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
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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
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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
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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.
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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.
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