HAECKEL, DARWIN, AND ECOLOGY
BY ROBERT C. STAUFFER
University
ofWisconsin
T INTRODUCTION
bya
H HE TERM ecologywas invented
theoreticalmorphologist who was not
a distinguishedstudentof ecologyand
who was outspokenlyhostileto one of
the most significantprogramsof eco-
logical researchof his day, the controversial but
pioneering ventureto establishplankton
influential
studieson a quantitativebasis. This paradoxofthe
positionof ErnstHaeckel (1834-1919) in the his-
toryof ecologyposes the problemforthispaper.
Haeckel's major contributions may be best under-
stood as an immediatebyproductof his aim to
interpret Darwin's thoughtforthe scientific world
ofGermanyratherthanas thelogicaloutgrowth of
his own research.In his workas a fieldnaturalist,
Haeckel showeda painter'seye forthebeautiesof
nature,and he was an ardentcollectorwhosetrav-
els rangedfromNorway,Helgoland,the Canaries,
and Messinato theRed Sea, Cylon,and Java. This
wide and vivid experiencebroughthim some ap-
preciationofecologicalpointsofview,but Haeckel
did not display any notable insightinto the dy-
namicprinciplesofecology.My purposehereis to
examinethenatureand thepersonallimitationsof
Haeckel's contributions to the science he named
and to demonstrate its Darwinianderivation.
HAECKEL AND DARWINISM
Ecology was introducedinto the vocabularyof
sciencein 1866 in Haeckel's Generelle
Morphologie.
This was the firstof a numberof books which
Haeckel wroteto forwardthe cause of Darwinism
as he saw it,and we muststartwitha glanceat the
generalbackgroundof Haeckel's earlyenthusiasm
forCharlesDarwin (1809-1882).
Even as a medicalstudentat Wiirzburgin 1853,
in writinghome to proposebooks forfamilyva-
cation reading,Haeckel had been appreciativeof
the "excellenttravelsof Darwin" (Haeckel, 1852-
138
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1856, p. 124). In the autumn of 1859, however,
whenDarwin's Originwas published,Haeckel was
far fromthe worldof scientificnews. He was at
Messina on the straitsbetweenSicily and Italy,
wherehe was followingthe exampleofhis beloved
teacher,JohannesMiiller(1801-1858),in studying
the richesof the sea to be capturedwith the col-
lecting net. Using the methods which he had
learnedfromMiillerat Helgolandin 1854 (Haec-
kel, 1852-1856,p. 251), Haeckel obtained many
new species of Radiolaria to delighthis artist's
eye. The resultsof thesemonthsof collectingand
study at Messina he broughtnorthto formthe
foundationfor his masterpiece-his first huge
monograph,Die Radiolarien,whichwas to secure
him his appointmentas ausserordentlicher Pro-
fessorin the medicalfacultyat Jena at the age of
28. It was thusonlylater,whilehe was working
up his Italian collections,that Haeckel became
familiarwiththeOriginofSpecies.
This encounterwas to be a major influenceon
Haeckel's personal beliefs and on his scientific
career."Aftermy returnfromItaly in 1860 when
I became acquaintedwithDarwin's work,whenI
saw in it the way preparedfor a mechanisticor
monisticsolutionofthemostdifficult philosophical
problems,thentherebegan to developwithinme a
unifiedview of lifein theface of whichonlya few
of my long-accustomedand beloved articles of
faithcould endureany longer" (Haeckel, 1874,p.
333). In his Radiolarien,Haeckel used Darwinian
conceptsin his discussionof the relationsof the
familiesof this order,in his considerationof a
genealogicaltable, and his speculationsregarding
the originalprimitivetypeformof the order;and
he emphasized the evolutionarysignificanceof
transitionalforms.Here Haeckel expressedhis
enthusiasmforthe stimulatingideas of the Origin
of Species and explicitlydeclared his beliefthat
Darwin's theories"had beguna new epochforthe
 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
swarms,theirabundant accumulationin 'zo6cur-
rents,'theirrelationto ocean currentsin general,
are onlya fewof the manygreatproblemswhich
planktologyoffers to theresearchimpulseofman"
(p. 334). In thewholeplanktoncontroversy, while
Haeckel may have been correctin importantmat-
tersofdetail,he persistentlyobscuredthepotential
value inherentin Hensen'sprogram.
But it is timeto contrastHaeckel's suspicionof
Hensen'squantitativemethodsin planktonstudies
with Darwin's approval and use of Hensen's
analogous methodsin ecological studies on the
earthworm.In his work on The Formationof
VegetableMould throughthe Action of Worms
(1881), Darwin was interestedin estimatingthe
weightof soil per acre broughtto the surfaceeach
year by earthworms. For thishe praisedHensen's
"admirablepaper" on theproblem,and he did not
hesitateto extrapolatefrommeasurements based
on a fewyard-squaresample areas. In regard to
skepticism concerninghis conclusions Darwin
simplyobserved: "Here we have an instanceof
thatinabilityto sumup theeffects ofa continually
recurrentcause, which has often retarded the
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ALLEE, W. C., A. E. EMERSON,0. PARK, T. PARx,
and KARLP. SCHMIDT. 1949. PrinciplesofAni-
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BURDON-SANDERSON, JOHNS. 1893. Inaugural Ad-
dress. Nature, Lond., 48: 464-72. Also in Rep.
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143
progressof science, as formerlyin the case of
geology,and morerecentlyin thatof theprinciple
of evolution" (quotations are from the end of
Darwin's Introduction).
CONCLUSION
Haeckel's original presentationof the term
"ecology" in 1866 embodiesconceptsthat come
straightfromthe Originof Species. His contribu-
tion to the scienceof ecologywas made in his role
of academic spokesmanfor Darwin; it was to
markout thisarea of biologyas a scientificentity
and to provideit with an effective name happily
reminiscent of the older conceptof the economy
of nature-one of the many fruitfulideas which
illuminatethe pages of the Originof Species. In
contrastto Darwin's appreciationof thepotential
value of quantitativestudiesas a meansof under-
standing the dynamics of ecological relations,
Haeckel's distrustof quantitativestudies seems
static and unexpectedlyconservative.As a source
fora vital stimulusto thecontinuingdevelopment
of ecologywe must look ratherto the work of
CharlesDarwin.
edition). Watts, London, [1950]. I have verified
the quotationsin the firstedition,but have given
page referencesto the more available reprintedi-
tion.
. 1860. OberdieEntstehungderArten....Trans-
lated by H. G. Bronn. Schweizerbart,Stuttgart.
For each of the passages quoted I have checked
Bronn's translationand have foundit reliable.
. 1881. The Formation of Vegetable Mould.
throughtheAction of Worms,withObservations of
theirHabits. London.
HAECKEL,ERNST. 1852-1856. The Storyofthedevel-
opment of a youth. . . Lettersto his Parents,
1852-1856. Translatedby G. B. Gifford. Harper,
New York, 1923.
. 1852-1919. Ernst Haeckel: Forscher,Kuinstler,
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