Nature and Structure of the Climax Author(s): Frederic E. Clements Source: Journal of Ecology, Vol. 24, No. 1 (Feb.

, 1936), pp. 252-284 Published by: British Ecological Society Stable URL: http://www.jstor.org/stable/2256278 . Accessed: 13/10/2011 14:29
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NATURE AND STRUCTURE OF THE CLIMAX

BY FREDERIC E. CLEMENTS
Santa Barbara, California) of Institution Washington, (Carnegie (WithPlates VI-XI)
CONTENTS
PAGE INTRODUCTION

...253

THE CLIMAX CONCEPT .253

NATURE

OF THE

CLIMAX

.254

Unity of the climax.254 Stabilisationand change.255 Originand relationship.257 Tests of a climax .257
CLIMAX AND PROCLIMAX

.261

Essential relations. Proclimaxes

261 262 265 266 267

Subelimax .262 Disclimax

Preclimax and postclimax. Preclimax

Postclimax .269
STRUCTURE OF THE CLIMAX .270

Community functions. Roles of the constituent species: dominants

270 270

. Influents Climax and seral units . Climax units

Association ..273

.271 . .271 .272

Consociation Faciation Lociation .. Society Sociation Lamiation Sation Clan

Seral units Serule
RANK TABLE AND

..274 ..274 ..275 ..276 ..277 ..277 ..278

. ..280
OF UNITS UNITS . . . . . . .

275

.278
280 281

CORRESPONDENCE AND SERAL

OF CLIMAX

Panclimax

and eoclimax

Prerequisites to research in climaxes

REFERENCES .. .

281 282 283

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INTRODUCTION

MOREthan a century whenLewis and Clarkset out upon theirmemorable ago journeyacrossthe continent NorthAmerica(1803-6), theywerethe first of to traverse greatclimaxesfrom the deciduouswoodsin the east through vast the expanse ofprairieand plain to the majesticconiferous forest the north-west. of At this timethe oak-hickory woodlandbeyondthe Appalachianswas almost untouchedby the ax except in the neighborhood a fewstraggling of pioneer settlements, and west of the Mississippi hardlyan acre of prairiehad known the plow. A few years later (1809), Bradbury states that the boundless prairiesare coveredwiththe finest verdureimaginableand willbecomeone of the mostbeautifulcountries the world,whilethe plains are of such extent in and fertility to maintainan immensenumberof aniimals.It appears probas able thatat thistimeno othergrassland the worldexhibited in suchmyriads of large mammalsbelonging but a fewspecies. to The naturalinference been that the prairies has weremuchmodified the by grazingofanimalsand the fires primitive of man, and thishas been reinforced by estimates of the population of each. Seton (1929) concludes that the originalnumberof bison was about 60 millionwith a probable reduction to 40 millionby 1800, and that both the antelope and white-tailed deer were equally abundant,whileelk and mule-deer each amountedto not more than 10 millionat the maximum. However, these were distributed over a billion or two acres,and the averagedensity was probablynevermorethan a scoreto the square mile. Estimatesofthe Indian tribesshowthe greatestdivergence, but it seems improbablethat the total populationwithinthe grasslandever exceeded a half million.The generalhabit of migration among the animals fromovergrazing insuredthat seriouseffects further and trampling were but whilethe influence fires by the Indians was even less of set local or transitory, in the thoseofthe northsignificant modifying plant cover. As to the forests, westwerestillprimevaland in the east theywereyetto be changedoverwide on and burning a large scale. areas by lumbering
THE CLIMAX CONCEPT

of The idea of a climaxin the development vegetationwas first suggested by Hult in 1885 and thenwas advanced moreor less independently several by investigators duringthe next decade or so (cf. Clements, 1916; Phillips,1935). It was applied to a more or less permanentand finalstage of a particular of area. The conceptofthe successionand henceone characteristic a restricted connected withits climateand often climaxas a complexorganism inseparably continental extentwas introducedby Clements(1916). Accordingto this in the and as suchforms the view,the climaxconstitutes majorunitofvegetation of The relationbetween basis forthe naturalclassification plant communities. to climateand climaxis considered be the paramountone, whilethe intimate

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of Nature and Structure the Climax

the from of by the bondbetween twois emphasized thederivation theterms climaxis invariably underthisconcept root. In consequence, same Greek the alone,namely, forcommunity to withreference the climatic employed mation itsmajordivisions. or that animalsmustalso be considered At the outsetit was recognized of for was and members theclimax, thewordbiome proposed thepurpose of 1916b; (Clements, rolesofplantsand animals uponthe mutual stress laying that the this 1936). With went realization theprimary and Clements Shelford, of by different virtue thefact to relations thehabitator ece werenecessarily On and that plantsare producents animalsconsuments. land, moreover, with in connection the matrix thebiome direct of the plantsconstitute fixed to as a dual relation, plants wellas to climate. bear the while animals climate, in upontheece,ofthe of effect theoneis displayed reaction The outstanding the bond of the upon plants,whichconstitutes primary otherin coaction community. biotic the climaxhas relation, term uponthe climatic Becauseofits emphasis as is which regarded an the formation, to and more replace word comemore to by mayhave beenfavored a tendency and exact synonym, thisprocess " " use. formation with avoidconfusion thegeological Thedesignationclimatic to its but beenemployed, thisis merely accentuate nature has nowand then climax biome and less it usages.Furthermore, andto distinguishfrom definite is though the when biotic communityto be indicated, synonyms arecomplete whenplants for to continue be employed the matrix climaxwillnecessarily aloneareconsidered.
NATURE OF THE CLIMAX

works detailin earlier in This themehas been developed considerable as wellas ina recent and 1929), 1928;Weaver Clements, 1920, 1916, (Clements, of a account (1935),andhence summary treatment Phillips by comprehensive in will place.Thesemaybe conveniently features suffice thepresent themajor and i.e. stabilization change, four the under following captions, unity, grouped tests. and and origin relationship, objective of climax Unity the rests uponthefactthatit is notmerely of unity' theclimax The inherent and the butis at thesametime expression climate, to theresponse a particular and the in variation space and time, of theindicator it. Becauseofextent, of to into climates, saynothing thehuman transition adjacent gradual usually are of nor neither physical humanmeasures a climate adequately equation, of nature the and the continuity, sessile By satisfactory. contrast, visibility, limits the in helpful indicating fluctuating are plantcommunity peculiarly and of growth in whileits direct response terms food-making, of a climate, factors. of Naturally, the integrationphysical possible life-form provides fullest

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both physicaland human values have a part in analyzingand in interpreting and not the climateas outlinedby the climax,but these can onlysupplement replace the bioticindicators. It may seem logical to inferthat the unity of both climax and climate but will shouldbe matchedby a similaruniformity, reflection make clearthat such is not the case. This is due in the first place to the gradual but marked one boundaryto the other, or from shift rainfall temperature in probablybest the illustrated the climateof the prairie. In termsof precipitation, latter by may range along the parallel of 40? fromnearly40 in. at the easternedge of 10 the true prairieto approximately in. at the westernborderof the mixed grassland,or even to 6 in. in the desertplains and the Great Valley of California. Such a change is roughly1 in. for50 miles and is regionally but all imperceptible. The temperature change along the 100th meridianfromthe mixed prairiein Texas to that of Manitoba and Saskatchewanis even more since only one association is concerned. At the south the average striking, but is frost about 9 months, at the north is less than 3, it periodwithout killing while the mean annual temperaturesare 70 and 330 F. respectively.The variation of the two major factorsat the extremesof the climatic cycle is likewisegreat,the maximumrainfallnot infrequently to amounting threeto fourtimesthat of the minimum. to The visible unity of the climax is due primarily the life-form the of dominants,which is the concreteexpressionof the climate. In prairieand steppe, this is the grass form,with which must be reckonedthe sedges, esthe three scrub climaxes of pecially in the tundra. The shrub characterizes North America, namely, desert, sagebrush,and chaparral, while the tree appears in threesubforms, coniferous, deciduous,and broad-leavedevergreen, and tropicalclimaxes.The lifeto typify corresponding the boreal,temperate, in is form naturallyreflected the genus,thoughnot withoutexceptions, since herbor shrub,deciduousor evergreen, or two or moreforms subforms, annual or perennial,may occur in the same genus. Hence, the essentialunity of a climaxis to be soughtin its dominantspecies,sincetheseembodynot onlythe a life-form the genus,but also denotein themselves definite and relation the to climate. Their reactionsand coactions are the most controlling both in kind and amount, and thus they determinethe conditionsunder which all the remaining speciesare associated withthem.This is trueto a less degreeofthe animal influents, thoughtheircoactionsmay oftenbe more significant than those of plants. Stabilization and change Under the growingtendencyto abandon static concepts,it is comprehensiblethat the pendulumshould swingtoo far and changebe overstressed. is This consequence fostered thefactthat mostecologicalstudiesare carried by is out in settledregionswheredisturbance the rulingprocess. As a result, the

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are or climaxis badlyfragmented evenabsentoverwideareasand subseres or impossible difficult entirely it legion. In all suchinstances is exceedingly to imperative and stability change, it becomes and between to strike balance a is control still climatic by muchless disturbed man,where turnto regions measure time, of a essential employ conceivable to It paramount. is likewise thanin eons. No rather of in terms millennia suchas canbe expressed human a have evolved, entertains doubtthat climaxes student past vegetation of of underthe compulsion greatclimatic changes and migrated disappeared that through but from Paleozoiconward, he is also insistent theypersist the and in of disturbances millions years theabsence suchchanges ofdestructive of the of within limitations evidence is goodand evenconclusive by man.There hasbeeninexistence several for climax millions that fossil materials theprairie of most thedominant of Thisis even at species to-day. ofyears leastandwith to on of true Coast, owing thewealth fossil more certainly offorests thePacific the of dominants thedeciduous evidence 1925,1935),while generic (Chaney, are similar. and of forests theDakota Cretaceous ofto-day strikingly is that affirmed stabilization the universal It can still be confidently and that climaxes underthe ruling are of climate, tendency all vegetation in when reckoned thousands even of or characterized a high degree stability by thanthedevotee succession more of of clearly millions years.No onerealizes but in the absenceof at and that changeis constantly universally work, and notdestructive it. of the manthisis within fabric theclimax of civilized as as West,the prairie developed the Middle Even in a country intensively and of almostcomplete exhibit in stability dominants subdominants relicts 1934). It is (cf. by spiteofbeingsurrounded cultivation Weaverand Flory, withthe season,year or obviousthat climaxes changes displaysuperficial the but and itself as in aspection annuation, thesemodify matrix little cycle, in be present millions yearand one of annuals thedesert ornotat all. The may one or the mayseemprevailing seasonand a grass absent next, onedominant are recurrent or one different the following year,but thesechanges merely the While modifications by represented bareareasand indeedonlyapparent. these in process striking, areall intheirresistible stagearemore byseres every and climate theinterference as as stabilized rapidly thecontrolling of ofbeing manpermit. or annuation natural due the coaction are In brief, changes to aspection, and orperiodic leaveno permanent while those of impress, fleeting superficial, are process.Man alone can succession an intrinsic part of the stabilizing the of the by destroy stability the climaxduring longperiodof control its in of this and climate, heaccomplishes byfragmentsconsequence a destruction renewed. and or thatis selective, partial complete, continually

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Originand relationship each climaxnot onlyhas its owngrowth organisms, Like otherbut simpler but succession, it has also and secondary in and development termsofprimary evolved out of a precedingclimax. In otherwords,it possessesan ontogeny studied,much and experimentally that can be quantitatively and phylogeny as with the individualsand species of plants and animals (Plant Succession, 1916, pp. 181, 342). Out of the one has come widespread activity in the on in of whileinterest the otherlingers the threshold, investigation succession, chiefly because it demands a knowledgeof the climaxes of more than one With increasing researchin these,especiallyin Europe and Asia, it continent. the willbe possibleto test critically panclimaxesalreadysuggested(Clements, the of 1916, 1924, 1929), as well as to determine originand relationships the constituent formations. This task will also require the services of paleo-ecologyfor the reconstructionof each eoclimax, which has been differentiated worldwide by climatic changes into the existingunits of the panclimax or panformation. of As it is, therecan be no seriousquestionofthe existence a greathemispheric and subtropical clisereconstituted the arctic,boreal,deciduous,grassland, by for an tropicalpanclimaxes. Desert formations the mostpart constitute exception and may well be regardedas endemic climaxes evolved in response 1935). to regionalchangesof climate(Clements, of It is a significant fact that the boreal formations NorthAmerica and ones of the former, than the coniferous but Eurasia are morecloselyrelated climaticdifferences have that thisseeming anomalyis explainedby the greater produced the forestsof the Petran and Sierran systems.The five climaxes and it is possibleto indicatetheirrelationwellknown, are concerned relatively shipswithsome assurance,and all the morebecause oftheirparalleldevelopand menton the two great mountainchains. In the case of deciduousforest in of onlya singleformation each is present NorthAmerica,and the grassland, the and relationship intotracing origin resolvesitself of problem differentiation of the several associations. It has been suggestedthat the mixed prairieby the virtueof its position,extent and commondominantsrepresents original reinforced its close resemblance in formation Tertiary by times,an assumption to the steppe climax (Clements,1935). It is not improbablethat the mixed hardwoodsof the southernAppalachians bear a similarrelationto the associationsof the moderndeciduousforest (Braun, 1935). Testsofa climax As has been previouslyindicated,the major climaxes of NorthAmerica, and prairie, stand forth such as tundra,boreal and deciduousforest, clearlyas distinctunits,in spite of the fact that the prairiewas first regardedas comas prisingtwo formations, a consequenceof the changes producedby overJourn.of Ecology XXIV 17

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Nature and Structure the Climax of

grazing.The otherconiferous and the scrub climaxes emergeless distinctly because ofthe greater of withineach group,and henceit is similarity life-form necessaryto appeal to criteriaderivedfromthe major formations just mentioned.This insuresuniformity basis and a highdegreeof objectivity, of both ofwhichare qualitiesofparamountimportance the naturalclassification for of biomes. In fact, entireconsistency the application of criteriais the best in warrant objectiveresults, of thoughthisis obviously procedure a thatdemands a first-hand acquaintance withmostif not all the unitsconcerned and over a large portionof theirrespectiveareas. The primary criterion that afforded the vegetation-form, is illusis by as tratedby thefourmajorclimaxes.The othersofeach group,suchas coniferous forest scrub,are characterized or also by the same form the dominants, in but this is not decisive as betweenrelated climaxes and hence recoursemust be taken to the othertests. The value of the life-form mostevidentwheretwo is climaxes of different are physiognomy in contact,as in the case of the lake forest pine-hemlock the deciduousforest hardwoods.The staticview of of and would make the hemlockin particular dominant the deciduousformation, a of but the evidence derivedfromthe vegetation-form supportedby that of is phylogeny and by early recordsof composition and timber-cut show that to two different climaxes are concerned. Secondary formsor subforms rarely if ever mark distinctions betweenclimaxes,but do aid in the recognition of associations.This is well exemplified the tall, mid and shortgrassesof the by and somewhat less definitely the generally prairie deciduouscharacter the by of Petran chaparraland the typicallyevergreen natureof the Sierran. As wouldbe expected,the mostsignificant ofthe unityof a formation test is afforded the presenceof certaindominantspecies through or nearly by all all of the associations,but oftennot in the role of dominantsby reason of reducedabundance. Here again perhapsthe best examples are furnished by prairieand tundra,thoughthe rule applies almost equally well to deciduous ones because of a usually smallernumber and onlyless so to coniferous forest of dominants. For the prairie,the numberof such species,or perdominants, found in all or all but one or two of the five associationsis eight,namely, Boutelouagracilis,Sporoboluscryptandrus, Stipa comata,Agropyrum smithi, Poa scabrella and Festucaovina. Even when Koeleriacristata, Elymussitanion, a speciesis lackingover most of an association,as in the case ofStipa comata and the true prairie,it may be represented a close relative,such as S. by spartea,whichis probablyno morethan a mesicvarietyofit. As to the three a associationsof the deciduousforest, stilllargernumberof dominantspecies occurto some degreein all; the oaks compriseQuercusborealis, velutina, alba, stellataand marilandica, and the hickories, macrocarpa, coccinea, muhlenbergi, Carya ovata,glabra,alba and cordiformis. It was the application of this test by specificdominantsthat led to the mantle of the Petran and of recognition the two climaxes in the coniferous

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Sierrancordilleras. The naturalassumption was that such a narrow belt could not contain more than one climax, especiallyin view of its physiognomic uniformity, thisfailedto reckonwiththe greatclimaticdifferences the but of two portionsand the corresponding responseof the dominants.The effect of altitude proved to be much more decisive than that of region,dominants commonto the montaneand subalpinezones beingpractically absent,though theruleforthe same zone in each ofthetwo separatemountain systems.Long afterthe presenceof two climaxes had been established,it was foundthat Sargenthad anticipatedthis conclusion, thoughin otherterms(1884, p. 8). As wouldbe inferred, dominants relatedassociationsbelongto a few the of commongeneraforthe most part. Thus, there are a dozen species of Stipa variouslydistributed dominantsthrough as the grassland,nearlyas many of Sporobolus,Bouteloua and Aristida, and several each of Poa, Agropyrum, Elymus, Andropogon, Festuca and Muhlenbergia.In the deciduous forest, Quercus, Carya and Acer are the greatgenera,and forthe various coniferous ones, Pinus, Abies and Picea, with species of Tsuga, Thuja, Larix and Juniperushardlyless numerous. The perennialforbsthat play the part of subdominants also possess considerablevalue in linkingassociationstogether, and to a higherdegreein the to deciduousforest than in the prairie,owingchiefly the factorsof shade and to protection.Over a hundredsubdominants belonging two score or moreof genera,such as Erythronium, Dicentra,Trillium,Aquilegia,Arisaema,Phlox, Asterand Solidago,range Uvularia,Viola, Impatiens, Desmodium, Helianthus, from Nova Scotia or New England beyondthe bordersofthe actual climaxto Nebraska and Kansas. Acrossthe wide expanse of the prairieclimax,species in commonare onlyexceptional, to thesefewbelonging mostly the composites, Artemisia notably Grindeliasquarrosa,Gutierrezia dracunculus sarothrae, and On the otherhand, the numberof genera of subdominants found vulgaris. the throughout grasslandis verylarge. of The greatermobility the largermammalsin particularrendersanimal but influents significant less than plants as a criterion, severalofthesepossess definite value and the less mobilerodentseven more.The antelopeand bison are typicalof the grasslandclimax,the first restricted it, to beingpractically while jack-rabbits, ground-squirrels and kangaroo-ratsare characteristic dwellersin the prairie,as is theirchieffoe,the coyote. or The remaining criteria derivedfrom are development directly indirectly, the ecotonebetweentwo associations. thoughthisis less evidentin the case of Here the mixing of dominants and subdominantsindicates their general in withinwhich range their preferences similarity terms of the formation, successionis associations.The evidencefrom assignthemto different primary of value only in the later stages as a rule,since initialassocies like the reedswamp may occur in severalclimaxes. With subseres,however,all or nearly all the stages are related to the particularclimax and such seres denote a
17-2

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Nature and Structure the Climax of

This is especiallytrueof all subelimaxes corresponding unityin development. still and most evidentlyin the case of those due to fire. More significant are postclimaxes bothgrasslandand forest.For example,the associesofspecies in of Andropogon, which is subelimax to the oak-hickory forest,constitutesa postclimax fiveout ofthe six associationsofthe prairie. On the otherhand, to the communityof Ulmus, Jubglans,Fraxinu?s, etc., found on flood-plains forms commonsubelimaxto thethree a the through regionofdeciduousforest, associations. In additionto such ontogenetic criteria, phylogeny suppliestests of even greater value. Thisis notablythecase withthetwo associationsofthe montane forests the west,thoughperhapsthe moststriking of and subalpineconiferous is this criterion in,connectionwith the lake forestof pineapplication of hemlock.Though the concreteevidenceforsuch a climax recursconstantly the regionof the Great Lakes to the Atlantic,it is fragmentary through and thereis no evidentrelated associationto the westward. However, the four genera are represented related species in the two regions,namely,Pinus by P. strobus P. monticola, banksianaby P. contorta, by Tsuga canadensisby T. Larix laricina by L. occidentalis, heterophylla, and Thuja occidentalis T. by plicata,thoughthe last two generahave changedfrom subelimaxrolein the a east to a climax one in the west. As suggestedearlier, evidence phylogenetic of still more direct nature is supplied by the mixed prairiewith the other enclosingassociationsand by the remnantsof a virgindeciduousforestthat exhibits a similar genetic and spatial relation to the associations of this climax (cf. Braun, 1935). Finally,it is clear that any test will gain in definiteness accuracy of and applicationwhenever dependablerecordsare available withrespectto earlier and structure. composition These may belongentirely the historical to period, as in the case of scientific reportsor land surveys,they may bridgethe gap betweenthe presentand the past as withpollen statistics, theymay reach or back into the geological record, as with leaf-impressions other further or fossils(Chaney,1925, 1933; Clements,1936). Two instancesof the scientific recordthat are of the first importance may be givenas examples.The first is the essentialrecognition Sargent of the pine-hemlock by climax under the name of the northern pine belt (1884), at a time whenrelatively littleof this had been logged,by contrastwith 90 per cent. or more at present(cf. also Bromley,1935). The second is an account,discoveredand communicated by Dr Vestal, of the prairiesof Illinois as seen by Short in ca. 1840. This is of interestsince its discoveryfollowedlittlemorethan a year after heightened repeated field trips had led to the conclusionthat all of Iowa, northern Missouri and mostofIllinoiswereto be assignedto the trueprairie,'a decision
1 The true prairieis characterized by the threeeudominants, Stipa spartea,Sporobolus asper and S. heterolepis. presenceof tall-grasses it to-day,particularly The in Andropogon fiircatu-s and nutans,is the markof the discimax due to the varied disturbances associated withsettlement.

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Phot. i. Sierran subalpineclimax: Consociation Tsuga of CraterLake, Oregon. mertensiana;

*....

..

Phot. 2. Proclimax Sagebrush of a disolimax (Artemisia due tridentata), to overgrazing mixedprairie;climatically preclimax. of a

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and supported by the more for confirmed Illinois by Short's description, generalaccountsof Bradbury(ca. 1815) and Greeley(1860).
CLIMAX AND PROCLIMAX

Essentialrelations in terminates the regularly withthe view that development In accordance itselfunder a particularclimate,except capable of maintaining community when disturbanceenters,thereis but one kind of climax,namely,that conin as trolledby climate.This essentialrelationis regarded not onlyinherent all in but naturalvegetation, also as implicit the cognatenatureofthe two terms. While it is fullyrecognizedthat successionmay be halted in practicallyany are invariablysubordinateto the true climax as stage, such communities by determined climatealone. From the verymeaningof the word,therecan routewitha genuineclimax alongthe developmental notbe climaxesscattered to questionthe realityof such pauses, but at the end. There is no intention orderfrom the climax. onlyto emphasizethe factthat theyare of a different thisdoes an Whileit is naturalto expressnewideas by qualifying old term, or not conduceto the clearestthinking the most accurate usage. Even more undesirableis the fact that the meaning of the originalword is gradually shifteduntil it becomes eitherquite vague or hopelesslyinclusive. At the disease,insects, this hands of some,climaxhas alreadysuffered fate,and fire, and human disturbancesof all sorts are assumed to produce corresponding climaxes(cf. Chapman,1932). On such an assumptioncornwould constitute one climax, wheat another,and cotton a third,and it would then become imperativeto begin anew the task of properlyanalyzing and classifying vegetation. In the light of two decades of continuedanalysis of the vegetation of NorthAmerica,as well as the applicationof the twinconceptsof climax and in complex organismby workers otherportionsof the globe and the strong and holism(Phillips, evolution to brought themby theriseofemergent support in of characterization the climax as givenin Plant Succession, 1916, 1935), the the accurate. " The unit of vegetation, still appears to be both completeand the arises, is climax formation, an organicentity. As an organism, formation grows,maturesand dies. Its responseto the habitat is shownin processesor that are the recordas well as the resultof these functions and in structures is each climax formation able to reproduceitself, functions. Furthermore, The life-history the withessentialfidelity stages ofits development. repeating is of a formation a complex but definiteprocess, comparable in its chief is of featureswiththe life-history an individualplant. The climax formation of of the adult organism, which all initial and medial stages are but stages in development.... A formation, short, is the final stage of vegetational of in development a climaticunit. It is the climax community a succession life-form in possiblein the climateconcerned." that terminates the highest

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of only wouldneedmodification to theextent subTo-day statement this and "biome" forclimaxor formation "biotic" forvegetational. stituting by and beenannotated confirmed Phillips' has Thischaracterization recently as above,a treatise organism, cited and of masterly discussion climax complex of in interested thefield dynamic by be thatshould readanddigested everyone and ecology itswideapplications. Proclimaxes that all includes the communities simulate term, proclimax As a general but or of in to the climax someextent terms stability permanence lack the were type of communitiesthis climate.Certain of sanction theexisting proper (p. in climaxes PlantSuccession 108; 1928,p. 109),and two calledpotential To and preclimax postclimax. avoid pronamely, kindsweredistinguished, was need,subclimax madeto do double in a posing newterm advanceofits as stage of succession, well as apparent both the subfinal duty,denoting by (1929, of climaxes otherkinds.This dual usage was criticized Godwin noteon theground in by justified Tansley an appended p. 144)and partially was thata newterm madeit evident thisdiscussion just given. However, 1934).While (Clements, suggested was desirable proclimax accordingly and in thistakescareoftheuse ofsubclimax thesecondsensenotedabove,it is to adaptedby reasonofits significance applyto all kindsofsubperbetter is an there still proper.However, manent other communities thantheclimax have been and postclimax preclimax subclimax, after residuum important later. as to and recognized, it is proposed call thesedisclimaxes, indicated or community as Theproclimax be defined anymore lesspermanent may replaceable but the resembling climaxin one or morerespects, gradually by is of by the latterwhenthe control climate not inhibited disturbance. for it function, maybe usedas a synonym any one ofits Besidesits general such further investigation, as in divisions, wellas in casesofdoubtpending as disclimax, are waterclimaxes. The fourtypesto be considered subclimax, and preclimax postclimax. Subclimax the and seres,primary As the stage preceding climaxin all complete well The is as secondary, subclimax as universal it is generally understood. the that especially belongto the subsere, of greatmajority such communities a is constant disturbanceto-day practically to that following owing thefact fire, in are processes cultivated of Fireandfallow recurrent feature mostclimaxes. subsere until corresponding the to and generally theyserve maintain regions the is the Though subclimax terminates disturbance. or protection conversion thesehavelongago endedin theclimax a of just as regular feature priseres, communities are area and the relatedsubclimax Qvermostof the climatic in muchrestricted size and widelyscattered.Smallnessis consequently

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Pinus and Quercus;the "Plains", Phot. 3. Fire subclimaxofdwarf New Jersey Pine-barrens.

of in purpurea field Phot. 4. Subclimaxof consocies Aristida abandonedI5 years; GreatPlains.

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being the of naturallya characteristic nearlyall subelimaxes, chiefexceptions due to fireor to fireand loggingcombined,but by contrastthey are often numerous. exceedingly the the subclimaxresembles preof its positionin the succession, Because respectsand in a few instanceseithertermmay be properly climax in some applied. The distinctionbetween subclimax and disclimax presents some now and then, as the amount of change necessaryto produce the difficulty the This arisesin part also from structural lattermaybe a matterofjudgment. the of as and of diversity formation association, a consequence which dominants areas. When thereis but a of a particulartype of subserevary in different no as singledominant, in manyburnsubclimaxes, questionensues,but if two or more are present,the decisionbetweensubclimax and disclimaxmay be in less simple,as is not infrequent scrub and grassland. cases beingmostly Examples of the subclimaxare legion,the outstanding or alone or afterlumbering clearing. Most typical are those comdue to fire, posed of " jack-pines" or species with closed cones that open most readily afterfire. Each greatregionhas at least one of these, e.g. Pinus rigida,virin giniana and echinatain the east, P. banksianain the north,P. murrayana and radiata on the Pacific muricata the Rocky Mountains,and P. tuberculata, rolein the " piney" woodsofthe and taedaplay a similar Slope. Pinus palustris The as does Pseudotsugataxifoliain the north-west. Atlantic Gulf region, are composedofaspen (Populus subclimaxesofthe borealforest characteristic (Betula papyriand paper-birch balsam-poplar(P. balsamifera), tremuloides), fera), eithersinglyor in various combinations. Aspen also formsa notable for subclimaxin the Rocky Mountains, the most part in the subalpinezone. Prisere subclimaxesare regularfeaturesof bogs and muskeags throughout the threedominantsbeing Larix much or all of the boreal and lake forests, oftenassociated as zonal conlaricina,Picea mariana and Thuja occidentalis, two xeric of the deciduousforest, socies. Wherepines are absentin the region may constitutea subclimax,and this oaks, Quercusstellataand marilandica, assumedby smalltrees,Sassafras,Diospyrusand Hamamelis roleis sometimes being especiallyimportant. usually Subclimaxesin the grasslandare composedlargelyof tall-grasses, this part is taken by Spartina in the formof a consocies. In the true prairie, while Elymus and in the desert plains by Sporoboluswrighti, cynosuroides a similarrole in the mixed prairieand in portionsof the condensatus plays is of The function thetall Andropogons morevaried; they prairies. bunch-grass are typicallypostclimaxratherthan subclimax,though they maintain the and in oak openings. forest of latterrelationalong the fringe the oak-hickory They occupy a similarpositionat the marginof the pine subclimaxin Texas " be especially,and hencetheyare what might termed" sub-subclimax in such situations. Beyondthe forestand in associationwithElionurus,Trachypogon, etc., they appeaT to constitutea faciationof the coastal prairie. Chaparral

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proper tobe regarded a climax, with change species extends is as a it but of into themontane even subalpine andthere and a fire the zone constitutes subclimax. In thefoothills southern of the in California, coastalsagebrush behaves like manner where liesin contact it with chaparral. the The disposition seralstages of belowthesubclimax exhibit distinct that a retardation haltfora longer shorter or is or period a debatable matter.It is themamongsubclimaxes, thiswouldagain entirely possible include to but failinaccuracy definiteness hence and and leadto confusion. decision The may well be leftto usage by providing termforsuch seralor sub-subclimax a communities persist a longorindefinite as for becauseofcontinued period or control human or disturbance. virtue itssignificance, recurrent of edaphic By " serclimaxis suggested, " and with related the brevity accord terms, designation withthemeaning a seralcommunity of one before the usually or twostages as subclimax, which persists such a period to resemble climax this for in the and one respect.For reasons brevity agreement, connecting of the is vowel but omitted, thee remains longas in sere. in serclimaxes found standing are Forthemost part, water in saturated or soils as a consequence imperfect of The universal is drainage. example the with reed-swamp one or moreof several suchas Scirpus, consocies, Typha, and this Zizania,Phragmites Glyceria: is typical thelower of reaches rivers, of of deltasand of certain kindsoflakes,the greattule swamps California of instances.Another affording in outstanding in typeoccurs coastalmarshes is which the Spartina often soleormajordominant, while have sedge-swamps a wider climatic rangebut are especially characteristic northern of latitudes and highaltitudes. The Everglades Floridadominated Cladium of by constitute perhaps most the extensive example thegeneral of group, though Carex often swamps covergreatareasand the grassArundinaria forms jungle-like cane-brakes the through south. Among woody species, Salix longifolia an is consocies sand-bars of omnipresent and river-sides, the most unique but is exemplar thecypress-swampthesouth, of typified Taxodium. boreal by In and subalpine districts distinctive the serclimax the peat-bog, is mooror moreor less regularly associated withother muskeag, seralcommunities of Carex and usually LarixorPicea also in theproper of region. or Frequent the burning mayretard prevent development thenormal of fire subclimax causeit to be replaced a preceding and by stage.Thismaybe a scrub or community onekeptin theshrub form repeated by fires, along but theAtlantic Gulf and Coastsit is usually ofAndropogon one virginicus, owing " toitssufferanceburning. so-called balds" of southern of The the Appalachians areseralcommunities heaths grasses of or initiated maintained and primarily there serclimaxes weeds, are byfire. Finally, of in especially annuals, cultivated and similar districts, a somewhat of communitynative annuals characteristic is ofwidestretches thedesert in region.

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Phot. 5. Serclimax Taxodium., of a Nyssa and Quercus forming cypress swamp; Paris,Arkansas. Face26 p.

Phot. 6. Disclimax of Bouteloua, Muhlenbergia and Opuntia, due to overgrazing of mixed prairie, Great Plains.

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Discllmax As withthe relatedconcepts, significance thistermis indicatedby a the of prefix,dis-, denotingseparation,unlikenessor derogation,much as in the Greekdys,poor, bad. The most frequent examples of this community result fromthe modification replacement the true climax,eitheras a whole or or of in part, or froma change in the directionof succession.These ensue chiefly in consequenceofa disturbance man or domesticated by animals,but theyare also occasionally producedby mass migration.In some cases, disturbance and the introduction alien species act together of throughdestruction and competition constitute quasi-permanent to a community withthegeneralcharacter of the climax. This type is best illustratedby the Avena-Bromus disclimax of California, whichhas all but completely replacedthe bunch-grass prairie.' A similarreplacement Bromustectorum morerecently by has taken place over large areas of the Great Basin, while Poa pratensishas duringthe last halfcentury steadilyinvaded the nativehay-fields pasturesofthe trueprairie, and an advancefirst notedbyBradbury 1809. An evenmorestriking in phenomenon is the steadilyincreasing dominanceofSalsola overrangeand cropland in the west, and this is imitatedby Sisymbrium and Lepidium in the north-west. It is obviousthat all cultivatedcropsbelongin the same generalcategory, but this point hardlyrequiresconsideration. Probably the example most cited in NorthAmericais that of the shortgrassplains,whichactually represent reductionof the mixedprairiedue to a overgrazing, supplemented periodicdrouth. Over mostofthis association, by the mid-grasses, Stipa, Agropyrum, etc., are still in evidence,though often reducedin abundanceand stature, in someareas theyhave beenpractically but eliminated. Similar though less extensive partial climaxes of short-grasses characterise pasturesin the true and both pasture and range in the coastal prairie, dominants the regularly belonging Bouteloua, to Buchloeor Hilaria. Of essentially the same natureis the substitution annual species of Bouteloua of
1 The grassland climax of North America comprisessix well-markedassociations (Plant Indicators,1920; Plant Ecology,1929). The mixed prairie,so-called because it is composed of both mid-grasses and short-grasses, more or less centralto the otherfiveand is regardedas is ancestralto them. To the east along the Missouriand MississippiRivers, it has become differentiatedintothe trueprairieformed otherspeciesofmid-grasses by pertaining mostly the same to genera,and this unitis flankedalong the western marginof the deciduousforestby a proclimax oftall grasses,chiefly Andropoqon. Southwardthe trueprairieis replacedby coastal prairie, which in the main occupiesthe GulfregionofTexas and Mexico and is constituted similardominants by but of different species. The desertplains are characterized primarily species of Boutelouaand by Aristida,whichrangefromwesternTexas to the edge of the desertsof Mexico and Arizona. In the north-west short-grasses the disappear and the Palouse prairie of easternWashingtonand adjacent regionsis formed mid-grasses the bunch-grass by of life-form, among whichAgropyrum spicatum the eudominant.The same life-form is signalizesthe California prairie,foundfromthe northern part of the state southwardinto Lower California, but its especial characteris derived from endemicspeciesofStipa. As indicatedin the discussion, short-grass the plains, composedof Bouteloua,Buchloe,and Carex,are not climaticin nature,and this statementapplies likewiseto the tall-grass meadows of Andropogon mentioned above.

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of Nature and Structure the Cllimax

which ones for plains perennial ofthesamegenera, in and Aristida thedesert ones. by being is a case ofshort-grasses followed stillshorter a is of the instances, effect disturbance to produce community In other is the concerned thatofan when life-form of the with appearance a postclimax, when the or undershrub tall grass.Thisis notably case in themixedprairie sod up of to is overgrazingcarried thepoint breaking theshort-grass andperIn orGutierrezia sarothrae. essence, of the mitting dominance Artemisiafrigida and tridentata) of creosote-bush of (Artemisia the wideextension sagebrush eachoftheseis a climax though is tridentata)thesamephenomenon, (Larrea life-form the in dominant its ownregion.In the case of Opuntia, peculiar speciesbehavein all but difference, the numerous important an suggests of like othershrubs, thaughwiththe two advantages respects significant and propagation. spines ready originally of by formed species Andropogon of Thecommunitiestall-grasses of have all theappearance a since somedifficulty, thesenaturally presented to are number to be assigned the to postclimax theprairie.Probably greater that indicates inthe and from reconnaissance record but this type, theevidence in furcattus partiAndropogon the portion, and trueprairie especially eastern and in some mowing a due to pasturing, now constitutes disclimax cular in disclimax miniature 1933). A characteristic also to measure fire (Clements, coaction where tundra, of gardens" thealpine in is to be found the "gopher to of the have removed climaxdominants sedgesand grasses and reaction and " forbs." Towns ofprairie-dogs of gardens perennial flower make placefor communities. extensive more but similar much often kangaroo-rats produce as disclimaxes, may likewise initiates not cutting infrequently Selective disease.The most or agentssuchas fire epidemic actionof other the similar from dentata) (Castanea of is example the elimination the chestnut dramatic has importance beentheextreme evengreater but canopy, of theoak-chestnut of the through overcutting of and reduction fragmentation the lake forest climatic from a mayresult whatis essentially disclimax white pine. Finally, such as in the Black Hills of SouthDakota has brought mass migration, climaxoftheRockyMountains from montane the Pinusponderosa together forest. from boreal the and Picea canadensis and Preclimax postclimax (1916,1928) in were advanced PlantSuccession first concepts Theserelated (1929)andin theorganisation in PlantEcology and havesincebeendiscussed of corollaries the principle method (1934).Theyare bothdirect oftherelict by thatare setin motion a major of the oftheclisere, spatialseries climaxes phases.The its suchas thatoftheglacialepochwith opposite climatic shift, or in is comprehended the case ofhighranges summits, clisere mostreadily and visible, is is of series climaxes readily the suchas PikesPeak where entire of journey 1700. in described MountArarat his famous on whatTournefort

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However,thisis but an expression the continental of cliserein latitude,which achievesperhapsits greatest in regularity NorthAmerica. A similar relation is characteristic the longitudinaldispositionof climaxes in the temperate of zone between the two oceans, the portion fromdeciduous forestthrough prairieto desertbeingthe most uniform. With the exception of the two extremes,arctalpineand tropical,each climax has a dual role, being preclimaxto the contiguouscommunity soof called higher life-form postclimaxto that of lowerlife-form. may be and This illustrated the woodlandclimax,whichis postclimaxto grasslandand preby climax to montane forest.The arctic and alpine tundras exhibit only the preclimax relation, borealand subalpineforest to respectively, sincea potential lichenclimax attains but incomplete expression northward upward.While or thegeneral primary relation one ofwaterin terms rainfall evaporation, is of and temperatureconstantlyenters the situation and at the extremesmay be largelycontrolling, in thetundraespecially. However,in ourpresent as imperfect knowledgeof causal factorsit is simplerand moredefinite determine to rankby positionin the cliseralsequence,each community higher altitudeor in latitude being successivelypreclimaxto the precedingone. This relationis likewiseentirely consistent the cliserefromdeciduous forestto desert,as in it is amongthe associationsofthe same climax,thoughin boththesecases the zonal grouping may be moreor less obscured. Wherever concrete preclimaxes or postclimaxes occur, either between climaxesor withina singleone, theyare due to the compensation afforded by edaphic situations.The major examplesof the latterare providedby valleys, especiallygorgesand canyons, long and steepslope-exposures, by extreme and soil-types such as sand and alkali. The serationis a seriesof communities produced by a graduated compensationacross a valley and operating withina formation throughadjacent ones, while the ecocline embracesthe differor entiationbroughtabout by shifting slope-exposures arounda mountainor on the two sides of a highridge. In the case of such soils as sand or gravelat one extremeand stiff clay at the other,the edaphic adjustmentmay sometimes appear contradictory. Thus, sand affords haven forpostclimaxrelictsin the a dryprairieand forpreclimaxones in the humidforest region, whilethe effect ofheavy soilsis just the reverse. However,thisis readilyintelligible whenone recalls the peculiar propertiesof such soils in termsof absorption,chresard and evaporation(Clements, 1933). Preclimax Since theyoccupythe same generalantecedent positionwithrespectto the climax,it is necessaryto distinguish with some care betweensubclimaxand preclimax, especiallyin view of the factthat theyoftenexhibitthe same lifeform. However,this is not difficult whenthe priseres and subsereshave been investigatedin detail, as the actual compositionand behavior of the two

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leads reaction in Moreover, thefirst, quitedifferent. are communities usually while the in conversion, withultimate of to the entry theclimaxdominants within to if is by the second compensation localfactors rarely ever, be overcome disturbance. of short man-made climate, theexisting are two adjacentformations where marked are Preclimaxes mostclearly and prairie.Examplesof the or and forest desert prairie either concerned, kind are foundin the grassy"openings"and oak savannahsof the first " of " parks alongthemargin the and forest in theso-called natural deciduous dry on Theyare also welldeveloped warm slopeforests. and montane boreal is In in or exposures xeroclines theRockyMountains. theone,compensation due by soil, or by afforded a sandy rocky intheother a localclimate to usually to bear relation circumjacent regularly theproper climaxes Desert insolation. which life-form, would by obscured theshrub but grassland, thisis somewhat This formation. maybe explained, the to be expected characterize less xeric dominants by shown suchmajor for by however, thewidecapacity adaptation in a quality thatis lacking most and tridentata, as Larreatridentata Artemisia plainsand in communities desert as associates.Leftstranded relict oftheir by phase, they haveprofited the of by mixed prairie therecession thelast dry thanthatin muchlarger acrossa territory to ofgrasses extend overgrazing of theyare climax. Here theyhave all the appearance a postclimax, which several attainsa stature in especially the case of Larrea,whichcommonly of outcome sincethisis thedirect in thatfound thedesert.However, times partias regarded a disclimax, it of in disturbance terms grazing, is better in still grasses persist it to somedegree. sincetheclimax cularly or the the Within sameformation, morexericassociations consociations the between relation are preclimax theless xericones.Thisis thegeneral to the forest, former of associations thedeciduous and oak-hickory beech-maple or by sitesproduced insolation type drier the in occupying thelatter warmer the mayobtainin the case offaciations, Quercus relation of soil. A similar preclimax of beinga border marginal often community stellata-marilandica persist naturally Suchpreclimaxes faciations. oak-hickory mesic to themore or in as proper relicts valleys sandysoils the of beyond limits theassociation a grassland, to and thenassumethe role of postclimaxes the surrounding " in exemplified the "Cross Timbers of Texas. In the strikingly situation is of the of forest the Rockies, consociation Pinus ponderosa premontane in recurs all condition and a similar taxifolia, climaxto thatofPseudotsuga of or there more lesssegregation consociations. is forests where plainsoccurall alongthe of fragments the desert In the mixedprairie, the one confronting Colorado as the margin preclimaxes, most extensive prairie The to it postclimax thedesert. mixed where is at thesametime Valley, The mostfrethe of it relicts thistypewhere meets trueprairie. constitutes cryptandrus, and gracilis Sporobolus by are quentexamples provided Bouteloua part. has role,grazing playedsome in all as though with theshort-grassesthis

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Phot. 7. Postclimaxof Quercus, Juglans,Ulmus,Fraxinus,etc., Canadian River,near OklahomaCity.

and Phot. 8. Postclimaxoftall-grasses, Andropogon, Calamovilfa Nebraska. Paniciumin sandhills;Thedford,

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As a generalrule,postclimaxrelictsare much moreabundant than those that represent preclimaxes, owingin the first place to the seculartrendtoward desiccationin climateand in the secondto the large numberof valleys,sandin hills and sandy plains, and escarpments the grassland especially. Postand of theirflood-plain climaxesof oak-hickory associates,elm, ash, walnut, featuresof the true and mixed prairies,holdingtheir etc., are characteristic in own farwestward major valleysbut limitedas outlierson ridgesand sandy by stretches the eastern edge. However, the compensationafforded the to as reducedto the last two is incomplete a rule and the postclimaxis typically feature where forest, woodland savannahtype.The latteris an almostuniversal or chaparraltouches grassland,owingto the fact that shrinkage under slow desiccation operates gradually upon the density and size of individuals. Savannah is derivedfromthe reductionof deciduousforest along the eastern edge of the prairie,of the aspen subclimax of the boreal forestalong the northern, and of the montane pine consociation,woodland or chaparral on and southern the western borders, recurring again on the flanksof the Sierras and Coast Ranges in California. On the south, the unique ability of the afterfire, thorniness, its mesquite(Prosopisjulifjora)to produceroot-sprouts it palatable pods and resistantseeds have permitted to producean extensive savannah that oftencloselysimulatesa true woodland climax. As would be expected,a pointis reachedin the reduction rainfall of westadequate for ward in the prairiewheresand no longeraffords compensation trees. In generalthisis alongthe isohyetof30 in. in the centerand south,and of about 20 in. in the north. Southwardfromthe parallel of 37? the further of ", shrinkage the oak savannah may be tracedin the " shinry whichdwindles fromfouror fivefeetto dwarfsonly "shin" high. With these are associated and Calamovilfa the form in gigantea.To tall-grasses, principally Andropogon the northof this line, the shin oaks are absent and the tall-grasses make a that extends into Canada, though the compensatory typical postclimax to of influence sand is still sufficient permitan abundance of such low bushes Artemisia as Amorpha, Ceanothus, filifoliaand Yucca, as well as depauperate and aspen. In the vast sandhillarea of centralNebraska,the tallhackberry whichis assumedto reflect the grasspostclimaxattainsits best development, and theirassoclimatewhen the prairieswere occupied by the Andropogons ciates some millionsof years ago. The gradual decreaseto the rainfallof the in presenthas led to the tall-grasses finding refuge all areas of edaphic compensation,not only in sand but likewiseon foothillsand in valleys, and in additionalong the frontof the deciduousforest.

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functions Community of and theirrelationto the structure functions The natureof community climax and sere have been discussedin considerabledetail elsewhere(Plant 1936), and for the Succession,1916, 1928; Plant Ecology,1929; Bio-ecology, in to presentpurposeit maywell suffice emphasizethe difference significance major or primaryand minor or secondaryfunctiohs.The former between cooperation, ecesis, reaction,competition, migration, compriseaggregation, complexes,invasion and coaction, togetherwith the resulting disoperation, upon community effect and succession. Anyone ofthesemay have a profound and of forcein the selectionand grouping life-forms but structure, the driving deals and coaction. Migration in speciesresideschiefly reaction,competition, forthe most part with the movementand evolutionof units under climatic of compulsion,and succession with the developmentand regeneration the or denuded areas. climax in bare that are concerned functions to In contrast thesestandsthe groupofminor is as with numbersand appearance or visibility it may be termed.The first annuation, in accordance with which the abundance of any species may fromdryto wet phases of the various climaticcyclesor the growth fluctuate beingcombined. not the in differ termsofprominence, two effects infrequently in moreor less extreme eitherdirection a For the grassland, season of rainfall thoughthe balance is oftenemphasizesone dominantat the expenseofothers, year, whilein the desertin particularthe redressed the following by usually swingin numberof annuals may be fromalmostcompleteabsence to seasonal again withone or fewspeciestakingthe major role. Aspectionis dominance, season, more each growing processionof societiesthrough mainlythe orderly by or less modified changesin numberensuingfromannuation. Hibernation seasonal appearance and are formsof aspection, and estivationmerelyaffect Whileusuallyappliedto the of suspension coactioneffects. withthe temporary responses of animalmembers thebiome,it is obviousthatplantsexhibitcertain Diurnationis likewisebest knownin the case of animals, of similarnature. of ones,but it is exhibitedalso by the verticalmovement especiallynocturnal the and formby the openingand closingof flowers planktonand in different of "sleep" movements leaves. species: dominants Roles of constituent werelong life-form species of characteristic The abundant and controlling ago termed dominants(Clements,1907, 1916), this propertybeing chiefly competition.In harmony determined the degreeof reactionand effective by withthe conceptof the biome,it has become desirableto considerthe role of to in is animalslikewise;sincetheirinfluence seen chiefly coactionby contrast has the reaction of plants, the term influent been applied to the important

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species of land biomes (cf. Clementsand Shelford, 1936). It is an axiom that the life-form the dominant treesstampsits character of and woodupon forest land, that of the shrub upon chaparral and desert,and the grass formon prairie,steppe and tundra. Thereare seral dominantsas well as climax ones, and these give the respectiveimpressesto the stages of prisereand subsere. Finally, there are considerabledifferences rank or territory in even among the dominants each formation. of The mostimportant those ofwide range are that bind together associationsofa climax; to thesethe termperdominant the (per,throughout) maywellbe applied. In contrast thesestandthedominants to more or less peculiar to each association,such as beech or chestnutin their respectivecommunities and Sporobolus asper in the true and Stipa comatain the mixedprairie, whicheudominant for may be employed. Subdominants different fromthat of the regularlybelong to a. life-form dominants and are subject to the controlof the latterin a highdegree,as the name indicates. They are best exemplified the perennialforbs,though by biennials and annuals may serve as seral subdominants;all three may be actual dominantsin the initial stages of succession and especially in the subsere. The term codominant has so far had no very definite status; it is hardlyneededto call attention the presenceoftwo or moredominants, to since this is the rule in all cases with the exceptionof consociationand consocies. In contrastto the types mentionedstands a large numberof secondaryor accessory species that exhibit no dominance,which may be conveniently referred as edominants, to pendingmoredetailed analysis. Influents As indicatedpreviously, designation influent applied to the animal the of is members the biome by virtueof the influence coactiontheyexertin the of or community. The significance this effect of depends much upon the life-form and to a large degreeupon the size and abundanceof the species as well,and is seen chiefly the coactions involved in food, material,and shelter. Inin fluents may be groupedin accordancewith these properties, they may be or arrangedwithrespectto distribution role in climax or sere,or to time of and appearance (Clements and Shelford, 1936). For generalpurposesit is perhaps most convenient recognizesubdivisions to similarto those fordominants and withcorresponding termsand significance. Thus, a perfluent would occurmore or less throughout the formation, while the eufluent would be more or less typicalorpeculiar an association.Subfluents to wouldmark nextlowerdegree the of importance, roughly comparableto that of subdominant, whileminuteor influents stillless significance of microscopic might wellbe knownas vefluents. Climaxand seralunits No adequate analysis of vegetationor of the biome is possible without taking full account of development. As the firststep, this involves a distinctionbetween climax communities proper and those that constitutethe

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differ comin The twogroups the stages. movement toward final successional of but and position, stability, typeof control, theyagreein the possession made differences it Theseprimary subdominants, influents. and dominants, viz. and seraland to of two desirable recognize series communities, climax to suffixes, -ation distinguished therespective by terms, propose corresponding comeintouse as thefeeling 1916).Thesehave gradually and -ies(Clements, basis a and fordynamic ecology grown bid fairto constitute permanent has finally embrace theunits all thatthey forall suchstudies.It is notsupposed to application thegreat but constant system, their necessary a complete for thattheymeet twodecadesindicates America nearly for climaxes North of of present needsin thematter analysis. category the at placedin theproper can Not all communities be certainly smalland fewofthese casesis relatively of outset, thenumber doubtful but research. and extensive intensive difficulty undercombined present serious wide an sufficiently andlong Thisstatement, however, presupposes experience of types proclimax, to permit climaxes thevarious and between distinguishing in of features subelimaxes particular. as wellas recognizing characteristic the and are overa wideregion indispensable thedifficulties studies Comparative and Whileecotones mictia willdisappear thedegree thatthisis achieved. to thesein turnare in necessarily riseto somequestions thisconnection, give as as resolved investigations extensive theyaredetailed. by units Climax of rank of In theorganization these, four types descending andimportance the consociation, weredistinguished within formation, namely, association, the two baseduponthe and itself, first were society, clan. Liketheformation whilethe last wereestablished upon the subdominant and its life-form, life-form. was recognized thetimethatthe It at and its different dominant unitsformed the dominants within itself other contained (cf. by association and faciation divisions, PlantIndicators, 1920,pp. 107,276),and twofurther weresuggested and locies, seral and with lociation, corresponding ones,facies Thesehave as for to desirability. submitted Prof. Tansley hisopinion to their in of field and beentested thecourse further studies havenowand thenbeen until1932 (cf. serieswas not published the used in print, though complete the viz. units3 1932). The climaxgroupnow comprises following Shelford, faciation, lociation,society,and clan. At the consociation, association, it to for society sociation thesakeofgreater by beginning,wasintended replace suchusagethattheidea was had in but uniformityterms, theformer attained in sense the However, useofsociety quitea different bystudents relinquished. of of socialrelations, againraisesthe question the among insects, especially in of uponbioemphasis desirability sucha substitution, viewofthegrowing ecology also Du Rietz,1930;Riibel,1930). (cf.

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Phot. 9. Associationof mixed prairie,Stipa, Agropyrum, etc.: Monument, Bouteloua, Colorado.

of Phot. io. Foothillfaciation the desert-plains Bouteloua association,

eriopoda, B. gracilis, B. hirsuta,B. fliformis,etc.: Safford,Arizona.

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Association. divisionofthe biome the Underthe climaxconceptthisrepresents primary unit of the plant the generalized from entirely and or formation, hence differs to be preferred.Each biome is for sociologists, which the term community and the of consistsregularly two or moreassociations,thoughthe lake forest to each seeming consistof one desertscrubembodytwo apparentexceptions, associationonly. However,these are readilyexplained by the fact that the has of member the former been obscuredby the expansionofmontane western whileone or moreadditionalassociations in and coast forests the north-west, of the desert climax occur to the southwardin Mexico, and apparentlyin South Americaalso. is The numberof associationsin a particularformation naturallydeterand differences thesein turndependupon the of minedby the number primary restsupon the presenceof eudominants. Just as,the unityof the formation so of wide distribution several major dominantsor perdominants, the associationis also markedby one or moredominantspeculiarto it, and oftenas in well by differences the rank and groupingof dominantsheld in common. Thus, in the true prairie association, the eudominantsare Stipa spartea, for Sporobolusasper and heterolepis; the desert plains, Bouteloua eriopoda, and radicosa and Aristida californica,while Stipa comata and rothrocki Buchloe take a similar part in the mixed prairie. In the deciduous dominantsof one associationare suppliedby the climax, the characteristic beech and hard maple, of a second by chestnut and chestnut-oak, association,of widerrange and greatercomplexity, though the oak-hickory is comparativelypoor in eudominants by contrast with the number of species. and phyleticrelationsof the associationsof a climax are The structural of whichis the mosthighlydifferentiated all best illustrated the grassland, by of its great extent. The largelyas an outcome North Americanformations, whichoccupiesa generally mostextensive and variedunitis the mixedprairie, median position with respect to the other five associations of this climax. Originally,it derived its dominants from three separate regions, Stipa, fromthe south, and Agropyrum Koeleria comingfromHolarctica,Sporobolus fromthe Mexican plateaux, and it still exhibitsthe and the short-grasses closest kinshipwith the Eurasian steppe. It contains nearly all the genera that serve as dominantsin the related associations,while many of the eudominantsof these have all the appearance of direct derivativesfromits The Poa, and Agropyrum. evolution species,as is shownby Stipa, Sporobolus, in is of both species and communities evidently responseto the various subof climates,that ofthe trueprairiebeingmoister, the coastal warmeras well; prairiemarkedby winter the California the desertplains are hotterand drier, rainfalland the Palouse by snowfall.
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Consociation. In its typical form consociation constituted a single the is by dominant, of the is butas a matter convenience term also applied casesin which to other dominants but sparingly are present and hencehave no real sharein the 'control the community. has likewise of It been convenient refer the to in abstract each majordominant theassociation a consociation, to of as though withtherealization it occurs that more frequently mixture in thanbyitself. In thissense maybe considered unitoftheassociation, it a though actual the area ofthe latter to be regarded divided is as intodefinite faciations. Consociation dominants intoa more lessregular fall or series with respect factor to requirements, especially watercontent, and often exhibit zonation conin sequence. Thisis a general feature mixedprairie of where Agropyrum smithi and Stipa comata thechief are mid-grasses, former the occupying swalesand lower slopes, latter the upper slopesand ridges. Theconsociation achieves definite expression a considerable only over area when factors the concerned fluctuate within limits bytherequirements the set ofthedominant when other or the The dominants notfound theregion. are in case may be illustrated Pinus ponderosa the lowerpart of the first by in montane forest byAdenostomafasciculatum Sierran and while inthe chaparral, is thesecond exemplified Picea engelmannitheFront by in RangeofColorado, itsusualassociate, Abieslasiocarpa, being absent from district. rolling the In like terrain thatoftheprairie, each consociation recur will in constantly the situation is necessarily proper but fragmentary nature.Such behavior in is of characteristic dominants witha postclimax tendency, with as Stipa minor and Elymus in condensatus swalesand lower levelsofthemixed prairie. Faciation. Thisis the concrete subdivision the association, entire of the area ofthe latter madeup ofthevarious being or faciations, except seralstages fragfor ments theseveral of consociations. Each faciation corresponds a particular to climateof real but smaller regional differences rainfall/evaporation in and It temperature. maybe characterized one or two eudominants, as by such HilariajamesiandStipapennata thesouthern in mixed prairie, more but often it derivesits individuality froma sorting or a recombination the out of of dominants theassociation. is evident, term formed As the is from stem the fac-,show,appear,as seen in face and facies,and the suffix -ation, which a denotes climax unit. the During pastdecade,much attention beengiven therecognition has to and limitation faciations the basis ofthepresence absenceofa euof on or such as Hilaria, Buchloe, Carex,or a changein the rankor dominant, or of grouping common dominants, Stipa,Agropyrum, like Sporobolus Bouteor this loua. In theprairie taskhas beencomplicated overgrazing, cultivation by andrelated while disturbances, selective and haveaddedto the lumbering fire

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especially. In general, appears in difficulties, the deciduousforest temperature sincetheyusually of to play the leadingpart in the differentiation faciations, fall into a sequence determinedby latitude or altitude, though rainfall/ the also. The mixedprairieexhibits largest evaporationis naturallyconcerned respect by the deciduous forestas a number,but it is approached in this dominants. Over the GreatPlains consequenceof wide extentand numerous Boutelouafromnorthto south,the successivefaciationsare Stipa-Bouteloua, Bouteloua-Buchloe, Carex,Stipa-Agropyrum-Buchloe, Hl'ilaria-Stip'a-Bouteloua, are communities to be However,the short-grass and Agropyrum-Bouteloua. have been eliminatedor regarded as disclimaxeswhereverthe mid-grasses in that fluctuates relationto dry and wet phases of the nearlyso, a condition climaticcycle. Lociation. the of In its turn,the lociationis the subdivision the faciation, termbeing derivedfromlocus,place, as indicatinga generallocalityratherthan a large a region. Nevertheless, lociationmay occupy a relativelyextensiveterritory withseveralhundred milesor morein extent,by comparison up to a hundred in by forthe faciation. It is characterized more or less local differences the abundance and groupingof two or more dominantsof the faciation.These or slope-exposure altitude, in variations soil,contour, to correspond considerable lociations concerned.As a consequence, the but all within limitsofthefaciation witheach other, hereand thereas alternes recurring are veryoften fragmented, of and frequently withproclimaxes varioustypes.Like mostclimaxunits,they have been modifiedby disturbancein some degree,and this fact must be subelimaxor themfrom kept in mindin the task of distinguishing constantly disclimax. to of A detailed knowledgeof the faciationis prerequisite the recognition the various lociationsin it. The numberfor a particularfaciationnaturally of concerned. dependsupon the extentofthelatterand the number dominants lociations are more numerousin the faciationsof the mixed Consequently, forest. As prairie and desert plains, of the chaparral and the oak-hickory would be expected,they are oftenmost distinctin ecotonesand in districts with of wherethereis local intrusion anotherdominant. In correspondence or diminishsuperimposed theirlocal character,it is importantto eliminate of differences throughrestoration the originalcover by means of protection enclosures and thus renderit possibleto disclosethe true composition. Society. This termhas had a wide range of application,but by dynamicecologists of it has generallybeen employedfor various groupingsof subdominants, whichthose constituted aspects or by layers are the most important. In by in formed cryptogams the by addition,thereis a host of minorcommunities a and othermatrices.The soil itselfrepresents groundlayeror on host-plants
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Nature and Structureof the Climax

major layer,divisibleinto moreor less definite sublayers. Animalsregularly assume roles of varyingimportancein all of these, especially the insects, arachnids and crustacea, and hence most if not all societiescompriseboth subdominant plants and subinfluent animals. It is doubtful whether animals form true societies independentlyof their food-plantsor those used for materialsor shelter,but this is a questionthat can be answeredonly after the simplestunits, namely, family and colony, have been recognizedand in coordinated termsof theircoactions. In view of what has been said previously, seems desirableto employ it society as the general term for all communities subdominantsand subof above the rank of familyand colony, much as community the is influents inclusivetermforall groupings whatsoever of rank.This thenpermits carrying made two decades ago that the major typesof societiesbe out the suggestion set apart by distinctive names. In accordance,it is here proposedto call the aspect societya sociationand the layer societya lamiation, while the correspondingseral terms would be socies and lamies. Many of the societies of and minuteanimals would findtheirplace in these,particularly cryptogams so for those of the surfaceand soil layers,but many otherstake part in a sereor serule, such as that of a moldering and may best receive miniature log, that suggestthis relation. designations a societiesare welldeveloped,theyregularly Sociation. Wherever manifest seasonal sequence, producingwhat have long been known as fairlydefinite aspects (Pound and Clements,1898). As phenomenaof the growing season, as these were firstdistinguished early spring or prevernal,vernal proper, estival, and serotinalor autumnal,but there may also be a hiemal aspect, especiallyfor animals, in correspondence with an actual and not merelya calendarwinteras in California. Sociationsare determined primarily the relationbetweenthe lifecycle by and the seasonal marchof directfactors, of the subdominants in temperature faras the matrixofplantsis concerned, constituent particular. So the species may be in evidencethroughout season, but theygive character it only the to duringthe periodof flowering, fruiting the case of cryptogams. or in They are presentlargelyor whollyby sufferance the dominants, of and they are to be related to the reactionsof these and competition among themselvesrather more than to the habitat factorsas such. In grasslandand desert,they are often more strikingthan the dominants themselves,sometimesowing to as stature but chiefly an effect color and abundance, and they may also of in attain muchprominence woods withthe canopy not too dense. Sociations are usually most conspicuousand best developedin grassland, fouror even fivedistinct aspectsoccurring thetrueprairie in from earlyspring to autumn. In the mixed prairiethese are usually reduced to three,and in the desertplains and desertproper,to two major ones, summerand winter, in which howeverthere may be subaspects marked by sations,as indicated

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eS

Phot. ii.

SociationofErigeron and Psoralea,estivalaspect; Belmont Prairie, Lincoln,Nebraska.

Phot. I2. Lamiationofmid-herbs, Laportea, Physostegia, Impatiens, etc., in Oak-Hickory forest. CLEMENTS-NATURE
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less opporlatitude or altitude afford later. Short seasons due to increasing tunity,and the tundra, both alpine and arctic, usually exhibits but two role as in the prairie. In aspects,the sociationshowevertakinga conspicuous of and character the sociationadepend largelyupon the woodlandthe number regularly sociations the natureofthe canopy,and fordeciduousforest flowery developing belongto the springand autumnaspects,whenthe foliageis either or disappearing. It is convenient distinguish to sociationsas simpleor mixedwithrespectto of the plant matrixin accordancewiththe presenceofa singlesubdominant,or such a two or more. However, when animals are includedin the grouping, appears misleadingand may well be dropped.The word "mixed" distinction applied to plant-animalsocietieswereit not foTthe would be more properly factthat thisappears to be the universalcondition. Since seasonal insectsare in manyofthesocieties whichtheytake partare bestdenotedas sations. legion, the Lamiation. The termfora layersocietyis derivedfrom stemlam-,seen in lamina and lamella. As is well known,layersare best developedin forests so witha canopyofmediumdensity, that underthe mostfavorableconditions as many as fiveor six may be recognizedabove the soil. In such instances, muchinterrupted, often an stories, upperand lower, are there usuallytwoshrub of followedby tall, medium and low forblayers,and a groundcommunity mosses,lichensand otherfungi,and usually some delicate annuals (cf. Hult, 1894). The soil populationis perhapsbest treatedas a single 1881; Grevillius, sublayers.When the various unit,thoughit may exhibitmoreor less definite each is regardedas a lamiation,but are layersbeneaththe dominants distinct, designation, to organized warrant in manycases onlyone ortwo are sufficiently lamiation. e.g. shrubor tall-herb Layers are oftenreducedto a singlelamiationof low herbsin the climax lackingin dense and even this may be entirely forest, especiallyof conifers, chaparral. Two or three layers of forbs may be presentin true prairie in and oftenconthe upper lamiationbeingmuchthe most definite particular, of in cealingthe grassdominants the estivalaspect,but the structure grassland of the reflects greater importance sociations. Climaxesof sagebrush generally and desertscrubexhibitno properlamiations,owingto the intervalbetween show but the individualdominants, the herbaceoussocietiesofthe interspaces of something this nature. As a rule, well-developedlamiations also manifesta seasonal rhythm, to corresponding aspects or to subaspects. These constituterecognizable and in groupings the lamiationand forthe sake ofdetermination analysismay be termedsations.This wordis a doublet of season, both being derivedfrom of interplay the root sa-, sow, hence growor appear. Because of the frequent aspects and layers,the sation may forthe presentbe employedforthe subdivisionofboth sociationand lamiation,and especiallywhereseasonal species of invertebrates play a conspicuousrole.

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Nature and Structure the Climax of

Clan. This is a smallcommunity subordinate but of importance commonly of distinctive character. It is markedby a densitythat excludesall or nearly all competingspecies,in consequence of types of propagationthat agree in the possessionof short offshoots. Extensionis usuallyby bulb, corm,tuber, stolonor shortrootstock, each ofwhichproducesa moreor less definite family grouping;in fact,most clans are familiesdevelopedin the climax matrixand sometimes witha blurred outlinein consequence. Clans ofa particular species such as Delphiniumazureumor Solidago mollis are dotted throughout the respectivesociation, often in large numbers,and contributea distinctive impress muchbeyondtheirabundance. Like all units, smallonesespecially, but theyare subjectto muchfluctuation withthe climatic cycle,as a resultofwhich theymay pass into societiesor be formed the shrinkage the latter. by of Seral units. The conceptsof dominanceand subdominance apply to the sereas theydo to the climax, as does that of influence also, and the corresponding sets of units bear the same generalrelationto each other. Each of the fourmajor units is the developmentalequivalent of a similarcommunity the climax in seriesand thisis likewisetrueofthe variouskindsofsocieties.They constitute the successivestages of each sere,both primary and secondary, the including subclimax,wherethey oftenachieve theirbest expression. It has also been customaryto employ seral terms for preclimax and postclimax,and this appears to be the betterusage fordisclimaxand proclimaxes general.From in the fragmentary natureof bare areas and suitablewaterbodies in particular, seral communities oftenbut partiallydeveloped and one or more units are will be lackingin consequence. Thus, the reed-swampassocies is frequently represented a single one of its several consociesand the minorunits are by even morecommonly absent. The associes is the major unit of everysere,the numberbeing relatively large in the prisereand small in the subsere.The universaland best understood examples are those of the hydrosere, which Lemna, Potamogeton, in Nuphar, Nymphaea,Nelumboand othersformthe consocies of the floating stage, and Scirpus, Typha, Phragmites, etc., are the dominantsof the reedswampor amphibious associes. As alreadyindicated, every consocies mayoccur singlyand oftendoes whenthe habitat offers just the properconditions it for or the othershave failedto reach the particularspot. When the ecial rangeis wider,various combinationsof two or three dominantswill appear, to constitutecorresponding facies. Locies are less definitely markedas a rule,except in swampsof vast extent,but are to be recognized the abundance of reedby like dominantsof lower stature, belongingto other species of Scirpus, to Heleocharis, Juncus,etc. Both faciesand locies seemto be betterdevelopedin sedge-swampwith its largernumberof dominants,though the Everglades withthe singleconsociesof Cladiumforma striking exception.

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United States containa considerable of The tree-swamps the south-eastern Nyssa aquatica,bijioraand numberof consocies,such as Taxodiumdistichum, Magogeche, Carya aquatica,Planera aquatica, Persea palustrisand borbonia, and caroliniana,and Quercus profunda nolia virginiana, Fraxinus pauciflora, facies,thougha more nigra.These are variouslycombinedin severaldifferent detailed and exact study of the swamp sere may show the presenceof two by woody associes,distinguished the depth or durationof the water. As with the otherscrub communities, heath associes of peat-bog and muskeag comwealth of prises a large numberof dominantsand presentsa corresponding faciesand locies. the In the hydrosere the deciduousforest, typicalsubelimaxis that of of associes, composed of species of Quercus,Ulmus, Fraxinus, the flood-plain Platanus,Liquidambar, Celtis, Populus and Salix forthe Acer,Betula,Juglans, facies, namely,northern, most part. There are at least three well-marked locies. The each with a numberof moreor less distinct centraland southern, consistsof Larix swamp associes or subelimaxof the lake and boreal forests oftenas consocies occurring laricina, Picea mariana and Thuja occidentalis, subclimaxes in but generally the formof zoned facies. A large numberof fire appear in the formof consocies,as with many of the pines,but associes are frequentalong the Atlantic Coast, as they are in the boreal climax, where concerned.The numberof shrubs and small aspens and birchesare chiefly treesthat play the part of seral dominantsin the deciduous climax is much a not larger, producing merely wide rangeofassociesbut offaciesand lociesas well. Morethan a dozen generaand a scoreor so ofspeciesare involved,chief Asimina,Hamamelis,Prunus, llex, among them being Sassafras,Diospyrus, for Crataegusand Robinia. The subclimaxof the xerosereis constituted the and a most part by species of Quercus,forming eastern,a south-eastern an facies, one of stellataand westernassocies, the last with two well-marked and Carya ovata. the marilandica, otherof macrocarpa those of grasslandand scrub possess a large Amongpostclimaxassocies, varietyoffaciesand locies, numberof dominants and exhibita corresponding withfairly consocies. In the sandhillsof Nebraska,the talldefinite together hatli,furcatus and nutans, Calamovilfa grasses concernedare Andropogon and Panicumvirgatum; some trichodes, Elymuscanadensis Eragrostis longifolia, and at ofthesedrop out to the northward othersto the south,thus producing least three regionalfacies. The mesquite-acaciaassocies of the south-west a possesses a largernumberof dominantsand manifests greatervarietyof its faciesthrough wide area, and this is likewisetrue of the coastal sagebrush of California. to to With reference seral societies,it must suffice pointout that theseare of necessitypoorly developed in the initial stages of both hydrosereand few. Even in the reed-swamp, true as xerosere, the dominantsare relatively to layers are the exception,being largelyrestricted such subdominantsas

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Nature and Strueture the Climax of

in Alisma,Pontederia, Hydrocotyle Sagittaria, and whichare foundmostly the borders intervals. and However, extensive in subelimaxes postelimaxes and situation quitedifferent. tall-grass is The is associes sandhills often of quiteas rich saties lamies thetrue in of and as prairie, while various the subelimaxes the several great forest types mayequal thelatter thewealth subdominants of in for each season and layer,the actual communities beingverymuchthe same. Serule. of Thisterm, diminutive sere, beenemployed a great a of for has variety within course miniature successions run short somewhat but that their complex thecontrol a majorcommunity, of the and especially climax subelimax. They of resemble ordinary seresin arising bare spotsor on matrices different in etc. Parasitesand sorts,such as earth,duff, litter, rocks, logs,cadavers, role saprophytes a prominent often play and exclusive inthem, plants and and animalsmay alternate the dominant in parts.The organisms rangefrom to on microscopic bacteria worms mites, and larvaeandimagoes theonehand and largefleshy shelf on The of and fungi theother. mostimportant thesein termsof coactionand abundance knownas dominules are and (Clements as subdominule edominule terms thetwodegrees and for Shelford, 1936),with oflesser importance. thesamemodel formed are On associule, consociule, and in sociule general of correspondence theunits thesereitself.In addition with there families colonies these are and of minute which essentially organisms, are similar those theinitial to of stages themajor of succession. tothepresent, Up little attention beendevoted thedevelopment structure serules, has to and of but theyare coming receive to in adequateconsideration connection with bio-ecological problems. Manyofthecoactions, however, havelongbeenthe subjectofdetailed in research theconversion organic of materials.
RANK AND CORRESPONDENCE OF UNITS

Thefollowing exhibits actualunits climax sere, wellas the table of and as their with for correspondence eachother.However, thecomplete accurate and of climax especially continental ofvegetation, and the analysis a great mass it is necessary invokeother to that of the proclimax of concepts, chiefly and communities mixedin space or in time.The several proclimaxes have been characterized 262-8), theecocline seration and (pp. and briefly defined 267). (p. To these to be addedtheecotone mictium, terms longstanding, are and both of theformer applied themixing dominants to of between units, latter two the to the mixedcommunity intervenes that between two seralstagesor associes. Finally,therewill be the severaltypesof seresin all possiblestagesof the in development, prisere the formof hydrosere, xerosere, halosereor in less psammosere regions disturbed a myriad subseres thoselong and of in settled.

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TABLE OF CLIMAX AND SERAL UNITS Climax Association Consociation Faciation Lociation Sociation Lamiation Sation Clan Sere Associes Consocies Facies Locies Socies Lamies Saties Colony Family

Climax (formation)

Serule Associule Consociule Sociule

as is As indicatedpreviously, wordcommunity employed a generalterm the may well be used to units,whilesociety to designateany or all ofthe preceding i.e. includethoseofthe seconddivision, sociation,etc. These are characterized group. It in by subdominants contrastto the dominantsthat markthe first has also been pointedout that the entirearea ofthe associationis dividedinto of local expression comfaciationsand that the consociationis the relatively completedominanceon the part of a singlespecies. The clan plete or nearly the corresponds the familyas a rule,but in some cases resembles colonyin to beingformed two species. by but Families and coloniesmay also appear in climax communities, this is withthe serule. regularly connection in Panclimaxand eoclimax The comprehensive treatmentof these conceptsis reservedfor the sucthem ceeding paper in the presentseries,but it is desirableto characterize the meanwhile. The panclimax(wav,all, whole) comprises two or morerelated the same lifeof climaxes or formations the same generalclimaticfeatures, is and commongeneraofdominants. The relationship regardedas due to form theiroriginfroman ancestralclimax or eoclimax(qcos', dawn), of Tertiaryor even earliertime, as a consequence of continentalemergenceand climatic In a differentiation. the past, eoclimaxesformed seriesofgreatbioticzones in withthe pole as a focus,and this zonal disposition the northern hemisphere of or clisereis still largely evident in the arrangement panclimaxes at the in present. It is striking the case of the arctic tundra and taiga or boreal and somewhat forest, fairlyevidentfordeciduous forestand prairie-steppe, whilethe positionof desertsis obscureforwoodland and chaparral-macchia, mountainranges. This is true likewiseof by largelydetermined intervening broad land connection grasslandin some degree,and taken withthe former betweenNorthand South Americaexplainswhyboth prairieand desertpanclimax containat least one australformation.

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of Nature and Structure the Climax

understood that it In the lightof whathas been said earlier, is readily as and are and panclimax panformation exactsynonyms, are eoclimax eoforthe terms are and mation.Panbiome eobiome thecorresponding when biotic research. thebasisfor is community takenas in to Prerequisitesresearch climaxes to in superfluous statethatthe majordifficulty the It wouldbe entirely the were is of analysis vegetation its complexity, it notfor factthatit is too in This viewpoint. was embodied the for takenas thewarrant thestatic often were communities assembled as original oftheformation a unitin which idea and composition phyletic of irrespectivegeneric basis, ona physiognomic quite should have thatthisviewand its corollaries It relationship. is notstrange whathappened since is of longpastitsperiod usefulness, this exactly persisted fully has but of system Linnaeus, thetime cometo recognize with artificial the upon mustbe builtjust as certainly of system communities that a natural families. Comthatofplant as relationship must and development consequent in it, thanagainst and especially view for is plexity an argument thisrather whenthetouchpattern a discloses definite ofthefactthatthe complexity to is stoneofdevelopment applied it. kaleidoscope to may of the Though mosaic vegetation appear be a veritable by wrought uponit are readily long in countries occupied man,thechanges the previously, As concerned. emphasized of in intelligible terms theprocesses thatcorrespond scale in is control thatofclimate, a descending ofunits primary are pattern wrought general Uponthis and association, faciation. toformation, and and of effects physiography soil,andbothclimatic themore circumscribed a with obscured or more lesscompletely and are edaphic figures overlaid often kinds.Even abovethismaybe of veneer appliedby disturbance all possible of apparent, such recurrent but transient nonetheless discerned effect, the the of pattern climate Moreover, orderly and as changes annuation aspection. and as suchclimaxes tundra so ranges that mountain is complicated great by is varied the by zone,andtheeffect further their proper far taigaoccur beyond of relative position theaxis. of source fragmentary in Themigrations climaxes thepastarea prolific of of in is except terms dynamics. the of relicts, interpretationwhich impossible the of and which represents shrinkage forest Thisis likewise ofsavannah, true or modified fire further by and is thenusually climate a scrubunder drying of causesis characteristic every from Fragmentation thisand other grazing. enters the utilization when human its and diversified terrain reaches maximum not in is though in process similar effect sceneupona largescale. Somewhat with consequence the that of of by thereduction number dominants distance, of into maybe converted a consociation one. Sucha an association several of also to climateand physiography, bearssomerelation naturally shrinkage in its and finds bestillustration the as especially seenin the glacialperiod,

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generalpovertyof dominantsin the coniferous and deciduous climaxes of Europe,by contrast withthoseof easternAsia and NorthAmerica. A similar contrastobtains betweenthe grasslandof Asia and of North America,the latterbeingmuchricher dominants, in whileSouth Americaapproximates it closelyin thisrespect. On the part of the investigator, difficulties the way of an extensive the in and thoroughgoing study of climaxes are usually more serious. They arise partlyfromthe handicap too oftenset by state or national boundariesand partlyfrom limitations fundsand time.They are also not unrelatedto the of the fact that it is easiest to know a small district well and to assume that it reflects largerones withmuchfidelity. a consequence, is impossible lay As it to too muchstressupon the needforcombining intensive and extensive methods in the research upon climaxes,insofar theirnature,limitsand structure as are concerned. The detaileddevelopment termsof primary in and secondary successionlends itselfmuch morereadilyto local or regionalinvestigation, but even here a widerperspective essentialto accurategeneralization. is
REFERENCES Bradbury, J. "Travels in the Interior NorthAmerica, 1815." In Thwaites' Early Western of ca. Travels, 1904. 5, Bromley, S. W. "The original forest typesofsouthern New England." Ecol. Mon.5,61-89,1935, Chaney, R. W. "A comparativestudy of the Bridge Creek flora and the modernredwood forest." Publ. Carneg.Instn,No. 349, 1925. Oregon." Publ. Chaney, R. W. and E. I. Sanborn. "The Goshen Flora of West-central Carneg.Instn,No. 439, 1933. Chapman, H. H. "Is the longleaf type a climax?" Ecology, 328-34, 1932. 13, Clements, F. E. Plant Physiology Ecology, and New Yoik, 1907. Clements, F. E. Plant Succession, Washington, 1916. Clements, F. E. "Development and structure the biome." Ecol. Soc. Abs. 1916. of Clements, F. E. Plant Indicators, Washington, 1920. Clements, F. E. "Phylogenyand classification climaxes." Yearb. Carneg.Instn,24, 334-5, of 1925. Clements, F. E. Plant Successionand Indicators, New York, 1928. Clements, F. E. "The relictmethodin dynamicecology." This JOURN. 22, 39-68, 1934. Clements, F. E. " Origin the desertclimateand climaxin NorthAmerica." This JOURN. 24, of 1936. Clements, F. E. and E. S. Clements. "Climate and climax." Yearb.Carneg.Instn,32, 203, 1933. Clements, F. E. and V. E. Shelford. Bio-ecology, 1936. Bot. Tid. 24,489, 1930. Du Rietz, G.E. "Classification nomenclature vegetation."Svensk. and of Godwin, H. "The subclimaxand deflected succession." This JOIURN. 17, 144, 1929. Greeley, H. An Overland Journey California 1859,New York, 1860. to in Grevillius, A. Y. "Biologisch-physiologische Untersuchungen einiger SchwedischenHainthiilchen." Bot. Z. 52, 147-68, 1894. Hult, R. "Fors6k til analytiskbehandling viLxformationerna." af Medd. Soc. Faun. Flor. Fenn. 8, 1881. Medd. utvecklingshistorie." Hult, R. " Blekingesvegetation. Ett bidragtill viaxformationernas Soc. Faun. Flor. Fenn. 12, 161, 1885. Bot. Zbl. 27, 192, 1888. Journals the of Lewis Lewis, M. and W. Clark. Journal, 1803-1806.In Thwaites' TheOriginal and ClarkExpedition, 1904-05. Phillips, J. "The biotic community."This JOuRN. 19, 1-24, 1931.

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