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HAECKEL, DARWIN, AND ECOLOGY

BY ROBERT C. STAUFFER
University
ofWisconsin

T INTRODUCTION

bya
H HE TERM ecologywas invented
theoreticalmorphologist who was not
1856, p. 124). In the autumn of 1859, however,
whenDarwin's Originwas published,Haeckel was
far fromthe worldof scientificnews. He was at
a distinguishedstudentof ecologyand Messina on the straitsbetweenSicily and Italy,
who was outspokenlyhostileto one of wherehe was followingthe exampleofhis beloved
the most significantprogramsof eco- teacher,JohannesMiiller(1801-1858),in studying
logical researchof his day, the controversial but the richesof the sea to be capturedwith the col-
pioneering ventureto establishplankton lecting net. Using the methods which he had
influential
studieson a quantitativebasis. This paradoxofthe learnedfromMiillerat Helgolandin 1854 (Haec-
kel, 1852-1856,p. 251), Haeckel obtained many
positionof ErnstHaeckel (1834-1919) in the his-
new species of Radiolaria to delighthis artist's
toryof ecologyposes the problemforthispaper.
Haeckel's major contributions may be best under- eye. The resultsof thesemonthsof collectingand
stood as an immediatebyproductof his aim to study at Messina he broughtnorthto formthe
interpret Darwin's thoughtforthe scientific world foundationfor his masterpiece-his first huge
ofGermanyratherthanas thelogicaloutgrowth of monograph,Die Radiolarien,whichwas to secure
him his appointmentas ausserordentlicher Pro-
his own research.In his workas a fieldnaturalist,
Haeckel showeda painter'seye forthebeautiesof fessorin the medicalfacultyat Jena at the age of
28. It was thusonlylater,whilehe was working
nature,and he was an ardentcollectorwhosetrav-
up his Italian collections,that Haeckel became
els rangedfromNorway,Helgoland,the Canaries,
familiarwiththeOriginofSpecies.
and Messinato theRed Sea, Cylon,and Java. This
This encounterwas to be a major influenceon
wide and vivid experiencebroughthim some ap-
Haeckel's personal beliefs and on his scientific
preciationofecologicalpointsofview,but Haeckel
career."Aftermy returnfromItaly in 1860 when
did not display any notable insightinto the dy-
I became acquaintedwithDarwin's work,whenI
namicprinciplesofecology.My purposehereis to
saw in it the way preparedfor a mechanisticor
examinethenatureand thepersonallimitationsof
Haeckel's contributions to the science he named monisticsolutionofthemostdifficult philosophical
and to demonstrate its Darwinianderivation. problems,thentherebegan to developwithinme a
unifiedview of lifein theface of whichonlya few
HAECKEL AND DARWINISM of my long-accustomedand beloved articles of
faithcould endureany longer" (Haeckel, 1874,p.
Ecology was introducedinto the vocabularyof
333). In his Radiolarien,Haeckel used Darwinian
sciencein 1866 in Haeckel's Generelle
Morphologie.
This was the firstof a numberof books which conceptsin his discussionof the relationsof the
Haeckel wroteto forwardthe cause of Darwinism familiesof this order,in his considerationof a
as he saw it,and we muststartwitha glanceat the genealogicaltable, and his speculationsregarding
generalbackgroundof Haeckel's earlyenthusiasm the originalprimitivetypeformof the order;and
forCharlesDarwin (1809-1882). he emphasized the evolutionarysignificanceof
Even as a medicalstudentat Wiirzburgin 1853, transitionalforms.Here Haeckel expressedhis
in writinghome to proposebooks forfamilyva- enthusiasmforthe stimulatingideas of the Origin
cation reading,Haeckel had been appreciativeof of Species and explicitlydeclared his beliefthat
the "excellenttravelsof Darwin" (Haeckel, 1852- Darwin's theories"had beguna new epochforthe
138

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HAECKEL, DARWIN, AND ECOLOGY 139

systematicscientificstudy of organic nature" self;theinfluence ofotherearlierwritersseemsless


(1862, pp. 231-32). As "incontestablyimportant immediateif not problematical.
principlesof the greatestsignificance,"Haeckel If we searchthe Originforecologicalideas, the
mentionednot only natural selection and the mostrewardingchaptersare thoseon the struggle
strugglefor existence,but also the mutual rela- forexistenceand on naturalselection,and herethe
tions of organisms(pp. 231-33). The year after most generaland fundamentalecologicalconcept
makingthisinitialdeclaration,whenHaeckel went is that whichDarwin usuallydesignatedeitheras
up to Stettinforthe 1863 Versammlung deutscher the "economyof nature" or as the "polityof na-
Naturforscher und Aerzte,he threwhimselfinto ture." Now, among the Linnaean dissertations
the campaignforDarwin by devotinghis speech publishedin the Amoenitates Academicaeare both
at the firstgeneralsessionto Darwin's theoryof an Oeconomianaturaeand a Politia naturae,and
evolution(Haeckel, 1863,1883). Darwin could veryeasilyhave derivedhis concept
The verytitleof Haeckel's nextbook, Generelle directlyfromthe Linnaean schoolsimplyby sub-
Morphologie derOrganismen. Allgemeine Grundziige stitutingthe principle of natural selection for
der organischenFormen-Wissenschaft, mechanisch Linnaeus' pious beliefin Providenceas the direct
begriindetdurchdie von CharlesDarwin reformirtesourceoftheequilibriumofspeciesand theadapta-
Descendenz-Theorie (Berlin,1866), showsthat this tionsdiscoverablein the orderof nature.Darwin
two-volumework was another episode in the gave thecats-to-clover chainas "one moreinstance
Darwiniancampaign.The generalaim and charac- showinghow plants and animals,most remotein
terof theworkcan be judged fromHaeckel's own the scale of nature,are bound togetherby a web
comments:(1873, p. [xiii]) "a somewhatcompre- of complexrelations"(p. 63), Today we wouldre-
hensivework,whichconstitutedthe firstattempt gard thiseconomyor polityof natureas a system
to apply the generaldoctrineof developmentto in dynamicequilibrium;foras Darwinsaid, "in the
the wholerangeor organicmorphology(Anatomy long-runtheforcesare so nicelybalanced,thatthe
and Biogenesis),and thusto make use of the vast face of natureremainsuniformforlongperiodsof
marchonwardswhichthe geniusof CharlesDar- time" (p. 63). Yet, in "the case ofa countryunder-
win has effectedin all biologicalscience by his going some physical change,for instance,of cli-
reformof the Descent Theory and its establish- mate. The proportional numbersof its inhabitants
ment throughthe doctrineof selection.At the would almost immediatelyundergo a change"
same time, in the 'Generelle Morphologie' the (p. 70). Or, as Darwin had expressedhimselfin his
firstattemptwas made to introducethe Descent firstsketchofhis speciestheory,theessayof 1842,
Theory into the systematicclassificationof ani- "let any singularchangeofclimate(occur)here(?),
mals and plants,and to founda 'naturalsystem' how astoundinglysome tribes(?) increase ... the
on the basis of genealogy;that is, to construct pressureis alwaysready. . . a thousandwedgesare
hypotheticalpedigreesfor the various species of beingforcedinto the oeconomyof nature" (1842-
organisms."Thus Haeckel intendedto presentnot 1844,pp. 7-8).
only a broad synthesisof morphologybut also a The economyof nature has places or stations
surveyof biologyfroma Darwinianpointof view, whichare involvedin thestruggleforexistenceand
fortheintroduction
and thelogical justification of natural selection; "for as all organic beingsare
ecology comes simply as the demarcationand striving,it may be said, to seize on each place in
namingof one of many notable aspects of Dar- the economyofnature,ifany one speciesdoes not
win's thought. becomemodifiedand improvedin a corresponding
degreewithits competitors, it will soon be exter-
THE DARWINIAN CONCEPT OF ECOLOGY minated"(1859,p. 87). Here Darwindevelopedan
The elementsofHaeckel's conceptofecologyare idea he had expressedin writingabout coral reefs
to be foundin Darwin's Originof Species (1859). (1842), that "no station capable of supporting
It is truethat othersourcespossiblyrelevantbut lifeis lost. . . thereis a struggleforeach station,
certainlymore remotemay be recognized.Some betweenthe different ordersof nature" (p. 62).
of these Professorvan der Klaauw (1936) has But on the otherhand, "the morediversified the
discussedin his extensiveand informative study descendantsfromany one speciesbecomein struc-
of thehistoryof thedefinitionsof ecology.But the ture,constitution,and habits,byso muchwillthey
importanceof Darwin is stressedby Haeckel him- be better enabled to seize on many and widely

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140 THE QUARTERLY REVIEW OF BIOLOGY

diversified places in thepolityofnature,and so be ternalrelationsof organismsto each other,etc.


enabled to increasein numbers"(Origin,p. 98). "-Oecologie . . . derWissenschaft vonderOecono-
Darwin explainedadaptationto the environment, mie, von der Lebensweise, von der ausseren
both living and non-living,in writingthat "the Lebensbeziehungender Organismenzu einander
expressionof conditionsof existence,so oftenin- etc." Only hundredsof pages fartheron, in the
sistedon by theillustrious Cuvier,is fullyembraced latterpart of his secondvolumewhenhe came to
by the principleof naturalselection.For natural discussDarwin's conceptof evolutionand of nat-
selectionacts by ... adaptingthevaryingpartsof ural selection,did Haeckel begin to present a
each beingto its organicand inorganicconditions fullerconsideration of the term.Firsthe discussed
of life.. . ." (pp. 176-77; see Cuvier, 1817, p. 6; it brieflyas the scienceof the economyof nature,
1835,pp. 50, 53). of the mutual relationsof organisms;and he de-
To proceed as we have done, by considering scribedDarwin's cats-to-clover chain as an illus-
selectedecologicalconceptstaken fromDarwin's tration(II, pp. 235-36). Then fifty pages laterhe
presentationof his views on the originof species, presentedthe followingformalaccount:
admittedlyignoresDarwin's emphasis and the
originalsequenceof his ideas, but it does not do Ecologyand Chorology
any violenceto his perceptionof biologicalprob- In thepreceding sectionswe have pointedout re-
lems and relationships.We may summarizeour peatedlythatwithoutthe theoryofevolutionall the
findingsas follows:Darwin's most fundamental big generalseriesofphenomena oforganicnaturere-
ecologicalconceptwas that of the economicsand main completely incomprehensible and inexplicable
riddles, whileby meansof this theorytheycan be
sociologyof organismsas a system-the economy
explainedsimplyand consistently.(') This holdsespe-
orpolityofnature.Withinthissystemoccurplaces
ciallytruefortwocomplexes ofbiological phenomena
in the economyof nature open to different or- whichwe nowin conclusion wishto singleoutin a few
ganisms accordingto theirhabits of life and to words.Theseformthesubjectoftwospecialbranches
theirspecificadaptationto theconditionsof exist- ofphysiology whichso farhavebeenlargely neglected,
ence. We can let Darwin conclude in the terms namely, theecologyandchorology oforganisms.(2)
which seem to have been most influentialon
[Haeckel'snotesl(1) This tremendous mechanico-
Haeckel: "Let it be bornein mindhow infinitely causal significance oftheevolution theory forall biol-
complexand close-fitting are the mutualrelations ogy,andinparticular forthemorphology oforganisms,
of all organicbeings to each otherand to their cannotoftenenoughnorinsistently enoughbe heldup
beforeits unthinking or blindlydualisticopponents,
physicalconditionsof life" (1859, p. 69; cf.pp. 3, whoseteleological dogmatism retainsitsstrength only
95, 115, 267). becausetheycannotelucidatethesebiggeneralseries
ofthephenomena oforganic natureat all.
IAECKEL AND ECOLOGY (2) OtLKOs, 6 household or housekeeping, livingrela-
tions;Xc&pa, lodwellingplace,distributional area.
If we turnto Haeckel's discussionof ecologyin
his "Darwin-Buch," his GenerelleMorphologie By ecology, we meanthewholescienceoftherela-
oftheorganism to theenvironment including, in
(see Haeckel, 1852-1919,p. 67) we findthat not tions
the broad sense,all the "conditionsof existence."
onlyare reflections of thesestatementsof Darwin in nature;
These are partlyorganic, partlyinorganic
clearlydiscernible,but also this topic occursas a both,as wehaveshown, areofthegreatest significance
logicalpartofthediscussionofthetheoryofevolu- fortheformoforganisms, fortheyforcethemto be-
tionand the theoryof naturalselection.The asso- comeadapted.Amongtheinorganic conditions ofex-
ciationwithDarwin could scarcelybe closer. istenceto whicheveryorganismmust adapt itself
When Haeckel introducedecology,he discussed belong, first ofall,thephysical andchemical properties
it as an area ofbiologyand classifiedit as oneaspect ofitshabitat,theclimate(light,warmth, atmospheric
of the physiologyof relationships, but he did not conditions of humidity and electricity),the inorganic
defineit in any cut and driedfashion.Thus in dif- nutrients, natureofthewaterandofthesoiletc.
As organicconditions of existencewe considerthe
ferentpartsofhis twovolumestheemphasisshifts
entirerelations oftheorganism to all otherorganisms
and different aspectsof the fieldare stressed.The
withwhichit comesintocontact,and ofwhichmost
termecologyfirstappeared in a footnoteon page contribute eitherto its advantageor its harm.Each
eightof thefirstvolume.Here Haeckel introduced organism has amongthe otherorganisms its friends
it as a substituteforan arbitrarily restricted usage and its enemies,thosewhichfavorits existence and
of theterm"biology"and explainedecologyas the thosewhichharmit. The organisms whichserveas
scienceof the economy,of the habits,of the ex- organic foodstuff forothersorwhichliveuponthemas

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HIAECKEL, DARWIN, AND ECOLOGY 141

parasitesalsobelongin thiscategory oforganiccondi- After1866,Haeckel continuedto make passing


tionsof existence. In our discussion of the theoryof references to ecology.Duringtheinaugurallecture
we haveshownwhatenormous
selection importance all in the philosophicalfacultyof Jena whichhe de-
theseadaptiverelations haveforthe entireformation liveredin January1869,he again discussedecology
oforganisms, and specially howtheorganicconditions
in surveyingthe fieldof zoologicalscience.Here
ofexistence exerta muchmoreprofound transforming
The extra- he made the most quotable yet representative of
actionon organisms thando theinorganic.
ordinary significance oftheserelations doesnotcorre- his statements on its nature: "By ecology we mean
spondintheleasttotheirscientific treatment, however. the bodyof knowledgeconcerning the economyof
So farphysiology, [thescience]to whichthisbelongs, nature-the investigation of the total relationsof
has,in themostone sidedfashion, almostexclusively the animalbothto its inorganicand to its organic
investigated the conserving functionsof organisms environment; including,above all, its friendly and
(preservation oftheindividual and thespecies,nutri- inimicalrelationswith those animals and plants
tion,and reproduction), and amongthe functions of with which it comes directlyor indirectlyinto
relationship [it has investigated] merelythosewhich contact-in a word,ecologyis thestudyofall those
areproduced by therelations ofsinglepartsoftheor- complex
interrelations referred to by Darwinas the
ganismto each otherand to thewhole.On theother
of conditions ofthe struggle for existence." (Although
hand,physiology has largelyneglected therelations
theorganism to theenvironment, theplaceeachorgan- thistranslation, givenon p. [v] in thePrinciplesof
ismtakesin thehousehold ofnature,in theeconomy Animal Ecologyby W, C. Allee et al., is a free
ofall nature,and has abandonedthegathering ofthe ratherthan a literalrendering, I considerit very
relevantfactsto an uncritical "naturalhistory," with- faithfulto Haeckel's meaning,and so I quote it as
out makingan attemptto explainthemmechanisti- independentcorroborationof my interpretation.
cally. For the originalpassage see Haeckel, 1869,p. 17;
Thisgreatgapin physiology willnowbe completely fora veryliteraltranslationsee Haeckel, 1883,p.
filledbythetheory ofselection and thetheoryofevo- 200).
lutionwhichresultsdirectly fromit. It showsus how This accountof ecologywas givenin thelecture
all the infinitely complicated relationsin whicheach
of Haeckel's which was mentionedexplicitlyby
organism occursin relationto the environment, how
thesteadyreciprocal actionbetweenit and all theor- John Scott Burdon-Sanderson in his presidential
ganicand inorganic conditions ofexistence arenotthe addressto theBritishAssociationfortheAdvance-
premeditated arrangements of a Creatorfashioning mentof Science (1893). And it was in this latter
natureaccording to a planbutarethenecessary effects widelydistributed addressthatBurdon-Sanderson
of existing matterwithits inalienable propertiesand publicizedecologyby recognizingit as an aspect
theircontinualmotionin timeand space. Thus the of biologyequal in rankwithphysiologyand mor-
theory ofevolution explainsthehousekeeping relations phology-and of thesebranches"by far the most
of organisms mechanistically as the necessaryconse- attractive"(p. 465).
quencesofeffectual causesand so formsthemonistic Three other brief referencesto ecology by
groundwork ofecology.Now just thesamethingalso
Haeckel occurredin writings ofhis whichwerealso
holdstrueforthechorology oforganisms....
(Haeckel,1866,II, pp. 286-87) translated into English and so are also involvedin
the historyof the termin our own language. Of
[Throughoutthis translationI have departed these,two werein popularbookswhicheach went
froma literalrenderingto the extentthat seems througha numberof editions in England and
necessaryto produceclear and tolerableEnglish. America:The Historyof Creationand The Evolu-
I have translatedHaeckel's phrase Descendenz- tionofMan, (see indexesunderoecologyor oekol-
Theorieas "theoryof evolution"because Haeckel ogy). A thirdreference in Englishappearedin the
had alreadyequated theseexpressionson p. 148 of translationof Haeckel's Plankton-Studien pub-
Vol. II. We can justifytranslatingNaturhaushalt lishedin the Reportof the U. S. Commissioner of
as "economyofnature,"forin theGermantransla- Fish and Fisheries.
tion of the Originof Species used by Haeckel, H.
QUANTITATIVE METHODS
G. Bronn used Haushalt der Natur or Natur-
Haushaltas equivalentsforDarwin's "economyof These Plankton-Studien, firstpublishedin 1890,
nature"-and forhis "polityofnature,"also. It is offerus a revealinginsightinto Haeckel's under-
a pleasureto acknowledgethe advice of Professor standingof ecologicalpoints of view. Here the
R.-M. S. Heffner, whosesuggestionshave been of significant aspect is Haeckel's attitudetowardthe
greatvalue in improvingthis translation.] effortsby VictorHensen (1835-1924) of Kiel to

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142 THE QUARTERLY REVIEW OF BIOLOGY

introducequantitativemethodsinto the studyof Hensen, 1891), and these can be dismissedas


marinebiologyin orderto attack the problemof regrettableaspectsof Haeckel's polemics.But the
estimatingtheproductivity ofthesea. Even popu- most revealingpart of his objectionsinvolveshis
lar writinghad mentionedthe importanceof for- suspicionof quantitativestudies as a matterof
aminiferaand diatomsin the economyof the sea principle."Statisticsin generalis wellknownto be
for decades (Hartwig,1861, chap. 17); the value a veryhazardoussciencebecause it is usuallyem-
of Hensen's approach lay in his aim to discover ployedto findthe approximateaveragevalue of a
the orderof magnitudeof theirrole in food pro- magnitudefroma numberof imperfectobserva-
duction. tions" (Haeckel, 1891a,p. 322).
As a firststep towardthisgoal, Hensenplanned Haeckel had spokenof the economyofnaturein
to concentrateon the basic food-stuff such as the 1866 whenhe was firstdiscussingecology,and in
diatoms and copepods whichformmuch of this this discussionof planktologyhe cited as ecologi-
assemblageof floatingmarinelifeforwhichhe in- cal problems"the interestingand involvedvital
ventedthe term"plankton"(Hensen, 1887,p. 1). relationsof pelagic organisms,theirhabits of life
To make a start he boldly assumed for normal and economy"(p. 250), but he could not appre-
planktondistributiona uniformity which is still ciate Hensen's aim to make this study of the
being debated (Hardy, 1955). It was on this as- economyof the sea a quantitativeone. He re-
sumptionthat he based his estimatesof plankton jected Hensen's discoveryof the greatabundance
populationsfordifferent areas; forhe made these of planktonin colder waters because it did not
estimatesby extrapolation fromcarefullycounted agree with his own subjective impressionsand
samples.These sampleswereobtainedby filtering qualitative observations.Though he denied it,
a water column of a calculated cross-sectional he was apparentlymisledin part by allowinghis
area by makinga verticalhaul of his specially attentionto be concentratedon the richnessof
designednet. It was the countsof these samples the tropicsin numberof species as opposed to
which led Hensen to the unexpectedconclusion numberof individuals(pp. 267, 303-308). In op-
that the quantityof planktonis greaterin the posingHensen's methodsHaeckel arguedthat the
colder waters of the northernseas than in the employmentof "exact or mathematicalmethods
warmerwatersof the tropics. is impossiblein mostbranchesof
. . . unfortunately
On thebasis ofhis owncollectingexperienceand science (particularlyin biology),because the em-
his interpretation of the collectionsof the great pirical foundationsare much too imperfectand
Challenger Expedition (1872-1876), for which the present problems much too complicated.
Haeckel had describedthe deep-seamedusae,the Mathematicaltreatmentof thesedoes moreharm
siphonophores,and the radiolaria,Haeckel did than good because it gives a deceptivesemblance
not hesitate to attack Hensen's methodsand to of certaintywhichis not actuallyattainable.Part
questionhis aims. Part of the objectionsHaeckel of physiologyalso involvesproblemswhich are
raised againstHensen's assumptionsof a uniform difficult or impossibleto resolveexactly,and these
planktondistribution can easily be understoodas include the chorologyand ecologyof plankton"
due to skepticismbased on personalexperience, for (Haeckel, 1891a,p. 331).
Haeckel (1891a, pp. 308-18), like Darwin (1839, In regardto the "metabolismof the sea" (Stoff-
pp. 14-20; cf. pp. 189-90) and many other ob- wechseldes Oceans),Haeckel's criticism, ifshrewd,
servers,had been struck by the appearance of was bothpartialand unhelpfulwhenhe remarked,
dense local accumulationsof planktonwhicheven "To me it seemswhollyindifferent whetheror not
had visibly distinct boundaries. Here Haeckel the numberof thesebillionsis indicatedto us by
seems to have been misledbecause he overrated quantitative plankton analysis. To understand
the total effectof spectacularsituationsand un- theirphysiologicalsignificance, it would be more
derrated the inconspicuous conditions which importantto determinethe speed of their in-
provedto be statisticallyfarmoresignificant. But crease" (p. 329). Haeckel continuedwith a few
this involvesa technicalthoughmost interesting generalcommentsfavoringa subjectiveevaluation
controversy whichcan hardlybe settledby non- of this "metabolic"cycle,and thenhe conduded
specialists. Anotherpart of Haeckel's plausible by suggestingwith greaterenthusiasmthat "the
argumentsturnsout to be based on misinterpre- watery transparencyof the pelagic fauna, the
tationof Hensen's procedureor his reasoning(cf. periodicappearanceof manypelagic organismsin

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HAECKEL, DARWIN, AND ECOLOGY 143

swarms,theirabundant accumulationin 'zo6cur- progressof science, as formerlyin the case of


rents,'theirrelationto ocean currentsin general, geology,and morerecentlyin thatof theprinciple
are onlya fewof the manygreatproblemswhich of evolution" (quotations are from the end of
planktologyoffers to theresearchimpulseofman" Darwin's Introduction).
(p. 334). In thewholeplanktoncontroversy, while
CONCLUSION
Haeckel may have been correctin importantmat-
tersofdetail,he persistentlyobscuredthepotential Haeckel's original presentationof the term
value inherentin Hensen'sprogram. "ecology" in 1866 embodiesconceptsthat come
But it is timeto contrastHaeckel's suspicionof straightfromthe Originof Species. His contribu-
Hensen'squantitativemethodsin planktonstudies tion to the scienceof ecologywas made in his role
with Darwin's approval and use of Hensen's of academic spokesmanfor Darwin; it was to
analogous methodsin ecological studies on the markout thisarea of biologyas a scientificentity
earthworm.In his work on The Formationof and to provideit with an effective name happily
VegetableMould throughthe Action of Worms reminiscent of the older conceptof the economy
(1881), Darwin was interestedin estimatingthe of nature-one of the many fruitfulideas which
weightof soil per acre broughtto the surfaceeach illuminatethe pages of the Originof Species. In
year by earthworms. For thishe praisedHensen's contrastto Darwin's appreciationof thepotential
"admirablepaper" on theproblem,and he did not value of quantitativestudiesas a meansof under-
hesitateto extrapolatefrommeasurements based standing the dynamics of ecological relations,
on a fewyard-squaresample areas. In regard to Haeckel's distrustof quantitativestudies seems
skepticism concerninghis conclusions Darwin static and unexpectedlyconservative.As a source
simplyobserved: "Here we have an instanceof fora vital stimulusto thecontinuingdevelopment
thatinabilityto sumup theeffects ofa continually of ecologywe must look ratherto the work of
recurrentcause, which has often retarded the CharlesDarwin.

LIST OF LITERATURE
ALLEE, W. C., A. E. EMERSON,0. PARK, T. PARx, edition). Watts, London, [1950]. I have verified
and KARLP. SCHMIDT. 1949. PrinciplesofAni- the quotationsin the firstedition,but have given
mal Ecology. Saunders,Philadelphia. page referencesto the more available reprintedi-
BURDON-SANDERSON, JOHNS. 1893. Inaugural Ad- tion.
dress. Nature, Lond., 48: 464-72. Also in Rep. . 1860. OberdieEntstehungderArten....Trans-
Brit. Ass., 1893: 3-31; Rep. Smithson.Instn, 1893: lated by H. G. Bronn. Schweizerbart,Stuttgart.
435-63. For each of the passages quoted I have checked
CUVIEER, GEORGES. 1817. Le Regne animal distribuc Bronn's translationand have foundit reliable.
d'apres son organisation pour servir de base 4, . 1881. The Formation of Vegetable Mould.
l'histoirenaturelledes animaux et d'introduction 4, throughtheAction of Worms,withObservations of
l'anatomiecomparee. Vol. 1, Paris. theirHabits. London.
. 1835. Lecons d'anatomie comparee. 2nd ed. HAECKEL,ERNST. 1852-1856. The Storyofthedevel-
Vol. 1, Crochard,Paris. opment of a youth. . . Lettersto his Parents,
DARWIN,CHARLES. 1839. Journalof Researchesinto 1852-1856. Translatedby G. B. Gifford. Harper,
theGeologyand NaturalHistoryofthevariouscoun- New York, 1923.
tries visited by H. M. S. Beagle.... Henry Col- . 1852-1919. Ernst Haeckel: Forscher,Kuinstler,
burn, London. [Facsim. reprint. Hafner, New Mensch. Briefe.... Edited by George Usch-
York, 1952.] mann. Urania, Jena, 1954.
. 1842. The Structureand Distributionof Coral . 1862. Die Radiolarien. Reimer, Berlin.
Reefs. [Reissued in Geological Observations..., 1863. Ueber die EntwickelungstheorieDar-
1851],London. win's. [In Gesammelte populdreVortrdge aus dem
. 1842-1844. The Foundationsof the Origin of Gebiete der Entwickelungslehre. Heft 1. 1878.]
Species: twoessayswritten in 1842 and 1844. Ed- Strauss,Bonn.
ited by Francis Darwin. Cambridge University . 1866. GenerelleMorphologieder Organismen.
Press, Cambridge,1909. Allgemeine Grundzige der organischenFormen-
. 1859. On the Origin of Species by Means of Wissenschaft,mechanischbegriindet durchdie von
Natural Selection,or the preservationof favoured Charles Darwin reformirteDescendenz-Theorie.
racesin thestruggle forlife. (A reprintof the first 2 vols. Reimer,Berlin.

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144 THE QUARTERLY REVIEW OF BIOLOGY

HAECKEL, ERNST. 1869. UeberEntwickelungsgang des


HENSEN, VICTOR. 1887. Ueberdie Bestimmung
undAufgabederZoologie. [In Gesammelte
popu- Planktons Materials
oderdesimMeeretreibenden
lare Vortriageaus dem Gebiete der Entwicke- an Pflanzenund Thieren. Wiss. Meeresuntersuch.,
lungslehre.
Heft 2. 1879.] Strauss,Bonn. 5: 1-107.
1873. Author'sprefaceto theEnglishedition. . 1891. Die Plankton-Expeditionund Haeckel's
[In The Historyof Creation.Vol. 1. 1887.] Apple- Darwinismus.UebereinigeAufgabenund Ziele der
ton,NewYork. Lipsius &
beschreibendenNaturwissenschaften.
. 1874. Aus einer autobiographischen
Skizze Tischer,Kiel.
vom Jahre 1874. [In HimmelhockJauchzend... JACKSON, B. DAYDON. 1913. Authorshipin the
Erinnerungenund Briefe der Liebe. Edited by AmoenitatesAcademicae[of Carl Linneaus, 1749-
Heinrich
Schmidt,
1927]. Reissner,
Dresden. 90]. J. Bot., Lond., 51: 101-103,224.
1875. The HistoryofCreation. Translatedby KLAAUW,C. J. VAN DER. 1936. Zur Geschichte der
L. D. Schmitz;revisedby E. R. Lankester.2 der Oekologie,
Definitionen besondersauf Grund
vols.,London. Disziplinen.Sud-
der Systemeder zoologischen
1879. The EvolutionofMan: a popular exposi- Arch.Gesch.Med., 29: 136-77.
hoffs
tion of theprincipalpointsof humanontogenyand -. 1951. Qu'est-ce que l'ecologie? Ann. biol.,
phylogeny.
2 vols. KeganPaul,London. 27: 103-15.
1883. The Pedigree of Man: and otheressays LiNNP, CARL VON, praeses. 1749. ... Specimenaca-
.... Translation[ofGesammelte populareVortrdgel demicumde oeconomianaturae .... Diss. Upsala
byE. B. Aveling. Freethought Com-
Publishing (I. J.Bibergrespondent).[AlsoinLinne'sAmoc-
pany,London. International
Libraryof Science nitatesacademicae,Vol. 2 (1751), pp. 1-58.1
and Freethought,
Vol.6. -, praeses. 1749. I. J. Biberg,respondent.The
-. 1890. Plankton-Studien. VergleichendeUnter- oeconomy of nature,pp. 39-129in Miscellaneous
suchungenitberdie Bedeutungund Zussammenset- tractsrelatingto natural history,husbandryand
zungderpelagischenFauna und Flora. G. Fischer, fromtheLatin [oftheAmoenitates
physick,translated
Jena. withnotesby B. Stillingfleet.
Academicael 2nd
--. 1891a. Plankton-Studien. Jena. Z. Naturw., ed. London,1762.
25: 232-336. -, praeses. 1760. ... Dissertatioacademicade po-
1891b. PlanktonicStudies:a comparativein- litia naturae. Diss. Upsala (H. C. D. Wilcke,
oftheimportance
vestigation and constitution
of respondent).[Alsoin Linn6'sAmoenitatesAcade-
thepelagicfaunaandflora. Translated
by G. W. micae,Vol.6, 1763,pp. 17-39.]
Field,Rep. U. S. Comm.Fish., 1889-1891:565-641. -, praeses. 1760. H. C. D. Wilcke, respondent,
"On the policeof nature,"pp. 129-64in Select
HARDY, A. C. 1955. A further
exampleofthepatchi-
Academicae,a
Dissertationsfromthe Amoenitates
nessofplankton
distribution.
Pp. 7-11in Papers Tractsrelating
to Mr. Stillingfleet's
supplement to
in Marine Biology and Oceanography. Supp. to
NaturalHistory, translatedby F. J. Brand. Vol.
Vol. 3 of Deep Sea Research. London. 1. London,1781.
HARTWIG, G. 1861. The Sea and its LivingWonders. McATEE,W. L. 1947. The Cats-to-Clover Chain.
Translatedfromthe fourthGermaneditionand Sci. Monthly,65: 241-42.
partlyrewritten by the author. 2nd ed. Long- SEARS, PAUL B. 1950. Charles Darwin. The Nat-
mans,Green,London. uralistas a CulturalForce. Scribners,New York.

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