomplete manuscript of the first volume of Le Léman– Monographie Limnologique to the printer. Aftermore than 18 months of proofs and painstaking cor-rections by himself and his colleagues, Volume I wasfinally published, in August 1892.Forel (1892) is a remarkable work in many ways. Itbegins by defining Limnology as a new branch of sci-ence, and articulates the vision that he had formu-lated as a young scientist : to describe Lake Genevafrom diverse yet connected points of view. He dedi-cates the book to his father, to whom he pays homagefor introducing him at the age of 13 to scientific studyand to the art of observing and interrogatingNature ; I have continued under the eyes of mydear [father] to work on the numerous and diverseproblems that a lake, a true microcosm, poses tohuman curiosity ; encouraged and guided by hiscounsel I have devoted the best of my activities as anaturalist to this research (p. V). The Preface also

explains the intended audience, actually many audi-ences : fellow scientists, other readers interested in anexplanation of natural features that they are intri-gued by or admire, the people who live around thelake for whom the vast water mass of Lake Genevais an ocean that intervenes in so many aspects oftheir individual lives and society, and the boatmenand fishermen who live on the lake, who live fromthe lake, and who hold the precise knowledge andmethods for their professional activities (p. XI). Inthis defining moment for limnology, FAF thus cap-tures the broad spectrum of end-users of lake science,from fundamental to applied limnology. In so doing, healso shows how human beings are an integral part ofthe lake ecosystem, a theme that he comes back to ingreat detail in the third volume.Volume I presents a rich compendium of information,from a summary of field apparatus for limnologicalstudies, to detailed descriptions of the geographic set-ting, hydrographic features, geology, climatology andhydrology. Throughout there is a meticulous attentionto earlier work, the history of development of knowl-edge on each subject and full acknowledgement ofprevious authors and investigators, including manyunpublished data sets. The volume finishes with hisbathymetric map of Lake Geneva, with the 25 m iso-baths based on 11 955 soundings. FAF notes that 4338of the soundings came from the distinguished Frenchscientist André Delebecque, his respected limnologi-cal counterpart who was based at Thonon, on theopposite side of the lake. FAF submitted Volume II of his treatise two yearsafter the first, and it was published in December1895. In terms of limnological theory, this has turnedout to be the most important of the three volumes,with sections on lake hydraulics (including seiches),thermal characteristics of the lake and hydrologicoptics. These were followed by a short section onlake acoustics (focusing on the speed of sound inwater and air), and a 70 page section on water chem-istry. In a chapter in the latter section entitledDissolved materials in the water of the lake heclearly links lake chemistry to catchment properties,for example : The sulfate concentration of 64 mg[L-1], chalk sulfate in particular, is considerable.It is due to the presence of gypsum in many val-leys of the watershed, in particular in the largevalley of the Rhone… (Forel 1895 ; p.608). He notedthat the dissolved organic carbon concentrationswere low but variable, perhaps associated with vari-ations in rainfall and runoff, and asked the question,still non-trivial today What is the nature of theseorganic materials in the lake water ? This ques-tion has not been sufficiently studied (Forel 1895;p. 616). On the sampling date of 30 November 1880,the lake was still stratified, but well oxygenated evenat the bottom, with 7.08 cm3 O2 L-1 at 300 m depth(Forel 1895 ; p. 622). FAF thereby established an

Fig. 4. The F.A. FOREL scale for lake color. From Forel (1895).

(Image by W.F. Vincent).

WVincent

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important baseline value for gauging the magnitudeof change in Lake Geneva during its period of rapideutrophication in the 20th century.Forel (1895) presented a huge amount of thermal datafor the Lake Geneva water column based on dozens ofprofiles from his own work (much of which was with ahigh resolution reversing thermometer), as well asdata from many colleagues including AndréDelebecque. He made heat budget calculations for thelake, and identified the importance of density currentsand convection. His thermal profiling allowed him todefine the annual pattern of stratification and mixing,as well as interannual fluctuations in hypolimnetictemperatures. FAF developed a logical way to classifylakes based on their stratification regimes : ‘polar lakes’defined as those in which ice and cold water overlywarmer water during winter stratification, followed bymixing in summer; ‘tropical lakes’ in which warmwater overlies cold during the stratified period fol-lowed by mixing in winter; and ‘temperate lakes’in which there is an alternation between the ‘polar’and tropical’ modes of stratification, separated byperiods of mixing. Hutchinson (1957) later acknowl-edged and adopted FAF’s approach, but replaced theconfusing terminology in which Lake Geneva is classi-fied as a ‘tropical lake’ despite its temperate locationby terms based on mixing each year (cold monomictic,warm monomictic, dimictic, etc). FAF’s approach was always to formulate questionsand to test ideas, often by way of competing hypothe-ses and an encyclopedic synthesis of observationsfrom all sources including his own original work. Thisled to rapid insights and rejection of alternatives, forexample in his work on surface seiches in which hereadily eliminated hypotheses for causal mechanismssuch as electrical storm effects and periodic injectionof melt waters from glaciers. Not all of his conclusionshave survived the test of time, and he himself fullyacknowledged that science is a continuously evolvingprocess. Glacial erosion, for example, is now thoughtto have played a much greater role in carving out theLake Geneva basin than FAF surmised (Wildi et al.,1999). FAF was reluctant to accept the presence ofinternal waves in the lake, despite evidence in his owndata of thermocline tilting, which he erroneouslyattributed to the mixing of more dense water from theRhone at the western end of the lake (Forel 1895,p.354). In fact the evidence and theory from Scotlandof internal waves came late in his career, and at anunfortunate time when he had a medical problem withhis hands that curtailed his fieldwork on the lake.

❚5. Human and biological limnology

At the age of 54, and after 25 years of academic serv-ice as a professor, FAF retired from the Académie deLausanne, and was then able to devote even more

time to his limnological studies. His massive collec-tion and synthesis of information continued, as didhis prolific output of publications. He produced hislimnological textbook in 1901 (Handbuch derSeenkunde : Allgemeine Limnologie), and pub-lished Volume III of the monograph in May 1904. Thelatter included the biological limnology of the lake, itshistory of human occupation going back to neolithictimes, the navigation history of the lake up to thepresent, and the fishery.From his childhood onwards, FAF was intenselyaware of the significance of Lake Geneva to the peo-ple who lived around it, including for the many gener-ations of his own family who lived at Morges. In hisdevelopment of the new science of limnology, FAFmade it quite clear that human activities, past andpresent, were part of the structure and functioning oflacustrine systems. In Volume III of the monograph,FF details the aquatic and semi-aquatic biota of thelake, and the first species he defines as follows :

VERTEBRATES;

MAMMALS, PRIMATES :

Man, Homo sapiens L., is not an essentially aqua-tic species, but has become so by way of his activi-ties; the calling of fishermen, sailors, ... etc resultsin many such people living a semi-lacustrine life,making mankind almost an erratic species of thelake fauna. (Forel 1904, p. 26). Forel goes on to notethat humankind also has negative impacts on the lakewaters : he builds ports and quays, he channels theinflows, he dams the outflow, he discharges intothe lake the products of his factories, the sewers ofhis villages, the ashes of his steam boats etc. All ofthese actions modify the lake environment, anddirectly or indirectly impact on the biologicalfunctions of its inhabitants. Mankind exerts amore powerful effect than any other animal onNature and its inhabitants (Forel, 1904 ; p. 26).On this same theme, FAF also documented the arrivalof invasive plant and animal species in the lakethrough human means. For example, he describeshow the North American macrophyte Elodeacanadensiswas imported to Geneva in 1869 and wasplanted in ponds and streams to favor fish produc-tion. By 1880 it had escaped into the lake, and threeyears later invaded the boat harbor near FAF’s resi-dence at Morges. After the first few years of spectac-ular growth and displacement of other aquatic plantspecies, by 1900 its population size had begun to sta-bilise and contract, and all worries about the uns-toppable invasion by this American weed disap-peared. After its phase of exuberant and frighte-ning expansion over several years, Elodea settleddown as a calmer vegetation type. It became of nogreater concern than the Potamogeton andMyriophyllum that grew beside it. (Forel 1904 ; p.



162 ; Fig. 5). He also describes the introduction andproliferation of mute swans on the lake and reportshis detailed observations on their phenotypic varia-tion and behavioural ecology, noting for example :Although of very little intelligence, swans canhave strong passions and even show signs of cor-ruption that are truly deplorable. (Forel 1904 ; p.38).FAF’s Volume III provides a detailed overview of theplant and animal communities of Lake Geneva, sepa-rated according to the semi-aquatic edge of the lake,the littoral and pelagic zones, and the abyssal regionwith its deep-living benthic communities. From hisdiscoveries in 1869 onwards, the latter communitieswere of special interest to FAF and he devotes manypages to describing the communities, including thedeep living moss Thamnium lemani; microbialmats, in part composed of oscillatorian cyanobacteria(especially abundant during the period of higherwater column transparency in winter and spring);and 79 species of animals ranging from protists tofish. He describes the monotonously cold, dark cli-mate in the deep region of the lake and asks the ques-tion, do the organisms of this zone show any generalcharacteristics ? He identifies several common fea-tures and then goes on to consider the origins andevolution of the Lake Geneva biota. He was alsogreatly interested in the deep-living populations ofcrustacean zooplankton and their nocturnal migra-tion habits : These animals, capable of extensiveactive movement, come to the surface during thenight – perhaps only the nights when the lake iscalm, perhaps also only those nights when themoon does not shine too brightly – and they des-cend during the day to depths that are less illumi-nated or in obscurity (Forel 1904; p. 222). FAF also

gives credit to Auguste Weismann (1877) for inde-pendently coming to the same conclusion in LakeConstance.

❚6. Microbial limnology

FAF’s Volume III also provides an introduction toaquatic microbial ecology, and he makes someinsightful observations that are still highly relevanttoday. The methods in microbiology available at thetime were of course extremely limited relative tothose now currently available, but the microscopeshown in the painting of his laboratory (Fig. 1) wouldhave provided him with the first indications of LakeGeneva’s microbial world. He collected together allthe available bacterial count data from water hygienerecords, and noted (despite estimates many orders ofmagnitude below those obtained with modern meth-ods) their great abundance everywhere : No seriousanalysis, to my knowledge, has indicated any lakewater completely free of microbes (Forel 1904 ;p 360). He pointed out from this synthesis that thebacterial population size is especially high in the sur-ficial sediments, in rivers, in waste water, and parts ofthe lake near sewage outfalls, with lowest concentra-tions in the offshore pelagic zone. He separated thebacteria into functional groups, and underscored theessential role that they play in the carbon cycle oflake ecosystems through the degradation of organicmaterials. In Volume II, FAF had observed that even clean natu-ral waters contain microbes, and he cautioned thereader not to be alarmed about this : Secondly, andthis is a very important point, all microbes are notnecessarily unhealthy. Much to the contrary, theimmense majority of these minute beings are com-pletely innocent (Forel 1894 ; p. 637). In Volume IIIhe goes further in a chapter entitled The Circulationof organic material to explain how mineral materialsentering the lake are converted to organic materialsthat are distributed among three forms : A. Dissolvedorganic matter forms the general reserve that isstored in the waters of the lake....B. Organic mate-rial in the bodies of dead organisms and detritus(débris morts)....C. Organic matter in the state ofliving plants and animals, and that these compart-ments are connected, ultimately forming substratesfor microbial decomposition and remineralisation :How many successive incarnations is this organicmaterial subjected to as it passes from plants to theanimals that feed on it ! An alga, for example a dia-tom, is eaten by a rotifer, which is eaten by a cope-pod, which is eaten by a cladoceran, which iseaten by a whitefish, which is eaten by a pike,which is eaten by an otter or by a human. Thesmall and the weak are the prey for the large andthe strong, and they in turn are eaten by the larger

Fig. 5. F.A. FOREL and associates collecting aquatic macro-

phytes from Lake Geneva. F.A. FOREL observed Elodea

canadensis with great concern when it entered the lake as an

invasive species. (Image courtesy of the Musée du Léman and

the Forel family).



and stronger, or if they escape, they will not avoidthe microbes responsible for decomposition, whichall organisms are directly or indirectly subject to.After this incarnation in the plant or animal, theorganic material passes from an organized state toa state of solution. (Forel 1904 ; p. 368) Decadeslater, the celebrated ecologist Raymond Lindemanpraised FAF for providing a brilliant exposition ofthe general nature of food cycles that will serveeven today as an introductory account of trophicrelationships (Lindeman 1941 ; cited in: Sterner,2012).Although FAF was unaware of the enormity of thetotal bacterial biomass and respiration in LakeGeneva, his rough calculations of the allochthonousinput of organic matter led him to conclude that alarge part of the organic material that enters thelake from its inflows is emitted into the atmo-sphere as a result of the gases that are products ofdecomposition and putrefaction over organicmaterial living and dead, escaping without stopinto the atmosphere (Forel 1904 ; p. 369). Perhapsthis is the first recognition of lakes as heterotrophicecosystems.

❚7. Lake ecosystem services

The historical importance of a lake to its people was atheme that FAF had been introduced to as a child. Hisfather was very interested in the palafittes, thebronze-age villages built on stilts within the littoralzone of Lake Geneva, and other lakes in the region(Fig. 6). FAF participated in his father’s archeologicalexploration of the extensive site at Morges, whereseveral thousand remnant pilings could be found inthe lake, and this stimulated his lifelong interest inthese ancient cités lacustres. In Volume III of his

treatise he devotes 78 pages to this subject, docu-menting the artifacts that had been collected (somedating back to the stone age) and formulatinghypotheses about the mode of life of these prehis-toric peoples, their origins, the plants and animalsthat they depended upon, and the value of the lake tothem for its multiple ecosystem services : Add to thisnotion of security against any kind of attacks, theundeniable allure that such constructions overthe lake would offer... Having the lake in front,around, below ; having only to jump down to takea bath or travel by canoe ; to simply throw a net toharvest abundant fish; to be warmed in winter bythe tepid atmosphere of the lake and to be refres-hed in summer by comforting lake breezes ; tohave clean sanitation for disposal to the lake ofhousehold waste ; to enjoy the constant changes inthe mood of the lake, sometimes calm, other timeswhipped up by storms, bathed in light, sometimesin the sad grey hues of mist... We the residents ofthe lake, we know the powerful charm, alwaysrenewed and rejuvenated, that these waters pro-vide... (Forel 1904 ; p. 448). More recent studies haveindicated that many of these houses may have beenoutside the lake, and that the stilts may have alsooffered protection against changing lake levels.In the final sections of Volume III, Forel (1904)addresses two aspects of Lake Geneva’s geo/ecosys-tem services in quantitative economic terms : shiptransport and commercial fishing. He describes thelong history of navigation on the lake, from thecanoes of the stilt villagers to naval vessels, merchantbarques, fishing boats and the arrival of steam ships.FAF quantifies the commercial steamship tonnage atthe time and its passenger capacity, and to illustratethe commercial value of these activities he presentsthe financial report of the steamship company CGNbased out of Lausanne. This company, even more

Fig. 6. A reconstruction of a bronze age stilt village on Lake Constance, Germany, similar to the type surmised from the archae-

ological remains at Lake Geneva. The archaeology of this “cité lacustre” and the close relationship of its people to the lake were

of great interest to F.A. FOREL and his father. (Image by D. Straile, Limnological Institute, University of Konstanz).



active on Lake Geneva today, reported that itsreceipts for passenger and freight services in 1901totaled 1.24 million Swiss Francs. He concludes thenavigation section with chapters on the laws and cus-toms for vessels on the lake, the evolution over hun-dreds of years of the distinctive merchant barquedesign and the history of ports on the lake. In considering the fishery, Forel (1904) was awarethat several fish species (notably lake trout, Arcticchar, whitefish, carp, pike and perch) had been partof the diet of lake residents from prehistoric timesonwards. He traces the laws governing the manage-ment of the lake fishery back to 1043, when KingHenry III granted Archbishop Hugo de Besançon theright to fish in Lake Geneva. FAF then details the har-vest data, the tax imposed on catches, and the >300-fold rise in fish prices per pound over 5 centuries. Hebegins a chapter entitled Statistics by statingStatistics on the Lake Geneva fishery are almostnon-existent. The State, which is uniquely compe-tent and capable of obtaining such information,has consistently retreated from the difficulties ofthese tasks. He also notes the natural suspicion offishermen towards the collection of such statistics,and their legitimate fear that this would translate intoyet more regulations and taxes. Not to be deterred,FAF combed the regions for all available numbers onthe French as well as Swiss sides of the lake, notingthe particular lack of data from the latter. He pres-ents the annual exports from France to Switzerlandof fish caught in the lake (49 to 122 tonnes per year),and provides the annual receipts for a wholesale fishmerchant at Geneva. All of these statistics allow himto make the first order estimates that 1200 fishermanworked at that time on the lake, and that the fisherygenerated retail sales of around 1.5 million SwissFrancs per year. Concerning the latter estimate, hecautions that it is Not necessary to emphasize theuncertainty and imprecision of this number(Forel 1904; p. 650).

❚8. Contributions outside limnology

The name Forel is well known to most limnologists,but several other branches of science have alsogreatly benefited from his observations and insight,including glaciology, seismology and archaeology. Hiscontributions were so numerous (a publication list of288 titles was published with his obituary; Blanc,1912) and diverse that at the commemoration cere-mony for FAF and the unveiling of a special medallion,two years after his death in 1914, Paul-LouisMercanton noted : François A. Forel was a greatnaturalist, his name was known everywhere andfor such diverse reasons that many a scholar out-side Switzerland thought that there must existeveral François Forels (p. 39).

Throughout his life, FAF enjoyed trips to the SwissAlps and explored many glaciers of the region. Hisfather had taken him at the age of 14 to the Sea of IceGlacier (Mer de Glace) on the northern slope of MtBlanc, and had stimulated his scientific interests bypouring a little red wine on the ice, showing how itflowed into the fissures to reveal the crystalline struc-ture of the glacier. FAF published many glaciologicalstudies and over the 30 year period, published anannual report on the state of Swiss glaciers. He wasinstrumental in setting up the InternationalGlaciological Commission, and was its first president,later honored by the naming of the Forel Glacier (lat.67.48 oS, long. 66.50 oW) in Graham Land, Antarctica.He was also active in the local alpine club, and amongother activities encouraged a Swiss expedition toGreenland, which in his honor gave the name MountForel to the highest peak that they encountered, oneof the highest in all of Greenland (lat. 66.93 oN, long.36.79 oW; 3383 m asl). FAF was also fascinated by earthquakes, and in collab-oration with the Italian geophysicist Michele de Rossihe developed the Rossi-Forel scale, a precursor to thenow familiar Richter scale for earthquake intensity. Atthe International Geophysical Conference in Zurichthere was a resolution to set up an InternationalEarthquake Commission, to which he was appointedvice-chair in 1907. Each year he continued to amassdata in the region including via appeals to the publicfor information, so much so that he was even knownlocally as the Director of Earthquakes.From his earliest days as a teenage amateur scientistwith his father, FAF was also interested in the peoplewho lived in the past around the lake, and he made anumber of important contributions in archaeology,and in the final two years of his life, he catalogued hisfather’s collection of bronze age artifacts. On top ofall this, FAF was also active in local governmentaffairs and held various positions on the MorgesCouncil and in the canton. He was the father of fourchildren : Cécile, Marie, Hilda and François. MmePorret-Forel tells us that her grandfather particu-larly enjoyed having his daughter Marie come out onthe boat with him during sampling expeditions. Byall accounts FAF was an affable and generous manwho greatly enjoyed the company of others andthrived on working and discussing his work with col-leagues in Switzerland and around the world. EvenAuguste Forel, who severely commented on manypeople he came in contact with, had nothing butpraise for his cousin : He was a keen scientific wor-ker and observer, and in later years he becamefamous for his important investigations of theLake of Geneva. He always signed his work ‘F.A.Forel’, so that at Morges he was known as ‘FAF’.Kindness and benevolence, together with a happyoptimism accompanied him throughout his life(Forel 1937 ; p. 83).



❚9. The F.A. FOREL legacy

François Alphonse FOREL (Fig. 7) was such a prolificscientist, author and scholar that it is difficult to sum-marize the full scope of his enduring contributions. Inthe aquatic sciences, however, his name is clearlyassociated with the birth of limnology. FAF concludesat the end his third volume (Conclusions and FinalReflexions; Forel 1904) that he is happy to payhomage to the geographic method that he hasemployed in his new science of limnology, an integra-tive approach that allows us to generalize, to aban-don specializations after having harnessed theirdiscoveries, and to contemplate the overviewmade up of the facts garnered from diverse disci-plines of science. (p. 669). However in these closingpages he also stresses the individuality of lakes, andwarns against simply extrapolating from his ownensemble of data at Lake Geneva, or uncriticallyapplying his approach : I have been led to theoreticaldissertation, discussion and development far inexcess of a simple statement of things. I do not reg-ret having so often succumbed to going beyond thepure objectivity of the facts to try to find the neces-sary theoretical explanations; but I cannot presentmy book as a model for the description of otherlakes (p. 668). He also notes how the quest to getcloser to scientific truths is an ongoing process thatprogresses rapidly from generation to generation, andhe describes at length the joy of doing science : a sim-ple fact well studied, a little hypothesis well sup-ported, can provide us with as much immediatepleasure as the grandest theories have given to aNewton, Darwin, Helmholtz or Pasteur (p. 673).

FAF logically combined the Greek word limne forlake (and also pond, swamp) with logos to create thenew word limnology, but was it the best choice?Today, limnology certainly extends from lakes intopools, ponds, swamps, mires, bogs and other wet-lands. In 1922, August Thienemann and EinarNaumann founded the International Association ofTheoretical and Applied Limnology (SocietasInternationalis Limnolo giae, SIL) and more broadlydefined Limnology as the study of inland waters,which now includes rivers and streams (e.g., Wetzel2001) and even estuaries (e.g., Horne and Goldman1994). Despite this increasing breadth, however,Limnology is still not widely known among the generalpublic. The limnologist Jonathon Cole, former presi-dent of the Association of the Sciences of Limnologyand Oceanography (ASLO), laments In some ways itis unfortunate that François Alphonse FOREL(1841-1912), the first to use the term limnology,was both conversant with ancient Greek, andunwilling to compromise linguistic purism bycombining perhaps a more widely recognizedLatin word for inland water with the Greek -logos(Cole 2009). But what exactly would that word be?The obvious alternatives are unattractive hybrids, andseem limited in meaning. At any rate, limnologyremains well established in the names of many of ourprofessional journals and societies, including ASLOand its publications.Most modern textbooks of limnology refer to FAF inpassing as the founder of limnology, and usually citethe three volume monograph on Lake Geneva (LacLéman). Few, however, acknowledge FAF’s publica-tion of the first textbook in limnology, entitled Hand-

buch der Seenkunde : AllgemeineLimnologie (Textbook of LakeScience : General Limnology).Forel wrote this in French at therequest of a German publisher, andthen asked his friend and colleagueDr. Wilcez at the University of Lau-sanne to translate it into German forpublication (Forel 1901). It wasreviewed for the Royal GeographicalSociety by H.R. Mills who read theGerman text soon after publicationand noted that the appearance of thishandbook of Limnology may belooked upon as the formal admis-sion of the science of lakes to itsindependent position somewhereon the borderlands of geography,very near the place occupied byoceanography. Prof. Forel is theone man capable of writing such abook… (Mills 1901). The book wasalso well received in the UnitedStates, where a review signed by

Fig. 7. François Alphonse FOREL (left) has been honored in many ways, includ-

ing by the founding of the Institute F.-A. FOREL (right) in his name at the

University of Geneva in 1980. This limnological institute is renowned for its

multidisciplinary studies on lakes (website : www.unige.ch/forel).

(Left : image courtesy of the Musée du Léman and the Forel family;

Right : image by W.F. Vincent).


R.D.S. in the Journal of Geology, a prestigious scien-tific journal published by the University of ChicagoPress, appeared the same year and stated : It bringstogether in concise, comprehensive and readableform the general principles of limnology…In anappendix is given an outline for the prosecutionof lacustrine studies, and also a bibliography. Thevolume is the best compendium on the subject...In surveying the current limnological literature, FAF’sthird volume (Forel 1904) seems largely unknown.Yet the concepts and approaches made in that volumeare vitally relevant to the application of limnologicalprinciples toward managing the world’s freshwaters.As described above, FAF saw human beings as a keycomponent of limnology, from two perspectives.Firstly, humans are part of the coupled lake-catch-ment ecosystem biota (his species list of lacustrinefauna began with Homo sapiens) and we dependupon lakes for reasons as diverse as safe drinkingwater, waste disposal, reliable food supplies and theaesthetic and psychological pleasure of being on andnear water. FAF would be pleased that in theEncyclopedia of Inland Waters, the article on theorigins and nature of limnology (Cole 2009) is fol-lowed by one on the aesthetic values of lakes andrivers, and then by another on aquatic ecosystemservices. More than 100 years ago, FAF not only iden-tified the many ecosystem services provided by LakeGeneva, he even quantified one of them, the fishery, inprecise economic units of Swiss Francs (Forel 1904). Secondly, FAF considered Homo sapiens to be themost powerful of all aquatic species, with a capacityto not only to benefit from lakes, but also to causethem great harm. His examples included the pollu-tion by cities and their sewage discharges, the waste

streams from industries, the pressure of overfishing,the release of coal ashes from boats and the introduc-tion of non-native plant and animal species (Forel1904). Once again his work underscored the interre-latedness of each component of the ecosystem, andhow the perturbation of one would ripple throughoutthe entire physical-chemical-biological-human sys-tem. Here now, in the 21st century at a time ofincreasing perturbation of the world’s freshwaters byclimate change, long range contaminants, speciesinvasions and multiple other stressors, it would seemto be a good moment to rediscover FAF’s greatestlegacy : his advocacy for an integrative approach inwhich each discipline informs the other, toward anoverall synthesis that includes humankind in thepast, present and future, and that informs policydecisions and environmental stewardship. If F.A.FOREL were here today to write a limnology ofglobal change, he would likely encourage us to payclose attention to the role of humans in the bio-sphere, and to understand the human presence as acomponent part of all aquatic ecosystems.

❚Acknowledgements

We thank Carlos Duarte, John Dolan, Michio Kumagaiand the Forel family for their support and encourage-ment of the preparation of this article, which was ini-tiated while WFV was a visiting fellow at the LakeBiwa Environmental Research Institute, supportedby the Japan Society for the Promotion of Science(JSPS). An abridged version of this article was pub-lished in the ASLO Bulletin (Vincent and Bertola2012).




Bibliographie

❚ BERTOLA C. 1999. François-Alphonse Forel (1841-1912), p. 3-17. In : Bertola, C., Goumand C., and Rubin J.F. (eds.), Découvrir le Léman : 100ans après François-Alphonse Forel. Actes du colloque pluridisciplinaire, Nyon 16-18 septembre 1998. Editions Slatkine, Geneva.

❚ BLANC H. 1912. Le professeur Dr. François-Alphonse Forel, 1841-1912. Actes de la Société helvétique des sciences naturelles, 95 : 110-148.❚ COLE JJ. 2009. Limnology as a discipline, p. 6-13. In : Likens GE (ed.), Encyclopedia of Inland Waters, Elsevier, Oxford, U.K.❚ DELEBECQUE A. 1898. Les lacs français. Chamerot et Renouard, Paris. 436 p.❚ EGERTON FN. 1962. The scientific contributions of François Alphonse Forel, the founder of limnology. Schweizerische Zeitschrift für

Hydrologie, 24 : 181-199.❚ FOREL A. 1937. Out of My Life and Work. W.W. Norton & Co., Inc. New York (First English Edition). 352 p.❚ FOREL F-A. 1873. Première étude sur les seiches. Bulletin de la Société Vaudoise des Sciences Naturelles, 12 : 213.❚ FOREL F-A. (ed.) 1879. Matériaux pour servir à l’étude de la faune profonde du Lac Léman. Collection du Bulletin de la Société Vaudoise

des Sciences Naturelles, 459 p. ❚ FOREL F-A. 1885. La faune profonde des lacs suisses. Mémoire de la Société helvétique des sciences naturelles, H. Georg, Basel, 234 p.❚ FOREL F-A. 1886. Le Lac Léman: précis scientifique. H. Georg, Basel, 76 p.❚ FOREL F-A. 1892. Le Léman - Monographie Limnologique. E. Rouge, Lausanne, Tome I, 543p.❚ FOREL F-A. 1895. Le Léman - Monographie Limnologique. E. Rouge, Lausanne, Tome II, 651 p.❚ FOREL F-A. 1901. Handbuch der Seenkunde : Allgemeine Limnologie. Bibliothek geographisches Handbücher herausgegeben von Prof. Dr.

Friedrich Ratzel. J. Englehorn, Stuttgart, 249 p.❚ FOREL F-A. 1904. Le Léman - Monographie Limnologique. E. Rouge, Lausanne, Tome III, 715 p.❚ HORNE AJ, GOLDMAN CR. 1994. Limnology. McGraw-Hill, New-York, 576 p.❚ HUTCHINSON GE. 1957. A Treatise on Limnology : Vol. 1.Geography and Physics of Lakes. Wiley, New-York,1016 p.❚ MERCANTON P-L. 1914. François-Alphonse Forel : Son œuvre en géophysique, p. 26-39. In : Discours prononcés à la cérémonie commémora-

tive à l’inauguration du monument de F.-A. Forel à l’aula du Palais de Rumine, à Lausanne le 29 novembre 1913. Société Vaudoise desSciences Naturelles (SVSN), Lausanne.

❚ MILLS HR. 1901. Prof. Forel on limnology. The Geographical Journal,17 : 296-298.❚ NEEDHAM JG, LLOYD JT. 1916. The Life of Inland Waters. The Comstock Publishing Company, Ithaca, N.Y. 438 p.❚ SECCHI A. 1866. Relazione dell’ esperienze fatte a bordo della pontificia pirocorvetta l’Immacolata Concezione per determinare la

trasparenza del mare. In: Cialdi A. Sul moto ondoso del mare, p.258, Roma.❚ STERNER RW. 2012. Raymond Lindeman and the trophic dynamic viewpoint. ASLO Bulletin, 21 : 37-51.❚ VINCENT WF, BERTOLA C. 2012. Forel’s limnology : From lake physics to ecosystem services. ASLO Bulletin, 21 : 69-77. ❚ WEISMANN A. 1877. Das Thierleben im Bodensee. Lindau.❚ WETZEL RG. 2001. Limnology. Lake and River Ecosystems. 3rd Edition, Academic Press, New York.1006 p.❚ WILDI W, MOSCARIELLO A, PUGIN A. 1999. Histoire glaciaire du Léman, p. 399-414. In : Bertola, C, Goumand C, and Rubin JF (eds.). Découvrir

le Léman : 100 ans après François-Alphonse Forel. Actes du colloque pluridisciplinaire, Nyon 16-18 septembre 1998. Editions Slatkine,Geneva.

❚ WILLIAMSON CE, SAROS JE, VINCENT WF, SMOL JP. 2009. Lakes and reservoirs as sentinels, integrators, and regulators of climate change.Limnology and Oceanography, 54 : 2273-2282.

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