Society of Systematic Biologists The Shape of Things to Come Author(s): Stephen Jay Gould Source: Systematic Zoology, Vol. 22, No. 4 (Dec., 1973), pp. 401-404 Published by: Taylor & Francis, Ltd. for the Society of Systematic Biologists Stable URL: http://www.jstor.org/stable/2412947 Accessed: 22-04-2015 16:08 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Taylor & Francis, Ltd. and Society of Systematic Biologists are collaborating with JSTOR to digitize, preserve and extend access to Systematic Zoology. http://www.jstor.org This content downloaded from 134.84.2.243 on Wed, 22 Apr 2015 16:08:30 UTC All use subject to JSTOR Terms and Conditions I. C. S. E. B. BOULDER-AUGUST 1973 SYMPOSIUM "EVOLUTIONARY DEVELOPMENT OF FORM AND SYMMETRY" CONVENOR-STEPHEN JAYGOULD THE SHAPE OF THINGS TO COME STEPHEN JAY GouLD Museumof ComparativeZoology HarvardUniversity Cambridge,Mass. 02138 but it mayfail a quiet certainlevel of generality; in thissymposium, We celebrate, oldestsubject.Dur- badlyin trying ofbiology's renaissance to explainboththeparticuin African ingthepastdecade,likea thiefin thenight, lar case of explosivediversity becomein- lake fishesand such generalproblemsas has surreptitiously morphology again. The descriptiveconcerns the increaseof organicdiversitythrough teresting by theirjargon time-(under billiardball models,each constrained of cataloguers, witha fewotherspecial- speciesis a blackboxofno uniqueconstructo communication orat leastaccommodating,tionand the causes of increasingdiversity ists,areyielding, approaches. throughtimemustbe soughtin such ento analyticaland explanatory parametersas increasingenThereare severalreasonsforthis: A pan- vironmental driftrather have demicity techniques and throughcontinental oplyofnewmachinery extendedthe bounds of both perception thanin classical,and basicallysound,noof structure bi- tionsof increasingcomplexity (the SEM) and analysis(multivariate ometry).But no machinecan matchthe in majorgroups,vide Vermeijand Stebfinestcomputerof themall. Evolutionarybins). Liem showselegantlythata small change (the shiftin insertheoryhas been so vigorousof late thatit morphological could scarcelyavoid a mutualrelationshiptionof the fourthlevatorexterni)and its correlate(the use ofpharyngeal with morphology-bysupplyingit with functional offoodand freeingof ideas to catalyzethe advanceof explana- jaws forpreparation alone) has trigjaws forgathering toryproceduresand by drawinginsights anterior radiationof Afrifromit in return.Thus, we findStanley geredthe trulyfantastic theories ofecologyto can cichlidfishes.I believe,in short,that usingthecompetition a true"scienceofform" elucidatean old dilemmaof morphologicalwe areapproaching diversity (patternsof radiationin "conser- (Gould, 1970and 1971)-a claimthathas sinceCuvative"vs. "dynamic"groups); while, in notbeen madewithjustification thereturn,Liem remindsecologiststhat the vier'stime(Haeckel had an historical are oftento be ory,D'ArcyThompsona functionalone: causesof relativediversity must scienceof morphology soughtnot in the favoredenvironmentala satisfactory ofstability ofhabi- integrateboth). anddiversity parameters werepreof morphological The papersof thissymposium tat but in differences adaptation."Billiardball"ecology,likethe sentedon August7, 1973at thefirstICSEB of ideal gases,has its place at a meetingin Boulder,Colorado. We regret modelling 401 This content downloaded from 134.84.2.243 on Wed, 22 Apr 2015 16:08:30 UTC All use subject to JSTOR Terms and Conditions 402 SYSTEMATIC ZOOLOGY that the limitationsof this venerable me- methodology. Vermeij and Stebbins trace dium cannot fullyconvey both the beauty patternsin diversityto morphologicalconand excitementthatfilmsand colored slides straints: (Vermeij attributesevolutionary to "complexadded to the presentations. increasein potentialversatility In organizingthe symposium,I avoided ity"as measuredby numbersof parameters of form; "state of the art" and "contemporaryprob- needed to specifythe construction lems in . . ." talks as obvious magnetsfor Stebbinstracespatternsof diversityin mulpotboilers,and opted instead for a set of tinucleate organisms to the preadaptive exemplifications-ofnew techniques, par- featuresof unicellular eukaryotes,primarticularlyinterestingor importantcase stud- ily to polarity,nature of the cell covering, and cell shape ies, and general evolutionaryhypotheses. autotrophyvs. heterotrophy, This leads to a certainincoherence;but this and size). Stanley emphasizes ecological may,itself,be a markof virility(any rigid constraintsin arguing that the exuberant portrayal of this unformed renaissance and rapid rates of diversificationin mamwould, in any event, be as gratuitousas mals vs. bivalves are primarilyrelated to Ichthyostegadiscoursingin Devonian times more intense interspecific competition on its futureestate). The sequence of pre- among the mammals. I have prefaced these papers with the sentationsis not, however,utterlyhaphazard. The firstpapers are case studiesusing manifestoof an architect (and sensitive both deductiveproceduresof physicalanal- naturalist) because Stevens shares D'Arcy ysis (Liem, Kokshaysky) and inductive Thompson's vision that the biological scimethods of multivariate biometry (Ox- ence of morphologycan expand well benard). They displaymanyof the new tech- yond the confinesof our discipline to enniques alluded to above (Liem on the SEM, lightenand be a part of the wider science electromyography and cineradiography; of form that applies to all objects. We and rarelyrecognizehow our own laws of form Oxnard on multivariatemorphometrics experimentalstress analysis). They treat regulate the properties of buildings, maparticularlyinterestingor importantcases: chines and planets as well. To choose a Oxnard reaches some ratherstartlingcon- simple-mindedexample: medievalchurches clusions on the morphologicaluniqueness are excellentexemplarsof the laws of form of australopithecines;Kokshayskyintrodu- in correlationsbetween size and shape-for ces us to the Soviet school of evolutionary they were built in an enormousrange of morphologyin a carefullychosen natural sizes to serve fairlysimilarpurposes before experimenton size-required allometryin technologicalinventions(fromsteel girders the dynamics and morphology of wing to electriclightingto air conditioning)alslots in four species of herons. Liem con- lowed architectsto circumventmany considers the most outstandingcase of explo- straintsimposed by increasing size. The sive evolutionin modern vertebrates. Sei- law of surfacesand volumesregulatesmuch lacher's paper is transitional: he finds the of the external and internal structureof same structuresas adaptationsfor burrow- large organisms: a remarkable array of ing in taxa of severalphyla and, notingthat featurescan be explained functionallyby the same constructional principlesare used the need to increase surfaceallometrically to build featuresof such diverse function in orderto allow something(food, oxygen, as ammonitesepta and the "windows" of light) to penetrateinto the interiorof large Corculum,he presentsa strikinghypothesis organisms. Churches are subject to the on the developmental basis of adaptive same constraints. Light must penetrate form.The last threepapers presentspecific throughwindows of the peripheryand ilhypotheseson the general developmentof luminate the entireinternalarea (candles formthroughtime. They are differentin do have their limitations). Peripheryinin creases as L, area of floorplan as L2; large approach and even slightlycontradictory This content downloaded from 134.84.2.243 on Wed, 22 Apr 2015 16:08:30 UTC All use subject to JSTOR Terms and Conditions I.C.S.E.B.-FORM 403 AND SYMMETRY -X --l-*. 9S ~- p --#se9e e ; Feet 0 o 0o 20 3O o Feet ilometres LittleTey. Essex. FIG. 1.-Ground plan of the parish churchof Little Tey, Essex. From Clapham, 1934. churches cannot maintain the same shape as small ones and receive sufficientillumination. This is especially true for Romanesque churcheswith theirmassive masonry and limitedpossibilitiesfor windows; (the inventionof the flyingbuttressreduced the need formassiveouterwalls and permitted the remarkable increase in window area and "lightening"that characterizes later Gothic architecture). In order to maintain sufficientperiphery,large churcheshad to be built as more elongated rectanglesand had to introduce"outpouchings"in the form of transeptsand chapels (the formof the cross may have dictated the shape and positionof transepts,but laws of size required their presence). Enlarge the tiny parish churchof Little Tey (Fig. 1) to the size of Norwich Cathedral (Fig. 2) and the interiorwould have been darkerthanthe deeds of Herod; reduce Norwich to the size of Little Tey and the Christ child himself could scarcelyhave squeezed into the apsidal chapels. A plot of square root of area vs. peripheryforall floorplans of post-conquest Romanesque churchesof Britaindepicted in Clapham (1934) illustratesthe positive allometryof periphery. The isometric line of slope 1, drawn throughan average small church, misses all intermediate and large churchesby a wide margin. NiORWICII CATIMDRAt FIG. 2.-Ground plan of Norwich Cathedral. From Clapham, 1934. The log correlationof .995 astoundedme-I would have thoughtthat the vagaries of human will would have imposed more scatter than the centrifugaladaptation of organisms. Medieval buildershad theirrules of thumb,but thereis no evidence thatthey knew anythingof these laws of form. I can only end, therefore,by repeating D'Arcy Thompson's hope (1942, p. 1026) that biology might form the central core for a trulygeneral science of form: "Our own study of organic form,which we call by Goethe's name of Morphology,is but a portion of that wider Science of Form which deals with the forms assumed by matter under all aspects and conditions, IsO a=l 100 a=.846 75. WI0-. 25 '5 0 50 1OO 200 300 400500 P E R I P H ERY FIG. 3.-Plot, on logarithmicscales, of square rootof area vs. peripheryforall groundplans of post-conquestRomanesque churches in Britain givenin Clapham,1934. Data compiledby author using map reader and polar planimeterdirectly fromscaled drawingsof Clapham. This content downloaded from 134.84.2.243 on Wed, 22 Apr 2015 16:08:30 UTC All use subject to JSTOR Terms and Conditions 404 SYSTEMATIC and, in a stillwidersense,withformswhich are theoreticallyimaginable." ZOOLOGY GOULD, S. J. 1970. Evolutionarypaleontology and the science of form. Earth Sci. Rev. 6: 77-119. GOULD, S. J. 1971. D'Arcy Thompsonand the REFERENCES scienceof form.New LiteraryHistory2:229258. CLAPHAM, A. W. 1934. English Romanesque afterthe conquest. Oxford,Clar- THOMPSON, architecture D. W. 1942. On growthand form. CambridgeUniv. Press,1116 pp. endonPress,180 pp. This content downloaded from 134.84.2.243 on Wed, 22 Apr 2015 16:08:30 UTC All use subject to JSTOR Terms and Conditions
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