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Linnaeus: King of Natural History Paul Adam' and Elizabeth May- ' School ofBiological, Earth and Environmental Sciences, University ofNSW, Kensington, NSW, 2052; ^ School ofBiological Sciences, University ofSydney, NSW, 2006 Adam, P. and May, E. (2009). Linnaeus: king ofnatural history. Proceedings oftheLinnean Societyof NewSouth Wales 130, 245-250. Linnaeus'legacywasfarmoreencompassingthantaxonomic. Wearguethat,whilethesystematicrecording of species remains fundamental to modem ecological concerns, Linnaeus also laid the foundation for other major areas of ecology, including comparative biogeography, plant demography, and comparative anatomy. Manuscript received 18 October2008, accepted forpublication 21 January 2009. KEYWORDS: Biodiversity, Linnaeus, natural history, natural philosophy, taxonomy. essential skills underpinningthe study ofbiodiversity INTRODUCTION were already in decline. (Biodiversity itselfis a word of recent origin - first coming to the fore with the Linnaeus is one of the towering figures in the publication ofWilson [1988].) history ofbiological science. He isrememberedtoday The taxonomic side of Linnaeus' achievements chiefly for his introduction ofthe binomial system of was outstanding. Whilehis sexual systemdidnotlong nomenclature and for his taxonomy. However, ifwe surviveasabasis forplantclassification,Linneaushad consider his skill as an observer ofthe relationships graspedthepotential forclassificationtobepredictive between the plants and animals he classified, and and 'natural', even ifhis particular approach had its their environment, he should also be regarded as limitations. He had recognized the importance ofthe one of the earliest practicing ecologists. He passed hierarchical approach and provided a nomenclatural on these skills to his many students, including those system that was functional and, importantly, had who travelled the world collecting and classifying practical application to the large number of new organisms and became known as the 'apostles'. species thatwere being discovered outside Europe. It In eighteenth-century Sweden, Linnaeus was was the first classification system that was accessible accorded high status, and this great respect continued to the non-specialist, with the work encapsulated in to hold internationally until the twentieth century. handbooks that were 'small enough to be carried into Lord Rutherford's throw-away comment that '(a)ll the field' (Koemer, 1999, p40). science is eitherphysics orstamp collecting'reflected Intheyear2007,wecelebratedthetercentenaryof the marginalization oftaxonomy and natural history Linnaeus' birth. It also happens to be the anniversary within science as technological advances in physics, of Georges-Louis Leclerc, Comte de Buffon, the chemistry and engineering attracted fimding and authoroftheHistoireNaturelle. Conniff(2007), inhis support. These advances also revolutionised biology, article aptly titled Happy Birthday, Linnaeus, argues permittingbreakthroughsinphysiology,biochemistry that Buffon should be regarded as at least the equal and the molecular sciences, but in this brave new of Linnaeus, and in particular suggests that Buffon world the diversity and distribution oforganisms lost had a superior understanding ofhabitat, anticipating their attraction as fields ofstudy. It was only towards the development of ecology as a science. Linnaeus the end ofthe twentieth century when environmental and Buffon were mutually fierce critics, and Buffon issuesbecame suchamajortheme inpolitics andwith was undoubtedly also a major figure in the history of the public thatthere was renewed interest in the study science:hehadbettergeologicalinsightthanLinnaeus, of biodiversity, and a need to bring new techniques and was closerto having an evolufionaryperspective. and approaches to 'old fields'. By then, many ofthe However, to suggest that Linnaeus' natural history LINNAEUS: KING OF NATURAL HISTORY was not ecologically focused is far from accurate. A the links to scientific botany, and ethnobotany has major part of Lirmaeus' teaching was based on field been given a new impetus as a field of study by the excursions: he had a very comprehensive knowledge regimeforrewardingtraditionalownersofknowledge of the local flora of southern Sweden, and while and resources, which was established by the United his sexual system was not appropriate for higher Nations Convention on Biological Diversity 1992. taxonomic ranks, Linneaus' species concepts have From the late 16* century onwards there was largely stood the test oftime. a considerable interest in collecting and studying We would argue that Linnaeus' taxonomic work 'curiosities' ofall kinds, and some ofthe collections was firmly underpinned by a deep understanding of natural history objects that were assembled were of natural history and that natural history in turn large. Some ofthe more academic natural historians provided the basis for ecology (Mayr, 1997; Blunt, associated with these collections were distinguished 1971). The ecological insights of Linnaeus are scientists whose work has stood the test oftime. An clearly seen in his botanical 'text book', Philosophia example is John Ray, who was the first to draw a Botanica (Linnaeus, 1751). Koemer (1999) notes that distinction between Monocots and Dicots and who is '...he described many ofthe mechanisms of species commemoratedinthe still existingRay Societyandin interdependence, as Charles Darwin noted on reading thenameoftheherbariumattheUniversityofSydney. his Oeconomia naturae of 1749' (pi5). Some ofthe impetus for collecting was stimulatedby the increasing numbers of exotic specimens being sentbackto Europe fromwiderexploration. Linnaeus LINNAEUS AND NATURAL HISTORY was very much part of this natural history tradition and although he did not travel beyond Europe, he 'Natural history', the advancement of which is actively encouraged his students to do so, and he was the prime objective ofthe various Linnean Societies familiar with non-European plants both in the form aroundtheworld,hasaverylonghistory. Earlyhunter- of herbarium specimens and in gardens. The non- gatherer societies could not have survived unless European species he described famously included members possessed what we might consider to be bananas, which would then have been regarded as an innate understanding ofnatural history, including very exotic. the ability to recognise different sorts of food and Linnaeus' own exploration was closer to home to distinguish between the edible, the toxic and the and included his early expedition to Lapland. dangerous. Cave paintings provide, in a tangible Although there are suggestions thathis account ofhis form, evidence for knowledge of natural history. travels is somewhat exaggerated' (Koemer, 1999), it In classical times, plants and animals were seen as established his reputation as an explorer and natural sources of medicines or as an element in a broader historian. Lapland in the eighteenth century was at natural philosophy, and the Greeks and Romans left a the edge ofthe world and formany Europeans would documentary record which at least in part survives to havebeenregardedwithasmuchtrepidationasAfrica. thisday andthatwouldhavebeenknowntoLinnaeus. Eventoday itremains one ofthe fewwilderness areas Knowledgeofnaturalhistorywouldhavebeencurrent in Europe (Ratcliffe 2006). amongst the broader population, the majority of Once he was established as a senior academic whichlivedinrural environmentsandwereintimately in Uppsala, field teaching became an essential and dependent on the natural world for survival, but in popular part of Linneaus' teaching. His excursions 'academic' circles natural history was increasingly attractedlargenumbersofsliidentsandwereorganized associated with medicine. For hundreds of years with almost military precision (Blunt 1971). Most herbalists recycled the writings of classical authors, attention was paid to the flora, although any matter without making original observations and with the of natural history interest was open for study and claims becoming more fancifiil on eachretelling. The comment. Many of the localities around Uppsala Renaissance then brought a new curiosity about the that were visited on excursions still support the same world andmore organized scientific inquiry, although species today, so thatitrequiresno great stretch ofthe the importance of the links to medicine continued, imaginationtovisitsitestodayandseewhatLinneaus' and old myths still retained currency. Linnaeus students wouldhave seen, andto experience the same himself was Professor of Medicine and Botany at excitement offirst encountering awetmeadow fiill of Uppsala University, and in some institutions close snakeshead fritillaries {Fritillaria meleagris) or a dry links between the two disciplines survived until the calcareous esker with a spring abundance of Pasque twentieth century. The recent growth of interest in flowers {AnemonePulsatilla). alternative medicine suggests a need for revitalizing 246 Proc. Linn. Soc. N.S.W., 130, 2009 ' ADAM AND MAY p. E. LINNAEUS AND ECOLOGY section 312), displaying evidence of very carefial observation. He also pointed out that variation could William Steam (in Appendix I of Blunt, 1971) be correlated with soil conditions and microclimate recognised that Linnaeus has been variously declared andadvocatedanexperimentalapproach{Philosophia 'a pioneer ecologist, a pioneer plant-geographer, a section 316: 'Cultivation is the mother ofvery many pioneerdendrochronologist, apioneerevolutionist... varieties and is the best means oftesting varieties'), but considered that the 'most influential and useful foreshadowing by a century and a half experimental of his contributions to biology undoubtedly is his taxonomy (genecology), which enjoyed its heyday in successfulintroductionofconsistentbinomialspecific the second halfofthe 20* century. nomenclature'. Chapter XI ofPhilosophia (entitled 'Sketches') ItistruethatLinneaus'contributiontoecologyand contains much ecological material. Section 334 'The plant geography is rarely acknowledged within these native locations of plants relate to region, climate, disciplines, and the recognized founding fathers were soilandground contains avery succinct introduction all much more recent. Nevertheless his Philosophia to ecology and biogeography (as well as some rather Botanica contains many ecological insights, which strange views about geology). The discussion about were in the published literature and were dormant the relationship between latitude and flora gives hints seeds formany decades. Giventhe very large number oftheideassubsequentlydevelopedingreaterdetailby of students who attended Linneaus' classes, and the Alexander von Humboldt. The relationships between wide circulation of his publications, the ecological soil types and the plants they support also introduce perspective he developed must have been assimilated topics that formed amajorpart ofecological research into the perceived wisdom of the day, and when in the twentieth century. Section 335 provides an ecology and plant geography developed as separate overview of phenology and indicates that Linnaeus disciplines, Linneaus' ideas would have been part of was well aware that factors such as temperature and the assumed background. day length were involved in controlling flowering, Today the major concerns of ecology include although itwastobemanyyearsbeforephysiological the identification and evaluation of biodiversity. Of understanding of the mechanisms involved was the three generally accepted levels of biodiversity, achieved. Even on botanical excursions students Linneaus was ignorant of genes, but he clearly recorded the plant species eaten by particular animal recognized the need to document species, and species 'while watching the botanical specimens recognized that species occupied habitats. In fact, he disappear at the moment they realized that they devotespartofthePhilosophiato discussingthemain needed to identify them' (Koemer 1999 p49). habitats (communities) in Sweden. He also indicated Chapter V of Philosophia (Sex) includes what notes should be made on field excursions. observations on annual seed production ofindividual Unfortunately, the details on many herbarium plants, probably the first scientific exploration of labels in current collections fail to provide any plant demography. The essential^ feature of the ecological information. Linnaeus' advocacy of the Philosophia is the importance of observation, and it systematic recording of habitat data was part of his is remarkable how much was achieved using lenses approachtocataloguinginformationandthesefeatures and microscopes that today would be regarded as are easily accommodated in modem databases. If woefully inadequate. Linnaeus were alive today, he would undoubtedly WhileLinnaeuswas acreationist,therecognition be active in the development of bioinformatics and ofvariation suggeststhathewasnotasrigidlyso ashe the creation and manipulation of databases. The is usually portrayed - he certainly recognized that the A omissions ofthe past cannot be corrected but today's appearance of species could change. synthesis of collectors shouldbe encouraged to recordmuchmore Buffon's and Lirmaeus' ideas could have accelerated than is often the case. Regrettably, ecologists are the development ofevolutionary theories, well ahead often amongst the worst offenders when it comes to a ofthe publications ofDarwin and Wallace in the mid lack ofdetail associated with voucher specimens. 19* century. Linneaus was well aware of the variability The Philosophia is also strongly focused on displayed by some species and devoted Chapter IX the utilisation of plants, not just as medicines but of the Philosophia to a discussion of 'Varieties'. for a whole range ofpurposes. The 365* (and final) He urged against giving taxonomic recognition to article states: 'The economic use ofplants is ofgreat environmentally determined phenotypic variation, utility to the human race.' (Linnaeus, 1751). One of as he recognized that a variety ofdiseases and insect the majorjustifications for biodiversity conservation attackcouldcauseabnormalitiesinplants{Philosophia is the maintenance of the ecosystem services that Proc. Linn. Soc. N.S.W., 130, 2009 247 LINNAEUS: KING OF NATURAL HISTORY biodiversity supports. This is a concept that would This tradition of the extremely skilled amateur was clearly have found favour with Linneaus, and the never as strong in New South Wales as it was in the sorts of observations he advocated are needed to UnitedKingdom (forreasons thatperhaps requirethe document ecosystem processes. He recorded details attention of a social historian). Certainly, there have ofthetrophic interactionsbetweenorganisms andhad been some very gifted amateurs, but their interests an appreciation ofthe recycling ofmaterials, noting tended to be restricted to groups such as birds or people used churchyard soil for growing cabbages, flowering plants; there was not the same interest or hence 'human heads ... turn into cabbage heads' expertise shown in, for example, cryptogamic plants (Koemer, 1999, p83). as was the case in the United Kingdom. Given the Robert MacArthur (1972) famously wrote that dearth of professionals in Australia, this means that 'to do science is to search for repeated patterns' there are major components of biodiversity about and stressed the importance ofnatural history as the whichwe remain still basically ignorant. starting point for ecological research. MacArthur Today collection is frowned upon, and in many pointed out that not every natural historian was cases(suchasthecollectingofbirdeggs) itisproperly a scientist (in terms of approach and method, not illegal. The shift away from collection partly reflects necessarily profession) and not all ecologists were lack of opportunity given an increasingly urbanized natural historians, but we would agree with him that population, rejection ofa 'stamp collecting' approach most ofecology has its roots in natural history: even to science and greater concern for conservation. theoreticalmathematicalecologystartswithideasthat Certainly, it would not be possible to condone egg- are ultimatelybased onfieldobservation. Underwood collecting or capturing and killing native vertebrates (2007) has recently observed 'one of the great joys outside specially-approved scientific licences, but of experimental ecology is that natural history is nevertheless it is probable that both school and so important in the development of explanatory universitystudentsaremissingoutonwhatpreviously models'. had been important educational experiences. The old collections remain of continuing value, providing comparative material for taxonomic COLLECTING BIODIVERSITY: PRESERVING studies, as well as evidence ofchanging distributions BIODIVERSITY? or ofenvironmental change. For example, the ability to measure long-term trends in thickness of the In the eighteenth, nineteenth and early twentieth shells of raptor eggs was extremely important in century much natural history involved collections; drawing attention to the effects ofthe new post-war and many large collections of, for example, insects, organic agricultural chemicals (Ratcliffe 1967, 1970; bird eggs, or plants were made both by, or for, major Olsen and Olsen 1979). Sadly, curation ofhistorical institutions and individual collectors. Many people collections in some ofour greatest museums is being who subsequently became famous scientists in other eroded as funding for the care of collections, in the fields (for example Macfarlane Burnet - Sexton management schemes of modem-day directors, 1999) were avid collectors in their youth. Charles is coming a distant second behind promotion of Darwin himself was an avid beetle collector in his exhibitions for entertainment's sake without the college days (preferring 'beetling' to mathematics underlying scholarship being obvious. - Desmond and Moore 1991). (Another suggestion Even in the absence of collections, skilled for Linnaeus' mis-representation of his travels in amateurs in the United Kingdom have been able Lappland was mis-calculation (Selander, 1947 in to systematically record distributions of large Koemer, 1999). Perhaps he sharedDarwin's aversion numbers oftaxa at the national scale. These spatially to mathematics?). The making of collections taught explicit data are of enormous value for monitoring theneedforcarefial observation, systematicrecording environmental change. Dataofthis sortwouldbevery of data, and provided in-depth understanding of difficult to collect (and inordinately expensive) ifwe particular groups oforganisms. had to rely on professionals, yet they will be crucial Collectors and recorders were notjust the clergy to our monitoring of biodiversity. In Australia the andthe landedgentry (ortheirspinstersiblings); there omithologists have pioneered systematic recording was, at least in the United Kingdom, a very strong of species distribution at the continental scale, working class element ofminers and factoryworkers, hamessing the skills and enthusiasm ofamateurs and who, in their very limited free time, spent many professionals. The differences in distribution and hours completing arduous hikes and making major abundance between the two editions ofthe BirdAtlas finds of often taxonomically-challenging organisms. (Blakers et al. 1984; Barrett et al. 2003) provides 248 Proc. Linn. Soc. N.S.W., 130, 2009 ADAM AND MAY p. E. compellingevidencefortheimpactsofenvironmental REFERENCES change, and the recording program will continue into the future. Other areas of natural history have not Barrett, G.W., Silcocks,A.F., Barry, S., Cunningham, R.D. been so well served. and Poulter R. (2003). 'The new atlas ofAustralian Teaching in thefield is still acomponentofmany birds'. (BirdsAustralia [RoyalAustralasian Universitycoursesbutisundergreatpressurebecause Omothologists Union]: Hawthorn East, Victoria) ofcost, occupational health and safety issues (which Blakers M., Davies S.J.J.F and Reilly, RN. (1984). can createbureaucraticnightmares), the largenumber 'The atlas ofAustralian birds'. (RoyalAustralasian Ornithologists Union and Melbourne University ofstudents with part-timejobs who find it difficult to Press: Carlton, Victoria) attend courses at weekends or during vacations, and Blunt, W. (1971). 'The CompleatNaturalist. ALife of becauseofthedeclineinthenumberofacademic staff Linnaeus'. (Collins: London). with knowledge of many groups of organisms. The Conniff, R. (2007). Happy Birthday, Linnaeus. Natural long-termfiitureoffieldteachingisveryuncertainand History 115: 42-47 this will have major consequences for our ability to Desmond,A. and Moore, J. (1991). 'Darwin'. (Michael produce graduates capable ofaddressing biodiversity Joseph: London). Koemer, L. (1999). 'Linnaeus: Nature andNation'. issues. Peter Marren (2002, 2005) has written on (HarvardUniversity Press: Cambridge, Massachusetts). a number of occasions, pointing out the loss of UK Linnaeus, C. (1751). 'Philosophia Botanica'. (Translated expertise in institutions and the decline in the by Stephen Freer.) (Oxford University Press: numbers of skilled amateurs. At the same time Oxford). NGO there is an increase in the membership of MacArthur, R.H. (1972). 'Geographical ecology: patterns conservation societies, indicating wide support for inthe distribution ofspecies'. Harperand Row: New natural history, but the deep engagement with some York. particularfieldwithin natural history is less common. Mayr, E. (1997). 'This is Biology. The Science ofthe There may be many and varied reasons for this, but LivingWorld'. (Belknap-HarvardUniversityPress: pMraersrseunressuogngemstasnyonpeeompaley,baendthatth,e waibtshentchee omfotdimeem, MarreCna,mbP.ri(d20g0e2,).Ma'sNsaatcuhrueseCtotnss)e.rvation -AReview of the Conservation ofWildlife in Britain 1950-2001'. natural history had become a spectator rather than (HarperCollins: London). praercteinctiTpaVntnsaptourrta.lThhisetoqruyalpirtoygarnadmsexipsesrutcishethsahtorwantheirn MarreBrnit,iPs.h(n2a0t0u5r)a.l'hTishteorNy.ew2nNdatEudriatliiosnt's.-(hHaalrfpaerceCnotlulriynso:f than encouraging participation they suggest that we London). already know everything. This is an idea that needs Olsen, P. and Olsen, J. (1979). Eggshell thinning in the further exploration. peregrine, Faicoperegrmus (Aves: Falconidae), in The achievements of Linneaus and his Australia. Australian WildlifeResearch 6, 217-226. Ratcliffe, D.A. (1967). Decrease in eggshell weight in students were remarkable and the detail of their certain birds ofprey. Nature (Lond) 215, 208-10. observations made with minimal technological aids, Ratcliffe, D.A. (1970). Change attributable to pesticides was particularly remarkable. Few students today in egg breakage frequency and eggshell thickness in would have the capacity or patience to make similar some British birds.JournalofAppliedEcology 7, investigations. Experimental science needs to be 67-115. underpinned by substantial bodies of observation in Ratcliffe, D.A. (2006). 'Lapland: anatural history'. (Yale order that appropriate hypotheses can be generated University Press: New Haven, Connecticut). and tested. Lirmaeus' legacy ofobservation has been Selander, S. (1947). Linne i Lule Lappmark. Svenska built upon for the last 250 years, but the capacity to LinnesdllskapetsArsskrift. XXX, 9-20 Sexton, C. (1999). 'Burnet, a life'. (Oxford University continue to do so is being lost. Press: Melboume). Contrary to the impression left by Conniff, we Underwood,A.J. (2007). Logic, design, analyses, wouldarguethatLinnaeus also laidthe foundation for interpretations: essentials ofecological other major areas of ecology, including comparative experimentation. British EcologicalSocietyBulletin biogeography (long before van Humbolt), plant 38, 24-27. demography, and comparative anatomy. His legacy Wilson, E.O. (ed) (1988). 'Biodiversity' (National was far more encompassing than taxonomic, even Academy Press: Washington DC). though the systematic recording of species remains absolutely fundamental to modem ecological concems. Proc. Linn. Soc. N.S.W., 130, 2009 249 LINNAEUS: KING OF NATURAL HISTORY (Endnotes) ' He variously reported traveling up to three times further than he really did and spoke of long periodsspentwith the native Sami, when infact it was afew weeks. Thefact that he was beingpaid by the milefor hisjourney by the Science Society ofUppsala is cited by way ofpartial explanation (Koerner 1999). 250 Proc. Linn. Soc. N.S.W., 130, 2009

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