A  philosophical  critique  of  the  arguments  presented  in  The  Spandrels  of  

San  Marco  and  the  Panglossian  Paradigm:    

A  Critique  of  the  Adaptationist  Programme  

The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Program Stephen Jay Gould & Richard Lewontin Proceedings of the Royal Society B: Biological Sciences 205 (1979): 581-589

Christopher M. Anderson, MS Department of Biology- Evolutionary Theory & Complex Behavioral Biology chrisanderson.bio@gmail.com

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The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the

Adaptationist Programme is one of the most infamous papers that exist in the field of

evolutionary biology. Stephen Jay Gould and Richard Lewontin, the authors of the 1979

manuscript, are two of the most influential scientists in the field of evolution. With such

power comes great responsibility, and they felt the need to debilitate the adaptationist

programme. Gould and Lewontin, here on referred to as Gould and Lewontin, wrote the

Spandrels of San Marco to argue against the adaptationist programme and it’s

methodology, believing it to be flawed and full of inaccuracies.

In this paper, I give a background on the field of evolutionary biology, the

adaptationist program, and the spandrel analogy in order to lay the foundation of the

debate. I then argue against 4 major points presented by Gould and Lewontin against

adaptationism. (1) I argue that their criticism of adaptationist’s “inappropriate

atomization” [of organisms into distinct traits] is unsupported, and that this argument is

overbearing in that it assumes that this is common practice amongst adaptationists. (2) I

argue that their antagonistic views of trying to explain the adaptive trait when one attempt

fails, is contrary to the axiological philosophy of scientific progress. (3) I argue that

Gould and Lewontin use Darwin’s late work, which contains pluralistic notions, as a way

to disprove adaptationism, but that in doing this, they choose only what they want and

disregard Darwin’s earlier work, which favors selectionism. (4) I argue against the most

integral component of their paper, the Panglossian paradigm. The original analogy

presented by Voltaire against Leibniz optimism is faulty, which leads to an attenuation of

Gould and Lewontin’s parallel use of the Panglossian Paradigm. Though The Spandrels

of San Marco is a staple in evolutionary literature, Gould and Lewontin do not

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successfully discredit adaptationism and prove to the readers that the adaptationist

program is flawed in theory or practice.

A Brief History of Evolutoinary Biology, Adaptationist Program, and Spandrels

Evolution is a concept that is relatively new, and not fully understood. The

scientist that really made it a branch of science was none other than Charles Darwin.

After he published On the Origin of the Species in 1859, the groundwork for evolutionary

biology had been laid. His ideas of the survival of the fittest, i.e. natural selection,

presented the public at large with the idea that the diversity of life came about from a

common ancestor, and that populations evolve over the course of generations (Darwin,

1859). When it was first published, the idea of transmutation of species, referred to now

as speciation, was highly controversial and conflicted greatly with the beliefs presented

by natural theology. At the time, the Church of England ruled highly and dismissed

Darwin’s ideas immediately, preaching to the public at large that he was wrong. Thus

started the 50 years known as the “eclipse of Darwinism,” where many scientists

accepted evolution, but the idea of natural selection driving it was believed false (Bowler,

1983). From 1880-1930, Darwin’s ideas on natural selection were all but lost in science,

to later surface and aid scientists in their quest to understand the process of evolution.

Fast-forward to 1966 and the publishing of the book entitled Adaptation and

Natural Selection: A Critique of Some Current Evolutionary Thought written by

evolutionary biologist George C. Williams. Williams wrote this book to outline what is

referred to as “gene-centered view of evolution,” which holds that evolution is driven by

competing genes, with the alleles that produce the more successful phenotypes increasing

in frequency (Williams, 1966). This view contrasted the popular organism-centered view,

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which was the adopted theory of evolutionary biologist during the eclipse of Darwinism.

It also strongly debated the idea of group selection. With popularization of Williams’

book and spread of ideas, the adaptationist programme was developed. The adaptationist

program’s followers, known as adaptationists, adhere to methods that distinguish

adaptations from traits arisen through other process, which is collectively known as

adaptationism.

Adaptationists, such as John Maynard Smith, W.D. Hamilton, Richard Dawkins,

and Ernst Mayr, work to show that organisms are reasonably well adapted, and that these

adaptations arose from natural selection (Mayr, 1983). However, adaptationism has many

critics, including Stephen J. Gould and Richard Lewontin. Critics of adaptationism

believe that adaptationists assume most, if not all traits, are “optimal” adaptations, and

that the logic behind adaptationism over emphasizes natural selection as having such

power as to cause an evolutionary optimum, i.e. organisms that are flawless.

Gould and Lewontin (1979) begin their infamous manuscript against the

adaptationist program with a description of St. Mark’s Cathedral’s dome. Gould and

Lewontin explain that the dome contains quadrants, of which each meets one of the four

spandrels; a spandrel being described as a “tapering triangular space… a necessary

architectural by-product (G&L, 1979). They start their long-winded analogy by stating

that spandrels exist because of architectural constraints, that in order to have a dome,

these open triangular spaces need to be there. Gould and Lewontin go on to say that

though they are by-products of a greater design, they are often used for ornamental and

visually pleasing effects (1979). Thus, because they are a necessary by-product, but also

have an appropriate secondary effect, spandrels are not immediate adaptations to local

conditions (1979). Just as Dr. Pangloss said “Everything is made for the best purpose.

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Our noses were made to carry spectacles, so we have spectacles,” so too would

adaptationists argue that spandrels are not a by-product, but exist for a specific reason.

A background into the history of evolutionary biology is needed to understand the

roots of natural selection and the debate that has been raging on since Darwin. An

explanation of the Adaptationist Program and it’s critics is needed to comprehend the

animosity that exists between Gould and Lewontin and adaptationists. Finally, an

introduction of the spandrel analogy that Gould and Lewontin employ is needed to serve

as the backbone for their critiques. Now, I will show that Gould and Lewontin’s attempt

to depose the adaptationist program was unsuccessful through the use of various

examples and analogies of my own. Though Gould and Lewontin do make some valid

points, their overall goal of showing that adaptationism is flawed, is ineffectual.

Argument 1: Atomization and Trade-offs

In this section, I am going to analyze Gould and Lewontin’s statement that the

adaptationist program generally proceeds in two steps: atomization followed by the

introduction of trade-offs, which is fallible in theory and practice. I will then present

evidence given in support and against their claim, with additional quotes from various

key researchers in evolutionary biology to contradict Gould and Lewontin’s argument.

Gould and Lewontin insinuate that atomization of an organism has the potential to

proceed to the smallest level, i.e. cannot break the organism down any further. However,

I believe that this argument is flawed in that researchers commonly examine lower levels

of analysis, which can be used to look at functional genomics and changing genotypes.

With this ability to see evolution at the most intricate level, it is possible to tell what the

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proper levels of organismal trait break up is. Therefore, atomization of an organism is not

a negative aspect of the adaptationist program.

The first argument presented by Gould and Lewontin that I believe to be unfair

and amiss is their statement that the adaptationist program proceeds in two ways, firstly

implying that all adaptationists atomize organism into distinct traits (Gould and

Lewontin, 1979). According to Gould and Lewontin, this initial atomization is followed

by an attempt of explanation for each trait as a structure formed by natural selection to

produce optimum function (Gould and Lewontin, 1979). Atomization is used here to

mean breaking the organism into pieces, looking at each individual component. However,

what Gould and Lewontin do not make clear is whether their issue lies in breaking the

organism into bits, breaking the organism into the wrong bits, or breaking the organism

repeatedly into the smallest possible bits. I do not believe that even Gould and Lewontin

would debate the need to look at single traits as being influenced by selection. Legs do

not simply walk around on their own, they must be attached to a body, so we see here that

a certain amount of atomization is needed. However, Gould and Lewontin do not give

detail about what level they believe to be inappropriate, resulting in an inherent flaw in

their argument against adaptationist’s atomization of traits.

For example, if we were to try and understand the function of a car, Gould and

Lewontin propose that adaptationists would look at differences in separate structures (like

the tires, hubcaps, engine, fuel rods, windshield, etc.) as a function of the total car.

However, as stated above, a certain level of atomization is needed, but we do not know

how far Gould and Lewontin believe atomization should be taken before it is considered

a vice. Furthering the car analogy, the question is posed: would breaking the car into 2

separate components be too much? Would breaking it down by mechanical systems be

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too much? How many pieces can the car be broken down into before Gould and

Lewontin believe it detrimental? Surely they do not believe the only way to understand

the function of the car is to stare at it from the outside; they must believe that explanation

of function requires details about specific parts of the whole, i.e. atomization.

Since organisms must be atomized to an extent when looking at specific traits, the

only argument that would make sense for Gould and Lewontin to pose against

adaptationism is as follows: adaptationists atomize so much in their attempt to explain a

trait, that they break it down into such vastly small components, causing the “big picture”

to be lost; i.e. looking at the screws, nuts, iron, paint, coating, etc. of the motor

independently would not be able to explain the function of the car at large.

A biological illustration given by Gould and Lewontin as “their favourite

example” is about the human chin (Gould and Lewontin, 1979). By regarding the chin as

a “thing” instead of the product of interaction between the alveolar and mandibular

growth fields, than we make a misguided conclusion of it’s origin (recapitulatory) that is

completely different to the generally accepted explanation (neotenic) (1979). An example

using a human anatomical trait lets people make the transition from the inanimate,

architectural spandrel to a part of a living organism that they understand well (humans).

Ernst Mayr states matter of factly that it would be senseless to continue to atomize

an organism into smaller traits, for this would just be a search for an ad hoc adaptation of

each smallest component (Mayr, 1983). By saying that adaptationists atomize organisms

into each distinct trait, Gould and Lewontin portray the adaptationist program in a way

that is easily refuted. However, a motif that Gould and Lewontin do not mention, but use

to their advantage, is the argument from a reductionist point of view (Mayr, 1983). It is

very possible that various traits of an organism could have been acted upon by natural

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selection independently of others; to say that this is implausible would go against most of

the research by evolutionary biologists. Though Gould and Lewontin do rightly point out

some instances where traits had been acquired as adaptations remote to distant ancestors,

which do not serve the same purpose anymore, i.e. vestigial organs and exapted traits,

they fail to explain that these traits are still fundamental to the developmental system at

large, so cannot be excluded (Mayr, 1983).

The second part of Gould and Lewontin ’s argument that “adaptationists studies

proceed in two steps,” is the introduction of the idea of “trade-off” after attempts of

atomization fails (Gould and Lewontin, 1979). Here, trade-off means an organism retains

the greatest concession amongst contending demands. For example, the balance between

natural selection for locomotive ability in Gambusia hubbsi, and the sexual selection for a

trait affecting locomotive ability, which have different optima under varying ecological

conditions. Though natural selection calls for speed, sexual selection could select for a

trait that reduces speed, and a trade-off has been made. Trade-offs are commonly seen in

nature and help explain evolutionary traits in many organisms.

With the understanding of trade-offs, we can see why the next point does not

make sense. In their use of the Panglossian Paradigm, Gould and Lewontin call upon

Voltaire’s character, Dr. Pangloss, and his explanation of why he suffered from a

venereal disease. Dr. Pangloss explains: “It is indispensable in the best of worlds. For if

Columbus, when visiting the West Indies, had not caught this disease, which poisons the

source of generation, which frequently even hinders generation, and is clearly opposed to

the great end of nature, we should have neither chocolate nor cochineal” (Voltaire, 1759).

By using this quote, Gould and Lewontin insinuate that adaptationists are Panglossian

and believe that each attribute and idiosyncrasy is meant to be.

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The argument posed by Dr. Pangloss, which Gould and Lewontin ascribe that the

utility of end means, i.e. chocolate, makes this a metaphor for adaptationist thought.

However, David Queller presents the idea that this Panglossian example corresponds

expertly with the 2nd of five alternatives that Gould and Lewontin present against

adaptationism (Queller, 1995); there exists no adaptation or selection on the part at issue,

rather form of the part is a correlated consequence of selection directed elsewhere (Gould

and Lewontin, 1979). As Queller (1995) describes, Dr. Pangloss’ “part” at issue is the

venereal disease, but he does not reason it as distinct or adaptive on it’s own. Rather,

Pangloss explains the meaning of this “part” in ambience with the chocolate that

Columbus procured on this perilous journey. Gould and Lewontin use this example to

argue that the functionality of chocolate makes it like the examinations made by

adaptationists; it is not. Further, we see the switch of Dr. Pangloss from a supposed

atomist adaptationist, to a pluralist at heart (Queller, 1995). Gould and Lewontin “pulled

a fast one” on the readers by making Dr. Pangloss fit their purpose, in order to continue

the Panglossian extended metaphor. By doing so, they illegitimated Dr. Pangloss as a

useful character for their argument, and further dashed the effectiveness of the paradigm.

Argument 2: If one adaptive argument fails, try another

In this section, I am going to address Gould and Lewontin’s distaste for the

supposed common styles of argument that adaptationists employ. By showing that

adaptationist logic allows for the inclusion of both sexual selection and natural selection

in explaining the formation of traits, I argue that adaptationists’ exploration for

alternative explanations is beneficial to understanding the evolution of traits. I am going

to show that their animosity towards this argument style does not sync with scientific

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theory, and does not work as an agent of defamation against adaptationism. Additionally,

I will show that such logic is an asset to the adaptationist program, leaving us unable to

conclude that adaptationists are wrong in practice and theory.

Adaptationists base their methods of theory and practice on the plausibility that

every trait is, matter of factly, formed by natural selection (Mayr, 1983). The opponent to

form by natural selection is a trait that is formed by chance. In current scientific aptitude,

the ability to tell matter of factly that traits seen in the course of evolution have been

caused by purpose or chance will never exist. Scientists, such as evolutionary biologists,

geneticists, behavior ecologists, etc. have so far been able to deduce causation between

natural selection and manifestation of a trait due to vast amounts of research on selection.

It is only after repeated unsuccessful attempts to explain the formation of a trait by

natural selection or sexual selection (or by reason of genetic drift) that researchers can

conclude generation by chance.

Gould and Lewontin support their argument against trying to find an alternative

adaptive argument when one fails by giving an example from a book by Davitashvili

(1961). This model system concerns traits once believed to be caused by defense against

a predator, which were later deduced to be formed because of intraspecific male

competition (Gould and Lewontin, 1979). Davitashvili reports that a plethora of external

structures, such as horns, antlers, or tusks, had previously been viewed as weapons that

males utilize against predators. However, in his 1961 book, Teoriya Polovogo Otbora

(translated into English as “Theory of Sexual Selection”), Davitashvili claims that these

structures are actually formed from intraspecific competition between males, which is a

vastly different (but still adaptive) explanation than the first.

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Though Gould and Lewontin present examples, they fail to prove them as

negative in terms of scientific prowess. As stated, Gould and Lewontin believe that

adaptationists use natural selection to try and explain all traits (Gould and Lewontin,

1979); but what about sexual selection? For instance, lets examine the model system

given by Davitashvili (1961), the external structures formed as a result of intraspecific

competition between males. Intraspecific competition between males could be a means of

creating hierarchy or caused over resource availability (indicating natural selection).

However, it could be a result of sexual selection; if the females select males with

enlarged external structures, than the males with those structures will more successfully

mate than those who lack them. Though it is the female’s choice (sexual selection)

playing a role, it creates competition between males because natural selection will take

out the genes of the males not mated with, i.e. the genes causing smaller external

structures. Here we see that sexual selection could explain the traits seen by Davitashvili.

Gould and Lewontin successfully provide examples of switches between adaptive

explanations that coincide with natural selection. However, I have shown above that

sexual selection can drive pressure for specific traits. Whether an organism’s fitness

increased because of its success in the wild, or it’s fitness increased because of additional

mating opportunities, an adaptationist approach could explain the evolution of the trait.

Note: it is important to note how fitness is measured. If fitness is measured by survival

alone, than natural selection is acting on the trait; if fitness is measured by ability to mate,

than sexual selection is acting via mate choice.

To justify this logic I present the example from current research being conducted

at North Carolina State University. In the Bahamas mosquitofish, Gambusia hubbsi, the

presence of a predator in certain blue holes has caused the rapid evolution of male

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genitalia; more specifically, the distal gonopodial tip (Langerhans, 2011; Rosen and

Gordon, 1953). Though gonopodium morphology is acted on by natural selection, i.e. a

bigger gonopodium would reduce burst speed-time of G. hubbsi, reducing it’s fitness and

increasing it’s likelihood of being caught by a predator (Langerhans et al. 2005; Heinen

and Langerhans, in review), evidence now exists supporting a distinct role of sexual

selection by the female for gonopodia with higher surface area (Langerhans and Reznick,

2010).

For the surface area of the gonopodium in G. hubbsi, natural selection is acting

directly by it’s influence on locomotive ability; too big of gonopodium is a reducer of

fitness. For the gonopodial tip, selection is acting indirectly, i.e. natural selection is not

explicitly acting on the trait, by changing the context of sexual selection (J. Heinen,

unpublished conversation; unreferenced). Here, we see that ecological factors behind

natural selection of phenotypic morphology also play an indirect role in driving sexual

selection to favor a specific trait.

Natural selection and sexual selection are intertwined and connected in such a

way that natural selection is a constituent of sexual selection in most cases (exception

being sexual selection of traits caused by random mutation). With both natural selection

and sexual selection fitting into adaptationist thought, the vast majority of explanations

for a trait to exist fall under their logic. This debunks the fundamentals behind Gould and

Lewontin’s argument that searching for an alternative adaptive explanation is a negative

idiosyncrasy. However, Gould and Lewontin cover their proverbial behind here by

stating that newer interpretations by scientists might be right, but that they don’t know

(Gould & Lewontin,1979), which opens a new set of philosophical doors to disparage

their deportment of the adaptationist program.

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From The Spandrels of San Marco, I quote “We do not attack these newer

interpretations; they may all be right” (Gould and Lewontin, 1979). By admitting that

newer interpretations may be right means that there is a chance they are not correct. On

top of this, they immediately follow with “…[we question if] the failure of one adaptive

explanation should always simply inspire a search for another of the same general form”

(1979). Either side does not debate the fact that adaptationists do look for an alternative

adaptive explanation in response to a failed attempt; they both admit that’s part of

adaptationist philosophy (though they disagree on its significance). However, nowhere

else in the paper do Gould and Lewontin state that adaptationists look for reasons of the

same general form. What constitutes an explanation to be of the same general form as the

previous? This accusation cannot be validated by Gould and Lewontin, as there is no data

showing that adaptationists only look for alternative explanations of same general form.

Further, Gould and Lewontin do not define the meaning of the “same general form;” it

could refer to another explanation which explains the role of natural selection, or it could

refer to a trait formed by any type of selection, or anywhere in between. By using the

phrase “same general form,” but not giving parameters on the extent that general refers

to, they lose the ability to garnish this point against the adaptationist program.

Finally, the last and most obvious squabble I have with Gould and Lewontin’s

logic is that it goes against the fundamental backbone of scientific inquiry. The

publication of new research sparks novel ideas and new hypotheses daily. As knowledge

is poured into our greater understanding of science, scientists constantly revise their

existing hypotheses in light of new evidence. Pre-existing hypotheses do not have to be

discarded, but many change slightly to take alternate forms. The existence of data that

falsifies an adaptive hypothesis does not mean that all attempts of adaptive explanations

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should cease; it means that ideas need to be modified. If all researchers stopped

investigating hypotheses because new data appeared that disproved their idea, science

would never move forward. I believe scientific investigation follows the age-old saying:

“If at first you don’t succeed, try and try again.”

Argument 3: Darwin’s own pluralistic approach

In this section, I will examine Gould and Lewontin’s use of Darwin throughout

the paper, and their claim of his pluralistic approach to identifying agents of evolutionary

change as a mechanism to admonish the adaptationist program. Their use of the logic

“Darwin thought it, and Darwin is a saint amongst biologists, therefore it must be right”

is adverse to all philosophical arguments. It is a form of elementary argument that proves

nothing. Further, I will cite Queller (1995) and show that of the multiple editions of On

the Origin of Species, Gould and Lewontin ignore the earlier works and try and convince

readers that his late edits were a true representation of Darwinism. I will present

Queller’s idea that Darwin was influenced by new hypotheses, for example: blending

inheritance, so tried “sloppily” to alter his previous work to reflect these new ideas. By

calling solely upon Darwin’s later work, which is the only portion that includes pluralistic

notions, I believe that Gould and Lewontin do not represent Darwinism, or Darwin

himself in a proper light. Therefore, their inaccuracy in portrayal of Darwin makes a

precarious argument at best, so does not remain useful in trying to disprove the

adaptationist program.

Firstly, it is important to note that justifying an argument on the basis of using a

figurehead, like Gould and Lewontin do with Darwin, is contemptible. No matter what

the argument, stances do not become true just because someone important says it is so.

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Gould and Lewontin (1979) appeal to Darwin, saying that he has attained sainthood (if

not divinity), in order to prove a point. It follows the logic, “if a saint like Darwin thought

it true, we need to listen.” This argumentation is improper and should never be used by a

researcher to prove a point. However, because Gould and Lewontin do use it on their

behalf, I will show that their exposition is weak at best.

On the first page of The Spandrels of San Marco, Gould and Lewontin state

outright: “We support Darwin’s own pluralistic approach to identifying the agents of

evolutionary change” (Gould and Lewontin, 1979). Further along in the paper, Gould and

Lewontin dedicate an entire section to Darwin, entitling it “The Master’s Voice Re-

Examined;” they use this section to persuade the readers that Darwin was more of a

pluralist than selectionist, i.e. many factors versus just natural selection. To an educated

individual, this section of the Spandrel paper proves that Darwin believed consistently in

pluralism. However, to those people educated in the history of evolutionary theory and

Darwinism, it feels more like somebody trying to put falsities in an existing knowledge

base. “The Master’s Voice Re-Examined” is a bold attempt to convince us that if Darwin

believed in pluralism all along, but it does not disprove the adaptationist program’s

theory or methodology.

The pluralist methodology that Gould and Lewontin believe in allows for modes

of change, i.e. evolution, other than natural selection. Gould and Lewontin (1979) claim

that Darwin was a pluralist all along, and that nothing “no argument angered him more

than the common attempt to caricature and trivialize his theory by stating it relied

exclusively upon natural selection.” This is true to a point, but not a valid representation

of what Darwin believed. It took a long time for Darwin to adopt pluralistic views, and

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did so in response to new theories. Only in his latest edition of On the Origin of Species

(1872) did he address pluralism.

One of the new theories that got Darwin thinking was that of Fleeming Jenkin

(Queller, 1995). Jenkin, and evolutionist, proposed the idea of blending inheritance in the

year 1867. The blending theory of heredity shows that natural selection cannot operate in

terms of single variations (Jenkin, 1867). For example, it could show additional benefit to

the organ of interest inside an organism, but it does not lead to the development of new

organs, i.e. it cannot originate species (Bulmer, 2004). However, serious flaws existed in

Jenkin’s theory that was not pointed out until 1871 by A.S. Davis (Davis, 1871).

Assumptions made by the theory did not hold true in all cases, and therefore was

falsified. Nonetheless, by this time the damage was done and Darwin began to doubt his

own theories and logic, incorporating false doctrines into his thought process.

The point of giving background into new additions of ideas into heredity at the

time of Darwin is to show why Darwin started to incorporate pluralism into his theory of

evolutionary mechanisms. On the introduction of new theories that debated his own

research, like blending inheritance, Darwin began to search desperately for answers.

Examples of his attempts to understand heredity better can be seen in his adoption of

Lamarckian inheritance (which he had previously disregarded for all intensive purposes)

(Queller, 1995). Near the end of his academic life, Darwin tried his best to hold onto his

mechanisms of heredity, but include what he thought to be correct research. Since many

theories, including blending inheritance, were later disproven… casting Darwin in the

light of a proponent and die-hard believer in pluralism is not a true representation of him.

As Queller says, he will take the younger Darwin and the first edition of On the

Origin of Species when discussing Darwin’s beliefs. Though Gould and Lewontin are not

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incorrect by saying Darwin believed in pluralistic notions, their use of his quote from the

last edition of On the Origin of Species is just a caricature of the true Darwin. Therefore,

I believe that their use of Darwin is a misrepresentation of a pioneer in Evolutionary

Biology, and that their attempt to bend the truth hurts their argument more than it does

debunk the adaptationist program and their theory and practice.

Argument 4: Fundamentals of the Panglossian Paradigm

In this section, I argue against the usefulness of the fundamental paradigm used in

Gould and Lewontin’s paper. I argue that Gould and Lewontin use the Panglossian

Paradigm unsuccessfully, in that Voltaire misinterpreted Leibniz Optimism from the

beginning. Leibniz Optimism, for function of this argument, notions that things are the

best of what is possible; not that they are the best overall. Voltaire ridicules Leibniz

Optimism through Dr. Pangloss and his teachings, which Gould and Lewontin quote

numerous times, elucidating to traits formed as the optimum (perfectionist ideology).

Here I show that Gould and Lewontin really argue against perfectionism, which is not the

basis for Leibniz Optimism. Finally, the concept of local optimality in adaptationism

refutes the arguments that Gould and Lewontin pose against the adaptationist program.

Therefore, the Panglossian Paradigm does not successfully show that the adaptationist

program is flawed in theory and practice.

One of the major staples of this 1979 paper is the Panglossian Paradigm, exploited

by Gould and Lewontin to demonstrate the “ridiculousness” of the adaptationist program.

The Panglossian Paradigm gets it’s name from Voltaire’s 1759 Candide, which is about a

young man who is lectured in Leibniz Optimism by his mentor, Dr. Pangloss. By

appealing to Dr. Pangloss, seen in modern times as an absurd laughable character, Gould

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and Lewontin insinuate that adaptationists assume that all traits are perfectly designed.

However, the flaw in Gould and Lewontin’s use of the paradigm is not one of immediate

evidence. Gould and Lewontin protest the perfectionist ideal of Leibniz Optimism that

Voltaire introduces in Candide, but Leibniz Optimism does not preach perfectionism.

One of the numerous quotes that Gould and Lewontin (1979) use from Dr.

Pangloss, theoretically to show the foolishness of adaptationist thought, is: “…

everything is made for the best purpose. Our noses were made to carry spectacles, so we

have spectacles. Legs were clearly intended for breeches, and we wear them” (Voltaire,

1759). In the context of adaptationism, Gould and Lewontin imply the belief of all traits

occurring for a specific, perfect purpose, is core to their philosophy. In the context of

Candide, Voltaire is using Dr. Pangloss to demonstrate that Leibniz Optimism is flawed

and comical. In order to understand why Voltaire would have Dr. Pangloss say such

absurd things, a base knowledge of Leibniz Optimism is needed.

Gottfriend Wilhem Leibniz, referred to here on out by his last name, was a

mathematician and Philosopher that lived from the mid 17th century to the early 18th

century (Adams, 1994). On top of his development of infinitesimal calculus independent

of Sir Isaac Newton (1994), he is well known in philosophy for his optimism, i.e. Leibniz

Optimism (Adams, 1994). Leibniz Optimism is built around his belief that God created

this world as the best of possible worlds (Franklin, 2003).

The 17th and 18th centuries were a time when Christian theism ruled over science

and philosophy. Leibniz believed that people were taught that God creates everything for

the best reason; that apparent flaws in the world must exist in order to permit over all

good. Therefore, he believed that this must mean that people believe this as the best

world, since God does not create imperfections (2003).

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Voltaire uses Candide to show his discontent with perfectionism, which as shown

above, is not part of Leibniz Optimism (Mayr, 1983). In theology, perfectionists believe

that God creates the best world; in evolution, Gould and Lewontin accuse adaptationists

of believing that evolution produces the best trait. However, the reality of the matter is

that Leibniz Optimism proclaims God creates the best world that is possible, i.e. not

perfect, but as good as it can be based on the circumstances. This philosophy is paralleled

in adaptationist thought concerning adaptations; traits are not perfect, but created to be

the best they can be with the various constraints placed on the organism.

To better explain the axioms of Leibniz Optimism, I present the evolutionary

concept of local optimality. Evolutionary Biologists use the term local optimality to

describe a specific trait as the best that can be from the given conditions, constraints, and

tradeoffs (Driscoll, unpublished work, uncited). This correlates to the theological

philosophy proposed by Leibniz. Adaptationists believe in local optimality, not

perfectionism.

In summation, Voltaire’s Candide (1759) was written to show distaste in Leibniz

Optimism, which was interpreted by Voltaire to mean that God created the best world

(Mayr, 1983). Gould and Lewontin (1979) used Candide’s character, Dr. Pangloss, as a

figurehead for their Panglossian Paradigm. The quotes Gould and Lewontin utilize in

their paper are meant to show the ridiculousness of the adaptationist program. However,

since Voltaire misinterpreted Leibniz Optimism, the reason behind Dr. Pangloss’ quotes

is nullified for the sake of Gould and Lewontin’s argument. Dr. Pangloss preaches

against the idea of perfectionism, not local optimality. Since adaptationists do not believe

in perfectionism, Gould and Lewontin’s Panglossian Paradigm does not serve the

function to discredit adaptationist thought. Further, since the Panglossian Paradigm, a

  19  

crucial portion of The Spandrels of San Marco and the Panglossian Paradigm (1979)

paper, is based upon shaky foundation at best, the overreaching power of the metaphor

does not prove to be as a tool for defaming adaptationism.

Importance of The Spandrels of San Marco and the Panglossian Paradigm

I have written this critique of The Spandrels of San Marco and the Panglossian

Paradigm: A Critique of the Adaptationist Programme (1979) to highlight the

shortcomings in argument against the adaptationist program. Gould and Lewontin, two of

the biggest names in modern Evolutionary Biology, attempted to discredit the

adaptationist program by writing a paper with intricate analogies and pristine writing;

though proving to be unsuccessful. Despite not believing that their 1979 polemic

produced a favorable outcome on their part, I do believe that it is one of the most

esteemed papers to all students and scientists whom study evolution.

Gould and Lewontin have been criticized for using colorful metaphors and

fanciful rhetoric, believed by some critics as a way to demonstrate their advanced

knowledge (Queller, 1995). Though I do believe that Gould and Lewontin use wordiness

and eloquent language excessively in their opinionated paper, I do not believe it should

be reprized. Gould and Lewontin’s target audience are educated individuals, i.e. those

that are capable of interpreting the analogies, metaphors, and verbiage. I do not think that

their use of big words is an attempt to prove their extreme knowledge on the subject; they

did not need to prove that to anyone as they were already prominent scientists at the time.

In having philosophical charity, I believe that the paper is well written and follows an

easy-to-read format despite the ostentatious rhetoric.

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In terms of the strengths of the paper, I believe Gould and Lewontin do great job

of supporting their case. Gould and Lewontin present strong motions because they give

evidence to justify their claims. Whether the evidence presented truly makes their point is

questionable in many areas, but that has already been discussed. Further, Gould and

Lewontin do not continuously cite themselves in the paper; they look to many outside

sources to aid their overture. A plethora of resources, such as non-fiction books,

textbooks, published studies, and manuscripts were used through out the paper. Unless a

reader is definitively attempting to critique their arguments, or find flaws in their

archetypes as I have, there is little flaw to be found in their manuscript. The strongest

argument is one that bestows reinforcement.

Besides the four flaws outlined by this analysis, the pure intensity and amount that

Gould and Lewontin attack the adaptationist program is a vice of the paper. Instead of

reading like an assessment, with presentation of counterarguments… it reads like a

personal vendetta that never ends. Upon disagreement on a matter in philosophy and

science, the proper course of action is to write a rebuttal in a respectful manner. Though

the purpose should be clear, it should not perpetually attack. Gould and Lewontin do not

simply present counter arguments, they generate claims (such as the atomization example

discussed above) that are not necessarily true.

Further, with apparent disregard of academic courtesy, Gould and Lewontin over

exaggerate many components of the adaptationist program in order to drive their point in.

They utilize splendid metaphors, such as the Panglossian Paradigm, in attempt to make

the adaptationist program look comical. Though I knew the intent of the paper, it was

hard to read at times because I felt the need to defend adaptationism. Gould and Lewontin

show a lack of respect for alternate philosophies, or for anyone that holds convictions that

  21  

are contrary to their own. For this, I lost a lot of appreciation for them as authors, as well

as a loss of respect for the arguments put forward.

As presented, I make my point that Gould and Lewontin’s (1979) piece of

literature has it’s frailties and faults; that many of the arguments presented are not useful

in argument against the adaptationist program. However, The Spandrels of San Marco is

a key scientific report that should be respected. Many ideals that Gould and Lewontin

possessed, such as Marxism, played a role in their academic lives and analyses of other

scholars, but are not so erroneous that they overshadow the men as scientists. Both Gould

and Lewontin augmented the fields of evolutionary biology, ecology, genetics, etc. by

adding vast amounts of knowledge to the existing expanse. These two men, along with

researchers like Ernst Mayr, A.R.B. Haldane, Theodosius Dobzhansky, and Richard

Lewontin (among many others), have paved the road for future researchers to follow. The

importance of Gould and Lewontin to science can not be over stated.

Gould and Lewontin, along with their celebreated manuscript, The Spandrels of

San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme

(1979), have given so much to the scientific community. I chose to critique this particular

piece because of the challenge it presented in logic, knowledge, and argumentation.

Though I feel I have effectively demonstrated portions of their critique to be flawed, I do

not wish to take away the significance of this paper. Reading The Spandrels of San

Marco has made me a better student, as well as scientist. For that, I am forever grateful to

Stephen Gould and Richard Lewontin. This literary monument will continue to be a

staple in the study of evolution across the world, and I feel honored to be a part (though it

is in opposition) of the immortality that is Stephen Jay Gould and Richard Lewontin.

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*I would like to thank Dr. Catherine Driscoll, Dr. Miles Engell, Dr. R. Brian Langerhans,

Justa Heinen, and Elizabeth Hassell for all their help in the formation and execution of

this paper. I want to thank Dr. Driscoll for her patience, guidance, and help with logic and

philosophical thought. I want to thank Dr. Engell for her continued mentorship and

altruism, as well as being my academic role model and the reason I pursued further

education in evolution. I want to thank Dr. Langerhans for his training & expertise in

evolutionary ecology, which proved vital in the fulfillment of this analysis, as well as for

allowing me to work in his lab and be a part of evolutionary research. I want to thank

Justa for her generosity, patience, and kindness towards me as I learn my way in higher

learning and research. Finally, I want to thank Elizabeth for the goodness, heart, and

thoughtfulness that she has shown to me this semester; I have never known a kinder

person. From all these people, I have learned more about evolution and myself this

semester than at any other time in my life, and I will forever be grateful.

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Works  Cited:      Adams,  Robert  Merrihew.  Leibniz:  Determinist,  Theist,  Idealist.  New  York:  Oxford,     Oxford  University  Press,  1994.      Bowler,  Peter  J.  The  Eclipse  of  Darwinism:  anti-­‐Darwinian  evolutionary  theories  in     the  decades  around  1900.  Johns  Hopkins  Univsersity  Press.  1983      Bulmer,  Michael.  “Did  Jenkin’s  Swamping  Argument  Invalidate  Darwin’s  Theory     Of  Natural  Selection?”  The  British  Journal  for  the  History  of  Science.     37:3  (2004):  281-­‐297      Darwin,  Charles.  On  the  Origin  of  Species  by  Means  of  Natural  Selection,  or  the  

Preservation  of  Favoured  Races  in  the  Struggle  for  Life.  United  Kingdon:  John  Murray.  24  November,  1859.  

   Davis,  A.S.  “The  ‘North  British  Review’  and  the  Origin  of  Species.”  Nature.  5  :     (1871):  161      Davitashvili,  L.S.  Teoriya  Polovogo  Otbora  [Theory  of  Sexual  Selection].  Moscow  :     Akademii  Nauk.  (1961)      Franklin,  J.  “Leibniz’s  solution  to  the  problem  of  evil.”  Think  5:  (2003):  97-­‐101      Gould,  Stephen  J.,  and  Richard  Lewontin.  “The  Spandrels  of  San  Marco  and  the     Panglossian  Paradigm:  a  Critique  of  the  Adaptationist  Programme.”     Proceedings  of  the  Royal  Society  of  London  B.  205  (1979):  581-­‐598        Heinen,  J.L.,  and  Langerhans,  R.B.  “Predation-­‐associated  Divergence  of  Male  Genital     Morphology  in  Livebearing  Fish.”  In  Review.      Jenkin,  H.F.  “The  Origin  of  Species.”  North  British  Review.  46:  (1867):  277-­‐318        Langerhans,  R.B.  Genital  Evolution.  In  J.  Evans,  A.  Pilsastro  and  I.  Schlupp,  eds.    

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  Ecology  and  Evolution  of  Poecillid  Fishes.  University  of  Chicago  Press.  2011.      Langerhans, R.B., C.A. Layman, and T.J. DeWitt. 2005. Male genital size reflects a Tradeoff between attracting mates and avoiding predators in two live-bearing fish Species. Proceedings of the National Academy of Sciecnes USA. 102: 7618-7623    Langerhans,  R.B.  and  Reznick,  D.N.  Ecology  and  Evolution  of  Swimming       Performance  in  Fishes:  Predicting  Evolution  with  Biomechanics.  Pages  200  –     248  in  P.  Domenici  and  B.G.  Kapoor,  eds.  Fish  Locomotion:  an  Etho-­‐     Ecological  Perspective.  Enfield  Science  Publishers.  2010      May,  Ernst.  “How  to  Carry  Out  the  Adaptationist  Program?”  The  American  Naturalist.     121:3  (1983):  324-­‐334      Queller,  David  C.  “The  Spaniels  of  St.  Marx  and  the  Panglossian  Paradox:  A  Critique     of  a  Rhetorical  Programme.”  The  Quarterly  Review  of  Biology.  70:4       (1995):  485-­‐489      Rosen,  D.E.,  and  M.  Gordon.  1953.  Functional  anatomy  and  evolution  of  male       Genitalia  in  poeciliid  fishes.  Zoologica.  38:  1-­‐47      Voltaire.  Candide,  Ou  l’optimise.  France:  Cramer,  March-­‐Michel  Rey,  Jean  Jourse,     Lanbert,  and  others.  1759