Dollo on Dollo's Law: Irreversibility and

the Status of Evolutionary Laws

S T E P H E N :lAY G O U L D

Museum of Comparative Zoology, Harvard University

Cambridge, Massachusetts

I. DOLLO'S F O R M U L A T I O N OF DOLLO'S LAW

Lrr~versibilit6: je s u i s b i e n t r a n q u i l l e s u r l ' a v e n i r et r u t i l i t 6 de cette
n o t i o n : s e u l e m e n t , p o u r l a souterfir o u p o u r l a c o m b a t t r e , il f a u t b i e n l a
c o m p r e n d r e , ce q u i n ' a r r i v e p a s t o u j o u r s !

L. Dollo in letter to T. Edinger, July 9, 1927

Othenio Abel l a u n c h e d Palaeobiologica with a wish t h a t the
ideas of Louis Dollo m i g h t flourish and bring prosperity to the
n e w journal. 1 Palaeobiologica did not survive the w a r ; Dollo's
n a m e lives as a m a s t h e a d to the l a w of irreversibflity, but his
forgotten work presents this notion in a fashion altogether
different f r o m the f o r m u l a t i o n s of our textbooks. Abel's j o u r n a l
m a y h a v e m e t a kinder fate. Dollo's law, moreover, h a s fallen
into disrepute along with the entire enterprise t h a t sought to
abstract historical laws f r o m the p h e n o m e n a of phylogeny. I
find this u n f o r t u n a t e for two r e a s o n s :
1. A p a r t f r o m a n y j u d g m e n t on the m e r i t of Dollo's law, I
regret this foreclosure of discussion since the debate on historical
laws illuminates so m a n y issues in the philosophy of biology
(reductionism, the n a t u r e of history).
2. Irreversibility, in its m o s t i m p o r t a n t sense, is a notion
quite different f r o m the s t a n d a r d set of such ' l a w s " those
n a m e d for Cope, Wflliston, etc. By a n ironic twist, as we shall
see, "Dollo's law" emerges as a particularized s t a t e m e n t of the
1. O t h e n i o Abel, "Die F e s t g a b e d e r "Palaeobiologica,' " Palaeobiologica, 1
(1928), 1-8.

Journal of the History of Biology, vol. 3, n o . 2 (Fall 1970), p p . 1 8 9 - 2 1 2 .

189

S T E P H E N J A Y GOULD

general reason for our rejection of an approach to phylogeny

based on a search for such historical laws.
Louis Dollo ( 1 8 5 7 - 1 9 3 1 ) was a m u c h misunderstood m a n .
Born and educated in Lille, he b e g a n a lifetime career at the
Brussels M u s e u m in 1889. a f t e r a brief stint as a m i n i n g engineer.
His open support of G e r m a n y during the First World W a r precluded any f u t u r e popularity with his Belgian colleagues; yet
with a stubbornness that he attributed to his Breton ancestry,
he r e m a i n e d at his post as a virtual recluse. While he m a i n t a i n e d
a f e w loyal friends and a uniquely high status in the paleontological c o m m u n i t y of Europe, his seclusion and avoidance of
the politics of scientific societies fostered his reputation as a
quietly dedicated, coldly dispassionate scientist. All that he hid
f r o m his colleagues he disclosed in the extraordinary correspondence with Dr. Tflly Edinger, f o u n d after the latter's
death in 1967; here, interspersed with W a g n e r i a n quotations of
d e a t h and yearning, we find the words of a lonely a n d t o r m e n t e d
idealist.
As he concealed his feelings by force of personality, so also did
he withhold his ideas by habits of writing. He wrote neither text
nor review article, and we paleontologists h a v e forgotten that his
paldontologie dthologique was the source for a type of research
t h a t we all pursue t o d a y - - t h e study of adaptation in reIationship
to environment. He wrote no discursive prose, no elaboration of
general ideas, but listed his contentions only as sets of s u m m a r y
propositions. The quoted source for all his evolutionary theorizing is a two p a g e r6sum~ in the Proc~s-Vebaux of 1893 for the
geological society of Belgium ( t r a n s l a t e d as a n appendix to this
article); rarely did he elaborate a n y f u r t h e r in his later works.
He wrote to Edinger of his views on scientific prose (September 21, 1929): "DoUo's style, a telegraphic style--difficult to
r e a d . . . Yes, but clear, brief, precise--that is m y purpose!
The consequence of a strong m a t h e m a t i c a l education! An original m e m o i r is not a storyl" These e x a m p l e s of his published
s t a t e m e n t s on irreversibflity are typical:
A Dollo epitome: 2
Nautilus does not yet h a v e fins; Octopus h a s t h e m no longer.
F r o m this point of view the series ends where it begins.
But there is n o t h i n g c o n t r a r y to irreversibility in this.
After all, Octopus h a s not turned into a nautfloid.
2. 1919., p. 117. A c h r o n o l o g i c a l b i b l i o g r a p h y o Dollo's w o r k o n Itr e v e r s i b i l i t y is p r e s e n t e d at t h e close of t h i s article. C i t a t i o n s h e r e i n a r e b y
d a t e a n d r e f e r to t h e s e w o r k s .

190

Dollo on Dollo's L a w
The didactic Dollo d e m o n s t r a t i n g t h a t unrolled a m m o n o i d s
h a v e not reverted to the ancestral straight nautfloid :3
N o n e of t h e m h a s b e c o m e the ancestral Orthoceras a g a i n
...
neither wholly, n o r partially: neither in the initial
c h a m b e r , n o r in the sutures, n o r in the aperture, n o r in the
siphon, n o r in the o r n a m e n t , n o r in the process of uncoiling.
Magnificent e x a m p l e s of irreversibflity [
Yet f r o m the totality of such a p o t h e g m s emerges a very definite
and consistent view of the n a t u r a l world, of evolution and of
paleontology. Dollo's thoughts on irreversibility flow naturally,
almost inevitably, f r o m this c o n c e p t u a l f r a m e w o r k . Divorced
f r o m it, his p h r a s e s are easily misinterpreted. W h e n understood
but depicted without the theoretical u n d e r p i n n i n g t h a t Dollo
provided, irreversibility a p p e a r s as a n isolated curiosity, and one
is left wondering why Dollo invoked it so often a n d with so m u c h
ardor.
Dollo on the natural world: Dollo was educated in the m e c h a n istic tradition t h a t d o m i n a t e d late nineteenth-century science.
His strong reductionist bias t a u g h t h i m t h a t the goal of biology
was to abstract f r o m the organic world a set of governing laws
p a t t e r n e d after the deterministic s y s t e m t h e n prevalent in
physics. This belief not only prescribed a general methodology
(to search for l a w s ) , but also led Dollo to an i m p o r t a n t particular
conclusion: the n e c e s s a r y association of a cause and its effect
m e a n t that a given e n v i r o n m e n t would always elicit the s a m e
type of adaptive morphological response.
W h e n L. Plate criticized the l a w of irreversibflity on the
grounds t h a t "the organic world c a n n o t be ordered according to
absolutely inviolable laws," Dollo replied: 4
I c a n n o t declare m y s e l f to be in a g r e e m e n t with him, because if there are n a t u r a l laws, they m u s t be as c o n s t a n t for
o r g a n i s m s as for the inorganic world. It seems only t h a t they
are m o r e complicated and, as a consequence, m o r e difficult
to discover a n d to define for organisms. To a d m i t the c o n t r a r y
would be to r e t u r n to vitalism.
And after both he a n d Tflly Edinger h a d written independently
to e a c h other of the garden at Stratford-On-Avon where the
flowers m e n t i o n e d in Shakespeare's plays are grown (letter of
J u n e 4, 1927), he c o m m e n t e d :
3. 192% p. 2m.
4. 1922, p. 223.

191

STEPHEN JAY GOULD

Shakespeare Garden: yes the coincidence is curious! But

there is something even more curiousl In a purely mechanical
explanation of the universe, this coincidence was already
scheduled, billions of years ago in the primitive nebula of Kant,
to occur on May 31, 19271 Otherwise, where would it have
come from? For there is no c h a n c e in nature! The famous
F r e n c h m a t h e m a t i c i a n Laplace (the successor of Kant in
natural cosmogeny) said that "we call chance the p h e n o m e n a
of which we are, for the moment, incapable of discovering the
causes." And another explanation of the universe? I do not
know of any.

DoUo on the nature of evolution: Three of Dollo's evolutionary

views are particularly relevant to his notion of irreversibflity:
1. Evolution is discontinuous. De Vries, in fact, credited Dollo
as the first to have stated this postulate on the basis of m o d e r n
evolutionary ideas. 5
2. In the course of evolution, different organs often evolve
independently of each other and at different rates. 6 In a paper
on lungfish evolution Dollo emphasized this "overlapping
(chevauchement) of specializations" and cited as an example: 7
"Hipparion has passed the Equus stage in its dentition; Equus
has passed the Hipparion stage in limb development." Although
he returned infrequently to this principle in his published work,
he considered it of great importance, for he listed it along with
irreversibflity, discontinuity, and limitation among "my laws of
evolution." s
3. Evolution is limited. Dollo used this phrase in two senses.
First, evolution is limited because a highly specialized organism
c a n n o t adjust to a rapidly altered e n v i r o n m e n t and becomes
e x t i n c t - - a n acceptable statement for m o d e r n evolutionists. Of
the specialized turtle, Lytoloma, Dollo wrote: 9
All is sacrificed to an overly precise purpose. They have lost
the necessary plasticity to continue to e v o l v e . . . A new proof
that evolution is limited, since it is the organisms whose
structure responds most exactly to a determined adaptation
which disappear without descendants . . .
We still have
5. 1912, 1). 140.
6. Paleontologists today refer to this p h e n o m e n o n as "mosaic evolution."
7. 1895, p. 88.
8. Letter to Tilly Edinger, November 26, 1927. It is often mentioned i n
subsequent letters.
9. 1903, pp, 25-26.

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Dollo o n Dollo's L a w
t u r t l e s , crocodiles, l a c e r t i l i a n s a n d e v e n r h y n c h o c e p h a l i a n s ,
b u t we h a v e n o m o r e d i n o s a u r s , i c h t h y o s a u r s , p l e s i o s a u r s ,
m o s a s a u r s , or p t e r o s a u r s .
S e c o n d , e v o l u t i o n is l i m i t e d b e c a u s e e a c h l i n e a g e h a s a d e f i n i t e
life cycle b a s e d o n a n i n h e r e n t l y finite c a p a c i t y f o r p h y l e t i c
v a r i a t i o n . 10
Dollo w r o t e i n h i s s h o r t p a p e r o n the l a w s of e v o l u t i o n : 11 "Is
e v o l u t i o n l i m i t e d or i n d e f i n i t e ? Does e a c h o r g a n i s m c a r r y w i t h i n
itself a b o u n d l e s s p o w e r of m e t a m o r p h o s i s or m u s t it n e c e s s a r i l y
b e c o m e e x t i n c t a f t e r h a v i n g r u n t h r o u g h a d e t e r m i n e d cycle?
. . All o r g a n i s m s m u s t n e c e s s a r i l y b e c o m e e x t i n c t a f t e r h a v i n g
r u n t h r o u g h a d e t e r m i n e d cycle w h i c h m a y , h o w e v e r , b e ext r e m e l y long." L a t e r , DoUo s u p p o r t e d t h e v i e w of h i s " ~ m i n e n t
m a i t r e , " A. G i a r d , t h a t ' 2 i v i n g fossils" s u c h as Lingula and the
opossum have stopped evolving "because they have no more
d i s p e n s a b l e p o t e n t i a l for m o d i f i c a t i o n a n d t h e y w o u l d die r a t h e r
t h a n c h a n g e . " 12
Dollo on the nature of paleontology: Dollo b e l i e v e d t h a t "phyl o g e n y ~ l l a l w a y s be t h e s u p r e m e goal of P a l e o n t o l o g y , " la b u t
h e d e p l o r e d t h e s p e c u l a t i v e a p p r o a c h , so c h a r a c t e r i s t i c of l a t e
nineteenth-century paleontology, that built lineages from morp h o l o g i c a l series w i t h o u t r e g a r d to the a d a p t i v e s i g n i f i c a n c e of
o b s e r v e d stages. A t r u l y e v o l u t i o n a r y p a l e o n t o l o g y c o u l d o n l y
r e s u l t f r o m t h e s y n t h e s i s of two a p p r o a c h e s : 14 " P h y l o g e n e t i c
p a l e o n t o l o g y w h i c h s t u d i e s i n h e r i t e d c h a r a c t e r s i n o r d e r to
establish fdiation and ethological paleontology which studies
a d a p t i v e c h a r a c t e r s i n o r d e r to r e c o g n i z e c o n v e r g e n c e s . " F a i l u r e
to r e c o g n i z e c o n v e r g e n c e w a s t h e p r i m e e r r o r of the s p e c u l a t i v e
school. 15 O n l y a n e t h o l o g i c a l a p p r o a c h c o u l d c o r r e c t s u c h errors.
lO. Such a belief is usually associated with various shades of vitalism,
but this was certainly not the case with Dollo. Lamarck was accused of
vitalism for his belief i n the sen~ment inf~rieur, but the existence of such
fluxes and/tows was central to his view of the physical world and carried
no implication of a special status for life. Likewise, Dollo believed that
phyletic life cycle was as natural an idea as individual life cycle. As a
convinced mechanist, Do]lo was a foe of vitalism in any of its forms. Never
could any internal force work to produce or even to preserve an inadaptive
configuration. An "old" species dies because it cannot evolve the required
adaptation to a changing environment.
11. 1893, p. 165.
12. 1905a, p. 131.
13. 1909a, p. 386.
14. Ibid., p. 387.
15. To emphasize this point, Dollo often and proudly cited his demonstration (1895) that the gephyrocercal tail of modern lungfishes is a secondary adaptation to benthic life and not a sign of primitive status. At that

193

S T E P H E N J A Y GOULD

Basic e n v i r o n m e n t s are few; stages in a lineage m a n y . At some

point in the history of m o s t lineages, a derived f o r m will r e t u r n
to the e n v i r o n m e n t of a distant ancestor. Since this e n v i r o n m e n t
requires a definite a n d predictable f u n c t i o n a l adaptation ( a consequence of Dollo's d e t e r m i n i s m ) , convergence to the external
f o r m of the ancestor m u s t occur. I f paleontology c a n succeed in
its " s u p r e m e goal," these convergences m u s t be recognizable; the
derived f o r m m u s t be distinguishable f r o m its distant ancestor of
the s a m e e n v i r o n m e n t .
It is at this point t h a t the concept of irreversibility enters
Dollo's system, for irreversibility provides the g u a r a n t e e that

A ~t

Fig. l a . Dollo's d e t e r m i n i s m ; tail s h a p e is a s u r e g u i d e to e n v i r o n m e n t . F o r m is detel

t e r m i n e d by m o d e of life ( p o i n t e d i n b e n t h i c e n v i r o n m e n t , f a n - s h a p e d i n nectic). F r o m let1
of A p r i l 7, 1928.

convergent f o r m s c a n be distinguished a n d placed in their

proper positions in a n evolutionary sequence. Irreversibility is no
isolated curiosity in Dollo's thought, b u t a n essential step in his
a r g u m e n t that paleontology is a worthwhile endeavor.
t i m e , m a n y p a l e o n t o l o g i s t s w a n t e d to v i e w l i v i n g l u n g f i s h as s u r v i v o r s of
a p r i ~ t i v e s t o c k t h a t h a d g i v e n rise to t e r r e s t r i a l v e r t e b r a t e s .

194

Dollo on Dollo's L a w
The definition of irreversibility is given in this context in
Dollo's work on secondary quadrupedalism in dinosaurs. 1 This
article begins:
I n all studies of adaptation, we m u s t distinguish with care
whether we are dealing with a primary or a secondary adaptation. I n other terms, whether the organism is evolving to

_~r

:ZIp2_

I
l b . Dollo's u s e of irreversibility. T h e r h i p i d o c e r c a l tail of Orl~hagoriscus is s e c o n d a r i l y
n e d f r o m d o r s a l a n d a n a l fins a f t e r irreversible loss of t h e c a u d a l fin. R e d e v e l o p m e n t
nectic m o d e of life p r e s c r i b e s t h e f a n - s h a p e d tail ( d e t e r m i n i s m of figure l a ) , b u t t h e
final r h i p i d o c e r c a l s t r u c t u r e could n o t r e a p p e a r exactly due to irrevocable m o d i f i c a t i o n s
:oduced d u r i n g a n i n t e r c a l a t e d b e n t h i c life. F r o m letter of J u n e 9, 1928.
16.

1905b.

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S T E P H E N J A Y GOULD

satisfy certain determined conditions of existence for the first

time, or, h a v i n g once left these conditions of existence, it is
r e t u r n i n g to t h e m , after h a v i n g adopted, for a m o r e or less
long time, another w a y of life.
He t h e n defines irreversibility as follows:
A n organism never returns exactly to a former state, even
if it finds itself placed in conditions of existence identical
to those in w h i c h it has previously lived. But by virtue of the
indestructibility of the past . . . it always keeps some trace
of the intermediate stages through w h i c h it has passed, iv

T h o u g h Dollo is often done the injustice of being l u m p e d with

the orthogeneticists, it is clear t h a t he n e v e r proposed irreversibility for all evolutionary events. Quite the contrary, for he
only invoked the concept w h e n he needed to show t h a t complete
structural reversal did not a c c o m p a n y reversal in f u n c t i o n or
e n v i r o n m e n t a l preference (see Fig. 1 for a n e x a m p l e illustrated
by Dollo). He applied irreversibility only to complicated morphologies: "Functional or physiological reversal occurs; structural or morphological reversal does not occur." 18
Within this t h e m e t h a t complicated structures are not reevolved, Dollo m i x e d two s t a t e m e n t s of r a t h e r different status.
1. The entire o r g a n i s m n e v e r r e t u r n s to a f o r m e r state: I f
o r g a n i s m s could reacquire an ancestral f o r m exactly, only a n
extremely complete fossil record of gradual change would p e r m i t
the a t t a i n m e n t of Dollo's " s u p r e m e g o a l " - - t h e establishment of
phylogeny. For, to d e m o n s t r a t e such a reversal, one would h a v e
to be able to link the second a p p e a r a n c e t h r o u g h all intermediate
stages to the f o r m m o s t different f r o m beginning and end points
(to a terrestrial m a m m a l for the whale t h a t b e c a m e a fish).
Given the i m p e r f e c t i o n of the geological record 19 and, especially,
Dollo's belief in the discontinuity of evolution, such a linking
could not be imagined. This first s t a t e m e n t is not a n empirical
postulate ( i f such reversals occur, we c a n n o t discern t h e m ) , b u t
an a priori methodological a s s u m p t i o n t h a t preserves the possibility of establishing phylogenies. Speaking of sawfish, Dollo
writes:20
Sawfish h a v e not b e c o m e sharks again. Otherwise, h o w
would we be able to k n o w that they h a d once been depressi17.
18.
19.
9,0.

Ibid., 10. 443.

1903, 10. 32.
1895, p. 88.
1922, 10. 218.

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DoUo on DoUo's L a w
f o r m r a y s i n a b e n t h i c life i n t e r c a l a t e d b e t w e e n the p r i m a r y
a n d s e c o n d a r y n e c t i c life. W e k n o w it b e c a u s e , i n r e a l i t y ,
sawfish are n o t s h a r k s , b u t s q u a l f f o r m rays.
He c a l l e d the n o t i o n of c o m p l e t e r e v e r s i b i l i t y a "postulate w h i c h ,
u n l e s s we p o s s e s s e d a n absolutely c o m p l e t e p a l e o n t o l o g i c a l
series ( w h i c h we are f a r f r o m h a v i n g ) , w o u l d d e s t r o y all possiblity of a r r i v i n g at p h y l o g e n i e s , the s u p r e m e goal of m o r p h o l ogy." 21
2. A c o m p l e x p a r t of a n o r g a n i s m n e v e r r e t u r n s e x a c t l y to a
f o r m e r state. This is a t e s t a b l e s t a t e m e n t a b o u t c o n v e r g e n t
s t r u c t u r e s . 22 Dollo c l a i m e d t h a t he b a s e d his p h y l o g e n i e s on this
s e c o n d s t a t e m e n t of irreversibility, b u t this w a s r a r e l y true. He
b a s e d t h e m on c a r e f u l m o r p h o l o g i c a l c o m p a r i s o n s of all p a r t s ,
not only u p o n the " i n d e s t r u c t i b l e " signs of a n c e s t r y p r e s e r v e d in
the c o n v e r g e n t s t r u c t u r e ; a p h y l o g e n y c a n be e s t a b l i s h e d e v e n
if one p a r t r e v e r t s e x a c t l y to a f o r m e r state. F o r e x a m p l e , he
s t a t e s t h a t m o d e r n p y c n o d o n t fishes are e i t h e r deep-bodied a n d
a d a p t e d to a p l a n k t o n i c ]fie or f u s i f o r m a n d a d a p t e d to n e k t o n i c
life. 23 A s s u m i n g t h a t the u l t i m a t e a n c e s t o r s of p y c n o d o n t s w e r e
f u s i f o r m , w h i c h of the two m o d e r n groups r e p r e s e n t s the p r i m i tive s t a t e ? N o t the f u s i f o r m species, says DoUo, b e c a u s e deepb o d i e d p y c n o d o n t s a p p e a r first in the geologic r e c o r d (chronology)
a n d m a i n t a i n p r i m i t i v e s q u a m a t i o n , teeth, a n d v e r t e b r a l c o l u m n
( m o r p h o l o g y ) . T h e following p h y l o g e n y is t h u s e s t a b l i s h e d
without irreversibility
fusiform pycnodonts

s e c o n d a r y n e k t o n i c life

deep-bodied p y c n o d o n t s

p l a n k t o n i c life

fusiform ancestors

p r i m a r y n e k t o n i c life

N o w we c a n d i s c e r n an e x a m p l e of i r r e v e r s i b i l i t y b e c a u s e the
21. 1907, p. 12,
22. It is a testable statement only if "good faith" is maintained i n
interpreting the qualifying term "complex." This term gives the statement
an "'open texture" that allows an unscrupulous supporter to exclude any
event from its domain by claiming that the event was not sufficiently
complex. See Friedrich Waismann, "Verifiability," in A. Flew (ed.),
Language and Logic (Oxford: Basil Blackwell, 1951). Flew claims that we
lose interest in such a hypothesis because it has suffered "death by a
thousand qualifications" (A. Flew, New Essays in Philosophical Theology
[London SCM Press], pp. 96-97). With a reasonable limit upon the term
"complex", Dollo's statement is testable; if "complex" is used to exclude
any possible counterinstance, the statement becomes unfalsifiable.
23. 1912, pp. 108-109.

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STEPHEN JAY GOULD

neural spines of secondarily fusiform pycnodonts still possess
the reinforcement layers needed by their planktonic ancestors
to support the great body height.
Likewise, this statement c a n n o t be used, as Dollo claimed, 24
to determine the direction of evolution. Chronology and not the
obviously reptilian n a t u r e of fish-like I c h t h y o s a u r u s teaches us
that the lineage of crossopterygian fish--terrestrial r e p t i l e - iehthyosaur did not proceed in reverse order. Were these stages
k n o w n only as an isolated structural sequence of m o d e m forms,
we could not establish direction without some additional postulates on the n a t u r e of evolutionary change. 25
Although we reject the more elaborate claims m a d e by Dollo
for this second statement of irreversibility, its importance is
by no m e a n s diminished. As an affirmation that secondary
convergences c a n be recognized morphologically by their preservation of some trace of an intermediate stage, 26 this statement of
irreversibility is central to Dollo's scientific study of adaptation.
While the status of each of these two statements of irreversibility is d i f f e r e n t - - o n e statement is an unprovable but necessary
assumption, the other a testable proposition--Dollo correctly
offered the same justification for both. The unifying theme is
complexity. Precise reversal does not occur because this would
require that the organism retrace, exactly and in the same order,
an extremely large n u m b e r of steps. Since the c o m p o n e n t s of
an organism c a n evolve independently of one another (his
belief in mosaic evolution), the reacquisition of a complex
structure will d e m a n d nearly as m a n y independent steps as
there are c o m p o n e n t s - - i t c a n n o t be claimed that the parts of a
structure are complex effects of a single cause. 2r The theory of
probability would not permit the second occurrence of such a
large series of independent events. Thus Dollo insisted, t h o u g h
the opposite claim is still being made, 28 t h a t irreversibility is
not a mere empirical generalization f r o m the facts of phylogeny.
Such a charge was insulting to a m a n who prided himself on the
deductive powers he h a d acquired t h r o u g h m a t h e m a t i c a l study.
24. 1922, p. 224, for example.
25. See 1905b, p. 442.
26. Osborn referred to irreversibility as the "law of alternate adaptation."
H. F. Osborn, T h e Origin a n d Evolution of Life (New York: Charles
Scribner's Sons, 1917).
27. Pleiot~opy, and Gregor Mendel for that matter, were unknown when
Dollo formulated his views.
28. Branislav Petronievics, "Sur la loi de l'~volution ir~versible," Sci.
Prog., 13 (1918), 406-419; O. H. Schindewolf, Grundfragen der Paliiontologle (Stuttgart: Schweizbart, 1950), p. 209.

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T h e i r r e v e r s i b i l i t y of e v o l u t i o n is n o t s i m p l y an e m p i r i c a l
l a w r e s t i n g on f a c t s of o b s e r v a t i o n , as m a n y h a v e believed. I t
h a s d e e p e r c a u s e s w h i c h l e a d it, in the l a s t analysis, to a
q u e s t i o n of p r o b a b i l i t i e s as w i t h other n a t u r a l laws. I n effect,
e v o l u t i o n is a s u m m a t i o n of p e r f e c t l y d e t e r m i n e d i n d i v i d u a l
v a r i a t i o n s in a p e r f e c t l y d e t e r m i n e d order. I n order for it to
be reversible, we would h a v e to a d m i t the i n t e r v e n t i o n of
c a u s e s e x a c t l y i n v e r s e to those w h i c h gave rise to the ind i v i d u a l v a r i a t i o n s w h i c h were the source of the first t r a n s f o r m a t i o n a n d also to their fixation in a n e x a c t l y i n v e r s e
o r d e r - - a c i r c u m s t a n c e so c o m p l e x t h a t we c a n n o t i m a g i n e
t h a t it h a s ever occurred. Otherwise, we m i g h t as well m a i n t a i n t h a t by t h r o w i n g into the air the c h a r a c t e r s n e c e s s a r y
f o r p r i n t i n g the Iliad, the p o e m would be c o m p l e t e l y c o m p o s e d
by the s i m p l e f a l l of these little m e t a l l i c blocks, z9
Dollo n e v e r s t a t e d it quite so explicitly, b u t I t h i n k it f a i r to
i n f e r t h a t he b a s e d the a p p l i c a t i o n of irreversibflity u p o n the
position of a p h e n o m e n o n i n a c o m p l e x i t y c o n t i n u u m . A n
i d e n t i c a l o r g a n i s m , he stated, is less likely to be re-evolved t h a n
an i d e n t i c a l o r g a n ; a simple f u n c t i o n c a n be reversed, a c o m p l e x
s t r u c t u r e c a n n o t , no W h e n a p h e n o m e n o n r e a c h e s a sufficient
degree of c o m p l e x i t y , r e q u i r i n g a sufficient n u m b e r of indep e n d e n t steps for its r e a l i z a t i o n , r e p e t i t i o n b e c o m e s "absolutely
u n i m a g i n a b l e - - t h e r e are too m a n y o t h e r possibilities, the probability is nil. T M I n his work on lungfish, he wrote: "Notice t h a t
we are n o t s p e a k i n g h e r e of an i s o l a t e d c h a r a c t e r b u t of a n e n t i r e
series of c h a r a c t e r s . . .
N o w it is, above all, i n its a c t i o n u p o n
h i g h l y m u l t i p l e e l e m e n t s , t h a t we c a n affirm with c e r t a i n t y t h a t
evolution is n o t reversible." 32
As w i t h all c o n t i n u a , there will be p r o b l e m s w i t h b o r d e r l i n e
cases, a n d Dollo cited as e x a m p l e s of irreversibflity some phen o m e n a t h a t m o s t of us w o u l d class i n the simple, r e v e r s i b l e
range. 8a I w o u l d p u t in this c a t e g o r y of m i s p l a c e d b o r d e r l i n e
cases the c l a i m s t h a t bone derived f r o m c a r t i l a g e c a n n o t r e v e r t
to c a r t i l a g e , a n d t h a t s e c o n d a r i l y m a r g i n a l trilobite eyes will
29. 1913, p. 59.
30. 1900, p. 14; 1903, p. 32.
31. 1922, p. 215.
32. 1895, p. 122.
33. It is, of course, well known that simple structures with a simple
genetic base can be reconstituted when lost. See Bjorn Kurt6n, "Return of
a lost structure in the evolution of the felid dentition," Soc. Scient. F e n n .
C o m m e n t . Biol., 26 (1963), 3-11; G. I-Iemmingsmoen, "Zig-zag evolution.'"
N o r s k Geol. Tids., 44 (1964), 341-352.

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not a s s u m e their original position relative to the g l a b e l l a . 34 These

e x a m p l e s are u n f o r t u n a t e , for they lead to the implication t h a t
Dollo h a d in m i n d m o r e t h a n just complexity as a justification
for irreversibility.
Irreversibility, as m o s t of the v e r n a c u l a r words we borrow for
scientific jargon, is not blessed with the u n a m b i g u i t y of a single
m e a n i n g . As we h a v e b e e n using the term, irreversibflity is a
function of the complexity of a series of i n d e p e n d e n t events.
T h e r e is, however, a n o t h e r sense of irreversibflity exemplified
by a notice I once f o u n d tacked to a coffee m a c h i n e : "Irreversibflity: you c a n ' t get y o u r dime back, b y pouring the coffee
b a c k into the m a c h i n e . " The dime is lost not because m a n y indep e n d e n t events are involved in its r e t u m (flipping 100 h e a d s in
a row with an honest coin), but because the m a c h i n e is prog r a m m e d in such a w a y t h a t once the dime is committed, it
c a n ' t be r e c l a i m e d without b r e a k i n g the rules ( j i m m y i n g the
lock, bribing the collector). The irreversibilities proposed b y
various orthogenetic theories fall in this second category. I f it
were true that successive m e m b e r s of phyletic series always increased in size, t h e n a reverse trend would be excluded not because smaller size is a complex genetic c h a n g e but because
evolution would t h e n be p r o g r a m m e d to p r e v e n t this tendency as
long as the rules were followed.
Did Dollo ever speak of a n irreversibflity in this second sense?
I n particular, were certain trends in evolution p r o g r a m m e d in
such a w a y t h a t e m b a r k m e n t u p o n t h e m c o m m i t t e d a lineage
to a definite and irreversible course of development? Here DoUo
is ambiguous. H e often wrote t h a t certain specialized f o r m s
c a n n o t b e c o m e generalized, b u t these short, d e c l a m a t o r y statem e n t s leave us w o n d e r i n g w h e t h e r this irreversibflity arises
because specialized f o r m s inevitably lose structures too intricate
to be regained ( t h e u s u a l a r g u m e n t b a s e d on c o m p l e x i t y )
or because s o m e t h i n g i n h e r e n t in the evolutionary process
dictates t h a t lineages m u s t proceed f r o m generalized to specialized f o r m s ( t h e orthogenetic a r g u m e n t b a s e d on p r o g r a m m e d
sequence). Refuting, for example, the link b e t w e e n ptyctodont
arthrodires and holocephalians, DoUo wrote: "If ptyctodonts are
holocephalians t h e n the m o s t ancient holocephalians are the
m o s t specialized. How, f r o m this state, could they give rise to
their successors. Impossible. And h e r e is a n e w application of
the irreversibflity of evolution." an
34. 1909b, 10. 430; 1909a, p. 410.
35. 1907, p. 7.

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Dollo on DoUo's L a w
T h e solution to this p r o b l e m is probably contained in DoUo's
views on the limitation of evolution, for h e r e we find the s a m e
a m b i g u i t y (see p a g e 193). Limitation b a s e d on the inability of a
very specialized f o r m to a d a p t to n e w conditions leads to a n
irreversibility d e t e r m i n e d by the loss or irrevocable modification
of c o m p l e x structures. Limitation b a s e d on the c u r t a i l m e n t of
"plastic potential" in racial old age leads to a n irreversibility
d e t e r m i n e d by p r o g r a m m e d sequence. Since he speaks of limitation in both senses, we m u s t conclude t h a t an orthogenetic interpretation of a f e w of Dollo's s t a t e m e n t s on irreversibility is
not inconsistent with his view of the evolutionary process. T h u s ,
Dollo m u s t share a portion of the b l a m e for the m o r a s s of
confusion t h a t his law generated in our literature. And yet, the
text t h a t defines irreversibflity as an a d j u n c t of orthogenetic
theories does Dollo a great injustice, for only a very f e w of his
s t a t e m e n t s could be fairly interpreted in this light and none need
be.
I n conclusion, three senses of irreversibility m a y be discerned
in Dollo's works:
1. A n a p r i o r i a s s u m p t i o n t h a t a whole o r g a n i s m n e v e r reverts
completely to a prior phylogenetic stage.

2. A testable hypothesis t h a t a c o m p l e x p a r t of a n ancestor

n e v e r r e a p p e a r s in exactly the s a m e f o r m in a descendant.
T h e s e two propositions are the h e a r t of irreversibflity; all of
Dollo's s t a t e m e n t s can, a n d probably should, be tied to one
of t h e m . T h e y h a v e the s a m e justification ( a probability argum e n t b a s e d on c o m p l e x i t y ) and play the s a m e role ( a m e t h o d
for the recognition of c o n v e r g e n c e s ) .
3. Certain evolutionary trends are necessarily unidirectional.
This i n t e r p r e t a t i o n c a n be attached only to a very f e w of Dollo's
s t a t e m e n t s . I f in Dollo's m i n d at all, it played a n e x t r e m e l y
m i n o r role in his t h o u g h t on irreversibility.
II. T H E DEBATE OVER IRREVERSIBILITY
Tis with our judgments as our watches, none
Go just alike, yet each believes his own.
A. Pope, E s s a y o n C r i t i c i s m
The literature on irreversibility is a quagmire. I h a d h a r b o r e d
the n a i v e hope of finding some chronological t r e n d in opinion;
either the h a p p y discovery t h a t a c c u m u l a t i n g w i s d o m drives out

201

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misinterpretation or the cynical result that the f u r t h e r we get

f r o m DoUo's writing, the less likely we are to r e a d it and the
m o r e likely, therefore, to misrepresent. I find, instead, t h a t
the tendency to m i s r e a d (or not r e a d ) is timeless: the s a m e
errors h a v e been cropping up with the s a m e f r e q u e n c y ever since
the debate started. I h a v e tried, therefore, to delimit w h a t I
consider the six m a j o r interpretations of Dollo's opinion. For
several of these we h a v e Dollo's own r e f u t a t i o n of views attributed to h i m , recalling M a r x ' s f a m o u s abjuration of
"marxism."
A p r i m a r y division arises f r o m the two principal m e a n i n g s
of irreversibility: Dollo's based on the improbability of reversion
in complex structures, a n d the orthogenetic b a s e d on the necessity of following a p r o g r a m m e d sequence.
A m o n g those w h o incorrectly i m p a r t e d an orthogenetic interpretation to Dollo's views, we see two tendencies.
1. Orthogeneticists w h o cited their m i s u n d e r s t a n d i n g to support a theory of ineluctable trends. Dep6ret defined DoUo's l a w
as "the f a c t t h a t a lineage, h a v i n g once e m b a r k e d on the p a t h
of a determined specialization, c a n in no case turn b a c k upon the
r o a d already travelled." We find him, a f e w pages later, using
this definition to support a claim t h a t "each phyletic b r a n c h h a s
a geologic career in which we c a n distinguish a p h a s e of youth,
a p h a s e of m a t u r i t y , a n d finally a p h a s e of senility or degeneration p r e p a r a t o r y to extinction of the type." 36 Likewise, Cu6not,
who believed that "orthogenesis is p r e p a r a t o r y to the future of
lineages just as ontogenesis is p r e p a r a t o r y to the f u t u r e of
a n individual," interpreted Dollo's l a w to m e a n t h a t certain
trends were necessarily unidirectional. 37
Beurlen, though not completely in the orthogenetic c a m p ,
shared the antimechanism of Dep6ret and Cu6not. He devoted
a n entire c h a p t e r to a supposed d e m o n s t r a t i o n that the validity
of Dollo's law demolishes a purely m e c h a n i s t i c interpretation of
the n a t u r a l world: "The knowledge t h a t phylogenetic history is
irreversible m e a n s that a mechanistic-causal interpretation, however it is constructed, either p r i m a r i l y L a m a r c k i a n or p r i m a r i l y
Darwinian, c a n n o t alone suffice for a n u n d e r s t a n d i n g of phylogeny." as Dollo, a convinced m e c h a n i s t , would h a v e t a k e n great
36. Chaxles Dep6ret, Les Transformations du m o n d e animal (Paris:
Ernest F l a m m a r i o n , 1919), pp. 243, 246.
37. L. Cu6not, L'Evolution biologique, (Paris: Masson, 1951), pp. 49-51,
537.
38. K. Beurlen, Die stammesgeschichtlichen Grundlagen der A b s t a m m ungslehre (Jena: Gustav Fischer, 1937), p. 42.

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Dollo o n Dollo's L a w
offense a t s u c h a u s e of h i s work, for h e h a d s t a t e d explicitly 39
t h a t i r r e v e r s i b f l i t y offered n o c h a l l e n g e to a m e c h a n i s t i c w o r l d
view.
2. D a r w i n i a n s w h o a t t a c k a f a l l a c i o u s o r t h o g e n e t i c i n t e r p r e t a t i o n , o f t e n w i t h t h e v e r y a r g u m e n t s DoUo u s e d to s u p p o r t h i s
v e r s i o n of irreversibility. E a s t o n , 40 for e x a m p l e , wrote: " A b o u t
the s a m e t i m e t h a t the d o c t r i n e of o r t h o g e n e s i s w a s p r o p o s e d ,
a c o l l a t e r a l h y p o t h e s i s w a s s u g g e s t e d i n E u r o p e b y Dollo. It w a s
DoUo's b e l i e f t h a t e v o l u t i o n w a s a l w a y s p r o g r e s s i v e - - t h a t is, t h a t
creatures, once they started down a certain path, never retreated
n o r c o u l d t h e y r e s u m e a f o r m e r c o n d i t i o n . " To u n d e r m i n e this
s u p p o s e d d e f i n i t i o n of Dollo's l a w , E a s t o n cited the r e t u r n to
w a t e r of i c h t h y o s a u r s a n d w h a l e s , a n e x a m p l e u s e d o f t e n b y
Dollo to s u b s t a n t i a t e h i s n o t i o n of irreversibflity.41 S i m i l a r
o r t h o g e n e t i e m i s d e f i n i t i o n s are p r o p o s e d as DoUo's o w n a n d
r e j e c t e d b y B e e r b o w e r a n d E h r e n b e r g . 42
F o u r m a j o r a r g u m e n t s are p r e s e n t e d b y s c i e n t i s t s w h o reco g n i z e d at l e a s t a p a r t of Dollo's c l a i m - - t h a t r e v e r s i b i l i t y of
s t r u c t u r e is the o b j e c t of d e b a t e .
3. Dollo c l a i m e d only t h a t c o m p l e x s t r u c t u r e s could n o t be
reacquired. 4a T h i s is n o t a l w a y s valid b e c a u s e ancestral structures are p r e s e r v e d i n early o n t o g e n e t i c stages (due to acceleration) a n d m a y again b e c o m e a d u l t i n p a e d o m o r p h i c f o r m s .
N o p c s a a t t r i b u t e d the r e a p p e a r a n c e of t h e p o s t o r b i t a l b a r i n
m a m m a l s to s u c h a process a n d s t a t e d : 44 " T h e u n e x p l a i n a b l e
b u t i m p o r t a n t f a c t , t h a t t h e l i f e - h i s t o r y of e a c h i n d i v i d u a l is
a l w a y s a d i s t o r t e d r e c a p i t u l a t i o n of t h e h i s t o r y of its w h o l e
p h y l u m , gives t h e clue b y w h i c h w e c a n u n d e r s t a n d w h y a
39. 1905a, p. 130.
40. W. H. Easton, Invertebrate Paleontology (New York: Harper, 1960),
19. 42.
41. 1912, 19. 106, and 1922, 19. 216.
42. ft. R. Beerbower, Search for the Past (Englewood Cliffs, N. J.: PrenticeHall, 1960), 19. 156; Kurt Ehrenberg, Paldozoologie (Vienna: Sl~ringer, 1960),
19.22.
43. The correct interpretation so far.
44. Francis Nopesa, "Reversible and irreversible evolution; a study
based on reptiles," Proc. ZooL Soe. London, (1923), 1058. The same 1point
has been made, with different examples in: G. J. Fej6rvary, "'Quelques
observations su~ la loi de Dollo et F619istr619hog~n~se en consideration
s19~eiale de la loi biog6n6tique de Haeckel," Bull. Soc. Vaud. Sci. Nat.,
53 (1920), 343-372; P. P. Sushkin, "'Notes on the 19re-Jurassic telxa19ods
from the U.S.S.R.," Tray. Inst. PaldozooL Acad. Sci. U.S.S.R. Leningrad, 5,
(1936), 43-91; and, more recently by: ~I. A. Shishkin, "Morphogenetic
factors and the irreversibility of evolution," Paleont. J., 3 (1968), 293-299.

203

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JAY

GOULD

l i m i t e d r e v e r s a l of e v o l u t i o n c a n occur." Dollo h a d a n t i c i p a t e d
this o b j e c t i o n a n d p r o v i d e d a c o u n t e r a r g u m e n t b a s e d o n the
f a c t t h a t " o n t o g e n y is n o t a c o m p l e t e a n d e x a c t r e c a p i t u l a t i o n of
p h y l o g e n y . " 45 N o t o n l y is r e c a p i t u l a t i o n a l w a y s i m p e r f e c t ( p r e c l u d i n g the e x a c t r e a c q u i s i t i o n of a s t r u c t u r e ) , it is also f a r f r o m
u n i v e r s a l i n o c c u r r e n c e . Of the o n t o g e n y of the t u r t l e Dermochelys coriacea, DoUo wrote: 46
There m u s t be a p e r t u r b a t i o n in r e c a p i t u l a t i o n - - s i n c e if
we r e f u s e to a d m i t p e r t u r b a t i o n , we a r r i v e at a p h y l o g e n y i n
o p p o s i t i o n to c h r o n o l o g y , p a l e o n t o l o g y , ethology, etc. Let u s
avoid a b u s i n g a l a w w h o s e f a l l a c i e s h a v e a l r e a d y b e e n e n u n c i ated b y t h e i l l u s t r i o u s F r i t z Miiller; it h a s v a l u e o n l y if we
a p p l y it w i t h d i s c r i m i n a t i o n .
Moreover, h e w r o t e to T i l l y E d i n g e r : "It is clear, a priori, t h a t
o n t o g e n y c a n n o t be a c o m p l e t e a n d e x a c t r e c a p i t u l a t i o n of phylogeny. J u s t t h i n k h o w l o n g it w o u l d t h e n take for e v e n a s i m p l e
i n d i v i d u a l to develop !" ~
4. Dollo c l a i m e d t h a t n o s t r u c t u r a l reversal o f a n y k i n d w a s
possible i n evolution. T w o m i s i n t e r p r e t a t i o n s are i n v o l v e d h e r e
s i n c e Dollo b e l i e v e d t h a t n o c o m p l e t e r e v e r s i o n of c o m p l e x
s t r u c t u r e s c o u l d occur. M a n y a u t h o r s h a v e cited i n c o m p l e t e
r e v e r s i o n s of c o m p l e x s t r u c t u r e s as e x c e p t i o n s to Dollo's law.
T h e s e c o n d a r y i s o d o n t t e e t h of c e t a c e a n s , r e c a l l i n g t h e d e n t i t i o n
of u l t i m a t e r e p t i l i a n a n c e s t o r s , is a p e r e n n i a l f a v o r i t e : S t r o m e r
a n d Scott i n v o k e d it a n d R e n s c h r e g a r d e d it as a n " u n q u e s t i o n able e x c e p t i o n to Dollo's views. 48 B u t Dollo h a d a n s w e r e d t h e
v e r y s a m e o b j e c t i o n i n 1907:
N o w the e v o l u t i o n of the d e n t i t i o n of w h a l e s is o n e of t h e m o s t
b e a u t i f u l e x a m p l e s of t h e i r r e v e r s i b f l i t y of e v o l u t i o n , s i n c e
t h e s e c o n d a r y i s o d o n t d e n t i t i o n is n o t a r e t u r n to t h e p r i m i t i v e
1 9 2 2 , p . 216.
46. 1901, p. 20.
47. Letter of November 17, 1928. Dollo had enormous respect for
Haeckel despite his doubts about recapitulation. He wrote to Tilly Edinger
(letter of June 30, 1928): "I do not want you to compare me with Haeckel!
. . . We exchanged publications, but I never had a personal relationship
with him. Nevertheless, a curious thing, he was interested in me and in
my work. Abel went t o s e e him several times and, each time, he asked:
'How is Dollo? What is Dollo doing?" "
48. E. Stromer, Lehrbueh der Paldozoologie, vol. 2, Wirbeltiere (Leipzig:
B. G. Teubner, 1912), p. 288; W. B. Scott, A History of Land Mammals in
the Western Hemisphere, (New York: Macmillan, 1929), p. 656; Bernhard
Rensch, Evolution Above the Species Level (New York: Columbia University Press, 1960), p. 124.
45.

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Dollo on Dollo's Law

isodont dentition but is an isodont dentition of completely
different morphology. It is infuriating that M. von Arthaber
chose to contradict before he h a d understood. ~9
Atavisms have often been cited as exceptions to Dollo's law. 5
Abel 51 devoted a lengthy chapter to dismissing this claim by
s h o ~ n g that atavistic structures never produce complete reversion. Dollo dismissed it more gently: "Have you noticed m y
simian atavism on this p h o t o g r a p h (Darwin's point on the ear).
But it is only partial and that is a beautiful example of irreversibflity I" 52
Other authors have tried to refute Dollo by pointing out that
complete reversion occurs in simple structures. Back m u t a t i o n
is often cited. 53 Stromer invoked the size reversion of d w a r f
hippos and elephants; Boulenger noted that lost vertebrae have
been regained in several fish lineages. 5~ Most ironic are the
statements of authors who unwittingly use DoUo's own formulation to refute their misconception of Dollo's law:
Deeply entrenched is the conception that phylogenetic development is irreversible (Dollo's L a w ) . But this is so only in
certain cases, especially w h e n we are dealing with the entire
o r g a n i s m or complex organs. A reversion to phylogenetically
earlier states c a n occur in simple structures . . . The unrestricted use of Dollo's l a w . . ,
as with the similarly extreme
use of the biogenetic law, has led the study of phylogeny into
too m a n y errors of interpretation.~5
5. DoUo did not claim that all evolution was irreversible, but
only that lost structures could not be regained in the same form.
Pressured by self-styled opponents of DoUo's law who b r o u g h t
up the irrelevant examples of reversion in simple structures cited
above, m a n y of Dollo's supporters restricted the law to a "sure
thing," but emasculated it in so doing. Diener, and especially
49. 1907, p. 7.
50. Rensch, i n Evolution, cites several s t a n d a r d examples.

51. Othenio Abel, Paliiobiologie und Stammesgeschichte (Jena: Gustav

Fischer, 1929).
52. Letter to Tilly Edinger, November 21, 1926.
53. Rensch, Evolution, p. 124; Walter Z i m m e r m a n n , "lVIethoden der
Phylogenetik," i n G. Heberer (ed.) Die Evolution tier Organismen (Stuttgart:
Gustav Fischer, 1954), pp. 25-102.
54. Lehrbuch, p. 285; G. A. Boulenger, "L'~volution est-elle r6versible?
considerations au sujet de certains poissons," Compt. Rend. Acad. Sci., 168,
(1919), 41--44.
55. Adolf Remane, "Die Geschlehte der Tiere," i n G. Heberer (ed.) Die
Evolution der Organismen (Stuttgart: Gustav Fischer, 1954), p. 419.

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STEPHEN JAY GOULD

Abel, are identified with this restriction, and their opinion was
c o p i e d b y m a n y l a t e r a u t h o r s . 56 S i m p s o n r e f e r r e d to t h i s p r a c t i c e
a s a " k i n d l y t r a d i t i o n , " b u t Dollo h i m s e l f h a d d i s a v o w e d it: " T h e
i r r e v e r s i b i l i t y o f e v o l u t i o n d o e s n o t o n l y a p p l y to l o s t or r e d u c e d
o r g a n s , b u t also to f u n c t i o n a l o r g a n s . " 57 T h e p o i n t is t h a t i n so
r e s t r i c t i n g Dollo's l a w , i t s e s s e n t i a l r e l a t i o n s h i p to c o m p l e x i t y
o f s t r u c t u r e is o b s c u r e d a n d its v a l u e a s a g e n e r a l s t a t e m e n t
destroyed.
6. D o l l o c l a i m e d o n l y t h a t c o m p l e x s t r u c t u r e s c o u l d n o t b e
r e e v o l v e d ; t h i s is a v a l i d s t a t e m e n t . S o m e s h o r t t e x t b o o k s t a t e -

m e n t s h a v e p r e s e n t e d Dollo's o w n p o s i t i o n i n a f a v o r a b l e l i g h t ; ~s
others, unfortunately a majority, expound the orthogenetic interp r e t a t i o n . O t h e r w o r k s , w h i c h c o n s i d e r t h e q u e s t i o n in g r e a t e r
detail, support both Dollo's formulation and his justification
b a s e d o n p r o b a b i l i t i e s . H e r e w e f i n d the w r i t i n g s o f S c h i n d e w o l f ,
Blum, Muller, and, especially, Simpson. Muller, arguing from
g e n e t i c s , s p o k e of " t h e s h e e r s t a t i s t i c a l i m p r o b a b i l i t y , a m o u n t i n g
to a n i m p o s s i b i l i t y , o f e v o l u t i o n e v e r a r r i v i n g a t t h e s a m e c o m plex genic end-result twice." Blum, from a thermodynamic
standpoint, stated that "the chances of retracing the steps of
e v o l u t i o n o v e r a n y d i s t a n c e b e c o m e s v a n i s h i n g l y s m a l l as t h e
c o m p l e x i t y o f o r g a n i s m s a n d t h e i r e n v i r o n m e n t i n c r e a s e . 59
T h u s f e w c r i t i c s w h o a t t a c h e d Dollo's n a m e to t h e i r d i s c u s s i o n
of i r r e v e r s i b i l i t y c o r r e c t l y p r e s e n t e d w h a t D o l l o h i m s e l f h a d
said. I c a n n o t a v o i d t h e f e e l i n g t h a t t h e s e m i s s t a t e m e n t s a r i s e
f r o m u n f a m i l i a r i t y w i t h t h e i n t e l l e c t u a l c o n t e x t of Dollo's i t r e v e r s i b i l i t y . F o r Dollo, a l a w o f i r r e v e r s i b f l i t y f u n c t i o n s a s a
guarantee that eonvergences can be recognized by preservation
of some ancestral structure(incomplete reversion). Convergence
56. Karl Diener, Palfiontologie u n d A b s t a m m u n g s l e h r e (Leipzig: Samml.
Goschen, 1910); Othenio Abel, Grundziige der Palaeobiologie der
Wirbeltiere (Stuttgart: Schweizbart, 1912), and Paltiobiologie; for example, in: W. K. Gregory, "On the meaning and limits of irreversibility
in evolution," Am. Nat., 70, (1936), 517-528; G. S. Carter, A n i m a l Evolution: A Study of Recent Views and Its Causes (London: Sidgwick and
Jackson, 1951).
57. G. G. Simpson, T h e Major Features of Evolution (New York:
Columbia University Press, 1953), p. 310; Dollo, 1909a, p. 397.
58. A. M. Davies, A n Introduction to Paleontology (London: Thomas
Murby, 1947); R. C. Moore, C. G. Lalicker, and A. G. :Fischer, Invertebrate
Fossils (New York: McGraw-Hill, 1952).
59. O. Schindewolf, Pal~iontologie; H. F. Blum, Time's Arrow and
Evolution (Harper Torchbacks, 1962), p. 200; H. J. Muller, "'Reversibility
in evolution considered from the standpoint of genetics," Biol. Rev., 14
(1939), 27; G. G. Simpson, Evolution, and This V i e w of Life (New York:
Harcourt, Brace and World, 1964).

9.06

Dollo on Dollo's L a w
is the m a j o r i m p e d i m e n t to phylogenetic interpretations; phylogeny is the goal of paleontology. I n this context, Dollo could
scarcely have been excluding a n y t h i n g but complete reversal.
W h e n this context is not known, criticism m a y be based solely
on the various v e r n a c u l a r senses of irreversibflity; n u m e r o u s
interpretations ( n o n e of them Dollo's) will then arise. The
seemingly endless and often acrimonious debates that have
raged about the concept of irreversibility have almost always
been based not on substantive disagreement concerning the
course of evolution, but rather on the sheer semantic misunderstanding generated by using one t e r m - - " D o l l o ' s law" for a
variety of contradictory concepts.
III. IRREVERSIBILITY AND T H E STATUS OF
EVOLUTIONARY LAWS
The nonrecurrence of experienced events must be one of the oldest
notions of the human mind, for in any real experience our sensation of
time is unidirectional and the ix'reversibility of history and of evolution
seem to be corollaries of this.
H. Blum, Time's Arrow and Evolution, p. 179
Our textbooks of evolution usually describe Dollo's notion of
irreversibility in c o n j u n c t i o n with other supposed "evolutionary
l a w s " - - u s u a l l y with "Cope's law" that body size tends to increase
in phyletic sequences and "Williston's law" that large n u m b e r s
of sJrnflar elements tend to be reduced to fewer differently specialized units. The attempt to order phylogenetic events into
regularities sufficiently pervasive to be termed laws was a popular strategy earlier in this century. It was centered on the
reductionist view that biology should be patterned on the formal
structure of the physical sciences. W. K. Gregory, a m a j o r prop o n e n t of this strategy, held that "we ought logically to begin
with the forces inside the h y d r o g e n atom and work outward and
u p w a r d t h r o u g h organic chemistry to m a n . " 6o The N e w t o n i a n
synthesis h a d produced a set of descriptive generalizations that
ordered complex results into a simpler lawful structure. Thus,
it was argued, the m a t u r a t i o n of evolutionary biology to true
science depended on the discovery of lawful structure a m o n g
phylogenefic events.
The laws of Cope and Williston are attempts at descriptive
60. w. K. Gregory, "Basic patents in nature," Science, 78 (1933), 561566.

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S T E P H E N J A Y GOULD

g e n e r a l i z a t i o n . Dollo's law, o n the o t h e r h a n d , h a s a v e r y

d i f f e r e n t status. P r e v i o u s l y , I d i v i d e d DoUo's l a w i n t o two statem e n t s : ( 1 ) T h e e n t i r e o r g a n i s m n e v e r r e t u r n s to a f o r m e r state.
( 2 ) A c o m p l e x p a r t of a n o r g a n i s m n e v e r r e t u r n s e x a c t l y to a
f o r m e r state. T h e first, as d i s c u s s e d o n p a g e 196, is n o s t a t e m e n t
a b o u t d a t a a t all, b u t a n e c e s s a r y a p r i o r i a s s u m p t i o n t h a t
p h y l o g e n i e s c a n be e s t a b l i s h e d , g i v e n the n a t u r e of e v o l u t i o n a n d
t h e fossil record. T h e s e c o n d s e e m s to h a v e t h e f o r m of a n
e m p i r i c a l l y g r o u n d e d s t a t e m e n t , b u t the a m b i g u i t y i n t r o d u c e d
b y v a r y i n g i n t e r p r e t a t i o n s of " c o m p l e x " h a s r o b b e d it of a n y
p r e c i s e m e a n i n g a n d g e n e r a l u s e f u l n e s s (see n o t e 2 2 ) . A l m o s t
a n y r e v e r s i o n c a n be e x c l u d e d f r o m its d o m a i n b y c l a i m i n g
e i t h e r t h a t t h e s t r u c t u r e s i n v o l v e d w e r e n o t sufficiently c o m p l e x
or t h a t the g e n e t i c b a s i s f o r a n e l a b o r a t e m o r p h o l o g i c a l c h a n g e
w a s simple.
If this s e c o n d p a r t of "Dollo's law" is n o t a u s e f u l s t a t e m e n t
i n a r g u m e n t s a b o u t specific p h y l o g e n i e s , w h a t is its s t a t u s a n d
h a s it a n y i m p o r t a n c e ? I f we a s s u m e t h a t c o m p l e x e v o l u t i o n a r y
e v e n t s g e n e r a l l y h a v e c o m p l e x c a u s e s f i 1 t h e n DoUo's l a w s i m p l y
affirms t h a t the r e s u l t s of e v o l u t i o n c o n f o r m to o u r g e n e r a l
n o t i o n of h i s t o r y as a s e q u e n c e of u n i q u e p h e n o m e n a . 62 A n d
Dollo's finest i n s i g h t w a s t h a t h e p r o v i d e d as his j u s t i f i c a t i o n
of e v o l u t i o n a r y i r r e v e r s i b f l i t y t h e v e r y s a m e a r g u m e n t a d v a n c e d
t o d a y for t h e u n i q u e n e s s of h i s t o r i c a l e v e n t s - - t h e s t a t i s t i c a l
i m p r o b a b i l i t y t h a t the i n c a l c u l a b l e n u m b e r of i n d e p e n d e n t conf i g u r a t i o n s a n t e c e d e n t to a n d c o m p r i s i n g a n y h i s t o r i c a l e v e n t
s h o u l d ever o c c u r twice. T h u s , Dollo's l a w is n o t a n a d j u n c t of
e v o l u t i o n a r y theory. I t is a s t a t e m e n t , f r a m e d i n t e r m s of a n i m a l s
a n d t h e i r e v o l u t i o n , of the n a t u r e of h i s t o r y ; or, p u t a n o t h e r way,
it is a n a f f i r m a t i o n of t h e h i s t o r i c a l n a t u r e of e v o l u t i o n a r y
e v e n t s . As is so o f t e n the case, we are i n d e b t e d to G. G. S i m p s o n
for this p e r c e p t i v e i n t e r p r e t a t i o n : " T h a t e v o l u t i o n is i r r e v e r s i b l e
is a special case of the f a c t t h a t h i s t o r y does n o t r e p e a t itself.
61. The argument leveled against Dollo's law by Nopcsa and Fej~rvary
(note 44) was based on a denial of this premise. They claimed that complex
reversions could be produced by the reasonably simple mechanism of
acceleration followed by paedomorphosis. In this sense, their argument is
potentially the strongest of any leveled against Dollo's law, but it failed
because nature doesn't work in the way they imagined. Yet even if it did,
we could stiU preserve Dollo's law by claiming that such reversions were
not really complex and that only reversions with complex causes should
be covered by the law. This is what I mean in stating that almost any
empirical challenge against the law can be ref~uted.
62. G. G. Simpson, This V i e w of Life, and quote of H. Blum introducing
this section.

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Dollo on Dollo's L a w
The fossil record and the evolutionary sequences that it illustrates are historical in nature, and history is inherently irreversible." 63
This interpretation of Dollo's law is involved in our j u d g m e n t
u p o n the descriptive generalizations of Cope and Wflliston and
u p o n the entire enterprise of l a w m a k i n g for phylogenetie results.
Simpson h a s distinguished i m m a n e n t f r o m configurational
properties of the universe (the f o r m e r as "the u n c h a n g i n g properties of m a t t e r and energy and the likewise u n c h a n g i n g processes and principles arising therefrom"; the latter as "the
actual state of the universe or of any part of it at a given time. 64
Laws are f r a m e d for i m m a n e n t properties: we are not interested
in the melting behavior of a particular ice cube but in the
properties of water in general. Physics rarely deals with the
eonfigurational; if its f o r m a l structure is lawlike, this is because
it has excluded the configurational f r o m its domain. The error
m a d e by reductionists who attempted to formulate laws for the
results of evolution was that they assumed a similar focus for
biology and physics. But biology often deals with the configurational and the search for so-called historical laws a m o n g such
properties is not a fruitful endeavor. While I agree with W a t s o n
and Siever that there is no f o r m a l distinction between historical
and non-historical science, 6~ there is a differenee of emphasis.
There are nomothetic undertones to the results of e v o l u t i o n - - t h e
principle of n a t u r a l selection is a m o n g t h e m - - a n d it is here that
our laws m u s t be formulated. They m u s t be based on i m m a n e n t
processes that produce events, not on the events themselves. The
'qaws" of Cope and Wflliston, based as they are on configurational properties, are not laws in the ordinary sense but descriptive generalizations of low-order probability that describe some
c o m m o n regularities without explaining anything. "Dollo's law"
is not a m o n g these. Quite the contrary: since irreversibility is an
a c k n o w l e d g m e n t of the historical n a t u r e of evolutionary events
and since that very n a t u r e precludes the f o r m u l a t i o n of laws for
these events, "Dollo's law" is a particularized statement of our
reason for rejecting the approach to evolutionary biology that
led to the laws of Cope and Williston. Dollo is done an injustice
63. This View of Life, p. 186.
64. Ibid., p. 122. Nagel makes a similar distinction between nomothetic
and ideographic properties (Ernest Nagel, "The logic of historical analysis,"
in H. Feigl and M. Brodbeck [eds.], Readings in the Philosophy of Science
[New York: Appleton-Century-Crofts, 1953], pp. 688-700).
65. R. A. Watson, "Is geology different? A critical discussion of The
Fabric of Geology," Phil. Sci., 33, (1966), 172-185; Raymond Siever,
"Science: observational, experimental, historical," Am. Scientist, 56,
(1968), 70--77.

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S T E P H E N J A Y GOULD

w h e n his views are relegated to the w a s t e b a s k e t of archaic

strategies, for irreversibflity is no outmoded historical law;
rather, the search for historical laws is outmoded because we
h a v e n o w recognized the significance of irreversibflity in Dollo's
sense.
Yet surely there is something ironic here, for DoUo h i m s e l f was
a reductionist and quite c o m m i t t e d to the search for historical
laws. In fact, w h e n he presents his justification for irreversibility,
his words b e t r a y the d e t e r m i n i s m t h a t characterized m o s t reductionist thinking prior to the advent of q u a n t u m physics: To
repeat "perfectly d e t e r m i n e d individual variations in a perfectly
determined o r d e r . . ,
we would h a v e to a d m i t the intervention
of c a u s e s . . ,
in an exactly inverse order." 66 I doubt t h a t DoUo
ever perceived the anti-deterministic implications of the larger
generality of which his irreversibflity f o r m s a special case. And
yet, if the operas of W a g n e r , which Dollo loved p e r h a p s m o r e
t h a n his fossils, are any guide, it is u p o n the content of the piece
itself and not the intent of its author t h a t we m u s t judge a m a n ' s
w o r k . 67
66. 1913, p. 59.
67. I t h a n k E r n s t M a y r , D i r e c t o r of t h e M u s e u m of C o m p a r a t i v e Zoology,
E v e r e t t M e n d e l s o h n , H i s t o r y of S c i e n c e D e p a r t m e n t , H a r v a r d U n i v e r s i t y ,
a n d Carl P u t z , P h i l o s o p h y D e p a r t m e n t , De P a u w U n i v e r s i t y f o r t h e i r c a r e f u l
a n d e x t e n s i v e c r i t i c i s m of t h e m a n u s c r i p t . A. S. R o m e r k i n d l y l e n t m e t h e
c o r r e s p o n d e n c e of Dollo a n d T i l l y E d i n g e r a n d r e g a l e d m e w i t h t h e bits
of h u m a n i n t e r e s t t h a t s u b s t i t u t e a l i v i n g m a n f o r t h e a b s t r a c t i d e a s
g l e a n e d f r o m h i s p u b l i s h e d work.

APPENDIX
T H E L A W S OF E V O L U T I O N , b y L o u i s Dollo
( T r a n s l a t e d f r o m Bull. Soc. Belge Geol. Paleontol. Hydrol., 7 [1893], 1 6 4 166).
I. A c c o r d i n g to t h e b r i l l i a n t c o n c e p t i o n of t h e i m m o r t a l C h a r l e s D a r w i n
(1809-1889,):
E v o l u t i o n - - t h e t r a n s f o r m a t i o n of o r g a n i s m s - - r e s u l t s f r o m t h e
f i x a t i o n of i n d i v i d u a l u s e f u l v a r i a t i o n s b y t h e i n f l u e n c e of n a t u r a l
selection provoked by the struggle for existence.
All species, a n i m a l or v e g e t a b l e , w h i c h e x i s t or h a v e e x i s t e d s i n c e
t h e a p p e a r a n c e of life o n t h e globe, owe t h e i r o r i g i n to t h i s f u n d a mental law.
II. But:
1. W h a t is t h e c a u s e of i n d i v i d u a l v a r i a t i o n s ?

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Dollo on Dollo's Law

2. W h a t is t h e i r a m p l i t u d e ? I s it s m a l l ? I s it l a r g e ? I s evolution
e x t r e m e l y slow, or does it o c c u r by r a t h e r r a p i d j u m p s ?
3. F r o m a n o t h e r v i e w p o i n t , is e v o l u t i o n reversible? C a n a n o r g a n i s m
r e t u r n ( e i t h e r totally or partially) to a f o r m e r c o n d i t i o n already
realized i n the series of its a n c e s t o r s ? - - e i t h e r by r e t u r n i n g i n
a single j u m p or b y p a s s i n g , i n r e v e r s e order, t h r o u g h t h e v a r i o u s
stages w h i c h led to its origin.
4. Finally, is e v o l u t i o n l i m i t e d or indefinite? Does every o r g a n i s m
c a r r y w i t h i n itself a b o u n d l e s s p o w e r of m e t a m o r p h o s i s , or m u s t it
necessarily become extinct after having r u n through a determined
cycle ?
IIi. T h e s o l u t i o n of t h e s e q u e s t i o n s is of g r e a t i m p o r t a n c e f o r the biologist,
n o t s i m p l y f o r the e n o r m o u s i n t e r e s t w h i c h t h e y offer i n t h e m s e l v e s ,
b u t also b e c a u s e of t h e i r applications.
IV. Mr. Dollo is of t h e o p i n i o n :
1. t h a t e v o l u t i o n o c c u r s b y r a t h e r r a p i d j u m p s .
2. t h a t a n o r g a n i s m c a n n o t r e t u r n , e v e n partially, to a f o r m e r state
a l r e a d y realized i n the series of its ancestors.
3. t h a t all o r g a n i s m s m u s t n e c e s s a r i l y b e c o m e extinct, a f t e r h a v i n g
r u n t h r o u g h a d e t e r m i n e d cycle w h i c h m a y , h o w e v e r , be e x t r e m e l y
long.
H e e x p r e s s e s t h i s b y saying:
E v o l u t i o n is d i s c o n t i n u o u s , irreversible a n d limited.
V. T h e a u t h o r t h e n p r e s e n t e d h i s r e a s o n s f o r t h i n k i n g t h a t this m u s t be
so. T h e n h e cited a l a r g e n u m b e r of e x a m p l e s , d r a w n f r o m living a n d
fossil a n i m a l s a n d f r o m living p l a n t s , to s u p p o r t h i s v i e w p o i n t s .
VI. Mr. Dollo is h a p p y to s a y t h a t h i s ideas h a v e b e e n a c c e p t e d b y h i s
m e n t o r , Mr. A. Giard, P r o f e s s e u r ~t la Sorbonne, a n d by h i s good
f r i e n d , Mr. P. Pelseneer, P r o f e s s e u r ~ l']~cole n o r m a l e de Gand.
He t h a n k e d t h e s e two n a t u r a l i s t s f o r the cases of d i s c o n t i n u i t y
a n d irreversibility w h i c h t h e y h a d so k i n d l y c o m m u n i c a t e d to h i m
(Mr. Giard: c r u s t a c e a n s , p l a n t s ; Mr. Pelseneer: m o l l u s c s ) .
He t h a n k e d two o t h e r of h i s b e s t f r i e n d s as well: Mr. J. M a s s a r t ,
A s s i s t a n t h l ' I n s t i t u t b o t a n i q u e de r U n i v e r s i t ~ de Bruxelles, w h o
p o i n t e d out to h i m m a n y i n t e r e s t i n g f a c t s related to d i s c o n t i n u i t y a n d
irreversibility a m o n g p l a n t s ; a n d Mr. G. B o u l e n g e r of the B r i t i s h
M u s e u m w h o called h i s a t t e n t i o n to n u m e r o u s a s p e c t s of t h e structure of living r e p t i l e s w h i c h h a v e a c o n s i d e r a b l e b e a r i n g o n t h e s e
questions.
He also m e n t i o n e d w i t h s a t i s f a c t i o n t h a t Mr. L. E r r e r a , P r o f e s s e u r
l'Universit6 de Bruxelles, agrees at l e a s t p a r t i a l l y w i t h h i s views.
Finally, i n c o n c l u s i o n , h e s t a t e d t h a t Mr. P. Hallez, P r o f e s s e u r
la Facult6 des Sciences de Lille, f o l l o w i n g h i s m o s t r e c e n t s t u d i e s o n
w o r m s , c o n c l u d e d t h a t evolution w a s d i s c o n t i n u o u s .
VII. Mr. Dollo w a s led to t h e s e g e n e r a l i z a t i o n s t h r o u g h h i s specialized
r e s e a r c h o n f o s s i l b o n e s t h a t h e h a s b e e n p u r s u i n g f o r twelve y e a r s at
the B r u s s e l s M u s e u m .
He first a n n o u n c e d t h e m i n h i s c o u r s e at the Solvay I n s t i t u t e
(University of Brussels) (Signed l e s s o n of N o v e m b e r 12, 1890).
Later h e r e t u r n e d to t h e m , n o t a b l y i n Giard's B u l l e t i n ( S e p t e m b e r

211

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20, 1891) a n d i n this Society's Bulletin (October 25, 1892).

VIII. T h e a u t h o r noted w i t h p l e a s u r e t h a t h i s ideas h a v e b e e n adopted
w i t h o u t r e s e r v a t i o n by Mr. A. L a m e e r e , P r o f e s s e u r ~ r U n i v e r s i t 6 de
Bruxelles, i n h i s E s q u i s s e de la Zoologie (Brussels, 1892) a n d i n the
syllabus of h i s Cours s u r le T r a n s f o r m i s m e (University e x t e n s i o n ;
l e s s o n 3; 1893).
IX. Mr. Dollo i n t e n d s to g a t h e r together i n a s m a l l illustrated v o l u m e all
t h e i m p o r t a n t cases of d i s c o n t i n u i t y , irreversibility, a n d l i m i t a t i o n
collected by h i m a n d h i s friends. 6s
X. Is this to say t h a t the laws e n u n c i a t e d above are t h e only ones w h i c h
direct t h e evolution of o r g a n i s m s ? Not at all. T h e r e are m a n y other
f u n d a m e n t a l laws: E x a m p l e s : the l a w of recapitulation, the law of
n e c e s s a r y regression, etc.
Short Bibliography of Dollo's W o r k on Irreversibility
"'Les Lois de l'$vohition," Bull. Soc. belge Gdol. Pal. Hydr., 7 (1893), 164-166.
"Sur la phylog~nle des d i p n e u s t e s , " Bull. Soc. belge Gdol. Pal. Hydr., 9
(1895), 79-128.
"Macrurus Lecointei, poisson abyssal n o u v e a u , reeueilli p a r l'exp~dition
a n t a r c t i q u e belge," Bull. Acad. r. Belg. Cl. Sc., (1900), 383-401.
"Sur l'origine de l a tortue l u t h (Dermochelys coriacea)," Bull. Soc. r. Sci.
mddic, naL Bruxelles, 59 (1901), 17-40.
"'Eochelone brabantica, Tortue m a r i n e nouvelle d u Bruxellien (Eocene
m o y e n ) de la Belgique," Bull. Aead. r. Belg. C1. Sci., (1903), 792-801.
" U n n o u v e l opercule t y m p a n i q u e de Plioplatecarpus, M o s a s a u r i e n plongeur,"
Bull. Soc. belge Gdol. Pal. Hydr., 19 (1905a), 125-131.
"Les D i n o s a u r i e n s adapt~s ~t la vie q u a d r u p ~ d e secondaire," Bull. Soc.
belge Gdol. Pal. Hydr., 19 (1905b), 441-448.
"Les Ptyctodontes s e n t des arthrod~res," Bull. Soc. belge G~ol. Pal. Hydr.,
21 (1907), 97-108.
"La Pal~ontologie dthologique," Bull. Soc. belge Gdol. Pal. Hydr., 23
(1909a), 377-421.
"Les P o i s s e n s voiliers,'" Zool. Jahrb., 27 (1909b), 419-438.
"Les C~phalopodes adapt6s ~t la vie n e c t i q u e secondaires et ~t la vie
b e n t h i q u e tertiare," Zool. Jahrb. Suppl., 15 (1912), 105-140.
"Podocnemius congolensis, tortue fluviatile nouvelle d u Montien (Pal~oc~ne
in~6rieur) d u Congo et r~volution des c h e l o n i e n s fluviatiles," Ann. Mus.
Congo belge, C~ol. Pal. Miner., serie 3, Bas et Moyen Congo, 1 (1913),
47-65.
"'Les C~phalopodes d~roul~s et l'irreversibilit~ de r~volution," Bijdragen
tot de Dierkunde, (1922), 215-227.
68. Such a v o l u m e w a s n e v e r published.

212