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Ancestor of the new archetypal biology: Goethe’s dynamic typology as a model

for contemporary evolutionary developmental biology

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 Studies in History and Philosophy of Biological and Biomedical Sciences 44 (2013) 735–744

Contents lists available at SciVerse ScienceDirect

Studies in History and Philosophy of Biological and

Biomedical Sciences
journal homepage: www.elsevier.com/locate/shpsc

Ancestor of the new archetypal biology: Goethe’s dynamic typology

as a model for contemporary evolutionary developmental biology
Mark F. Riegner
Environmental Studies Program, Prescott College, Prescott, AZ 86301, USA

a r t i c l e i n f o a b s t r a c t

Article history: As understood historically, typological thinking has no place in evolutionary biology since its conceptual
Received 1 June 2012 framework is viewed as incompatible with population thinking. In this article, I propose that what I
Received in revised form 17 May 2013 describe as dynamic typological thinking has been confused with, and has been overshadowed by, a static
Available online 18 July 2013
form of typological thinking. This conflation results from an inability to grasp dynamic typological think-
ing due to the overlooked requirement to engage our cognitive activity in an unfamiliar way. Thus, ana-
Keywords: lytical thinking alone is unsuited to comprehend the nature of dynamic typological thinking. Over
Archetype
200 years ago, J. W. von Goethe, in his Metamorphosis of Plants (1790) and other writings, introduced a
Evo–devo
Goethe
dynamic form of typological thinking that has been traditionally misunderstood and misrepresented. I
Morphology describe in detail Goethe’s phenomenological methodology and its contemporary value in understanding
Theoretical morphospace morphological patterns in living organisms. Furthermore, contrary to the implications of static typolog-
Typological thinking ical thinking, dynamic typological thinking is perfectly compatible with evolutionary dynamics and, if
rightly understood, can contribute significantly to the still emerging field of evolutionary developmental
biology (evo–devo).
Ó 2013 Elsevier Ltd. All rights reserved.

When citing this paper, please use the full journal title Studies in History and Philosophy of Biological and Biomedical Sciences

Form is a moving, a becoming, a passing thing. The doctrine of
forms is the doctrine of transformation. The doctrine of metamor-
phosis is the key to all signs of nature.
J. W. von Goethe (quoted in Richards, 2002, p. 454)
1. Introduction
There are few concepts identified by evolutionary biologists
that have received more criticism than typological thinking and
essentialism. As Mayr (e.g., 1963, 1982, 1991, 1997) never tired
of pointing out (see also Chung, 2003), Darwin (1859) addressed
these notions and hoped to put them to rest by proposing that only
populations of variable individuals evolve, so-called population
thinking. Typological thinking was viewed as incompatible with
this evolutionary principle and thus had no place in evolutionary
causality (see Amundson, 2005 for in-depth critique of this posi-
tion). Indeed, it has become anathema to even allude to typological
explanations in one’s research program, notwithstanding recent
conceptual explorations in evolutionary developmental biology
that are more accommodating (e.g., Amundson, 1998, 2005; Hall,
1996; Jenner, 2008; Lewens, 2009a).
In this paper I hope to show that, first, historically a static mode
of typology has overshadowed a dynamic mode of typology. Conse-
quently, a straw man has been erected and perennially attacked,
but this caricature bears little semblance to the dynamic typology
proposed herein. Secondly, I will attempt to demonstrate how the
dynamic typological thinking implicit in Goethe’s research ap-
proach is perfectly compatible with a notion of evolutionary
change and, if rightly understood, not only stands as a forerunner
to modern evolutionary developmental biology (evo–devo) but
also speaks to current questions regarding the nature of evolution-
ary dynamics. In fact, recent insights in evolutionary developmen-
tal biology, including genetic regulatory networks, developmental
constraints, analysis of theoretical morphospace, developmental
trade-offs, and recursive properties of morphological evolution,

E-mail address: mriegner@prescott.edu

1369-8486/$ - see front matter Ó 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.shpsc.2013.05.019
736 M.F. Riegner / Studies in History and Philosophy of Biological and Biomedical Sciences 44 (2013) 735–744
reaffirm a notion of dynamic typology. However, to grasp dynamic
typological thinking, it is necessary to engage our cognitive activity
in an uncustomary way, and this is the very reason why this way of
seeing has been historically distorted: our practice of analytical
thinking falls short of what is necessary to apprehend the dynamic
thinking implicit in Goethe’s way of understanding nature. Accord-
ingly, I will explore in detail the subtleties of Goethe’s methodol-
ogy to show how it is not just an historical curiosity but has
contemporary relevance for how we formulate questions regarding
understanding morphological evolution. In this vein, I concur with
Richards (2002, p. 408) ‘‘that Goethe’s understanding of scientific
procedure marked him not simply a good scientist for the time,
but a good scientist for all time.’’
2. The dynamic nature of the archetype
Central to typological thinking is the notion of ‘‘archetype.’’
Descriptions of archetypes and their relationship to physical enti-
ties date back to Plato’s Republic and other writings. His oft-quoted
allegory of the shadows on the cave wall, ultimately mistaken for
the full reality, identifies the relationship of, for example, actual
organisms and the informing ‘‘Ideas’’ that give them shape. Accord-
ing to this interpretation, there exists an archetype, or eidos, of Cat,
for instance, and all actual cats are but mere imperfect shadows, or
approximations, of this nonphysical entity. These archetypes have
been taken to be ‘‘perfect,’’ whatever that may mean (it’s rarely
defined), constraining, and static, that is, unchanging and
unchangeable. (Such interpretations, in fact, may stem from
misinterpretations of Plato. Bortoft (2012, p. 82), for example, cites
H-G Gadamer, the Plato scholar: ‘‘Plato was no Platonist.’’) Accord-
ingly, such notions cannot support organic transformation on an
evolutionary timescale and lead to views of species fixism. If the
archetype is a nonphysical entity—not of this earth—how can it
be influenced by earthly processes? And if it exists in a state of
eternal perfection, how, and why, would it change? Why are living
organisms so variable and thus only imperfect expressions of their
respective presumed archetype?
To illustrate how this static interpretation has influenced biol-
ogy, we need look no further than to the pre-Darwinian era, for
example to Richard Owen’s search for the archetypal vertebrate.
Owen was determined to discover the ‘‘essence’’ of the vertebrate
body plan, the unifying principle, or Unity of Type, common to all
living vertebrate forms (Amundson, 2005; Gould, 2002; Owen,
2007[1849]; Richards, 1992, 2002; Rupke, 1993). Accordingly, he
distilled what he believed to be the key ingredients, the common
Fig. 1. Richard Owen’s ‘‘archetypal’’ vertebrate (1848); reproduced in Owen (2007).
denominators, of vertebrate architecture and constructed a blue-
print of basic, repeating skeletal features (Fig. 1). Owen, therefore,
abstracted elements of the vertebrate body plan and juxtaposed
them in a generalized configuration; accordingly, his was a ‘‘reduc-
tive theory’’ (Richards, 2002, p. 302). Clearly, Owen’s schema
looked like no actual vertebrate, living or extinct, although it had
a vague resemblance to a fish skeleton. In 1859, with the publica-
tion of Origin of Species, Darwin supplanted Owen’s hypothetical
archetype with the presumed actual ancestral vertebrate—the ‘‘un-
known progenitor’’—which gave rise to all subsequent vertebrates
through descent with modification (Amundson, 1998; Brady,
1987). Consequently, Owen’s influence was demoted, and along
with him the Naturphilosophie movement of which he was a part,
and apparently the problem of the vertebrate archetype, and
archetypes in general, was resolved.
Earlier, in 18th century Germany, however, a previous attempt
was made to ascertain the nature of the archetype, but this time
not only animals but geologic formations, meteorological phenom-
ena, and especially plants provided the focus (Amrine, Zucker, &
Wheeler, 1987). J. W. von Goethe’s (1749–1832) original research
on morphology—a term he coined (Nyhart, 1995)—sought to grasp
the unity disclosed through the diversity of a given class of phe-
nomena through ‘‘disciplined and cultivated perception’’ (Steiger-
wald, 2002, p. 293). Regarding plants, after many years of
detailed botanical observations, culminating in his celebrated Ital-
ian journey, Goethe some years later claimed to have experienced
what he called the Urpflanze, the archetypal plant, the basic trans-
formative element of which he termed ‘‘leaf’’ (Richards, 2002; Tan-
tillo, 2002). Unlike Owen, Goethe did not attempt to express his
archetype in a visual schema except for a few hastily scribbled
lines he once showed to his philosopher friend Friedrich Schiller
during an animated conversation. Oddly enough, only subsequent
self-proclaimed interpreters of Goethe have taken liberty to illus-
trate his archetypal plant; as discussed below, these efforts were,
and continue to be, based on a misguided notion of Goethe’s
archetype.
According to philosopher of science Brady (1987), after 1859
Goethe’s notion of the archetype suffered the same ignominious
fate as Owen’s. But was this justified? A superficial analysis would
nod in agreement, but, as Brady points out, this is based on a mis-
reading of Goethe. In contrast to Owen’s abstract schema, Goethe’s
notion of the archetype does not necessarily imply an ancestral
form (nor does it deny one) and, more importantly, requires a dy-
namic mode of cognition to be apprehended. Unlike a static blue-
print that serves as a distilled generalization and representation
 M.F. Riegner / Studies in History and Philosophy of Biological and Biomedical Sciences 44 (2013) 735–744
of the phenomena under observation, Goethe’s archetypal Plant is
a kind of fluid concept that unites the changing structures of the
plant and enables the observer to see each structure not as sepa-
rate but as multiple expressions of one ideal organ (Bortoft,
2012). Thus, it is a dynamic property, a unifying principle without
uniformity (Bortoft, 1996), that comes to expression through the
details of the diverse, spatially separated form elements. In other
words, it draws upon a dynamic typology, not the static typology
that has dominated Western thought through a likely misunder-
standing of Platonism (Bortoft, 1996, 2012). An example from
geometry may be helpful at this point.
From a Goethean perspective, the archetypal Triangle can never
be represented or schematicized, but its essential, ideal principles
inform any number of triangles we may wish to construct. Clearly,
while an infinite number of triangles can potentially be con-
structed, no single triangle represents the archetype, but, rather,
the essential attributes of the archetype are represented in every
triangle, otherwise the shape would not be a triangle but some
other construct. Thus the archetypal Triangle is the idea that uni-
fies—but does not generalize!—the diversity of shapes that are so
configured by three lines and three angles. Any triangle that we
may draw would be a specific triangle but never the archetype,
which exists only as a sort of unifying, or relational, idea that de-
fines the limits and possibilities of all potential triangles and only
partially comes to presence in any given triangle. How does Goe-
the’s archetypal Plant fit with this notion?
Some visual examples may be helpful here. The Common Sow-
thistle (Sonchus oleraceus) and Common Ragweed (Ambrosia artem-
isiifolia), as in many plants, exhibit pronounced heterophylly, that
is, the leaf shape changes dramatically from the base of the stem
to the apex. If the leaves are removed from the stem and arranged
in a linear sequence, one can readily notice the graded series of
transformation (Fig. 2). Goethe (1790) had a particular interest in
this phenomenon and maintained that in a given specimen there
is ‘‘one leaf’’ that undergoes ‘‘metamorphosis.’’ In other words,
there exists a unifying idea that comes to expression in—or better,
through—the numerous related shapes. This idea—which to Goethe
was a cognitive experience grounded in sensory experience—is not
Fig. 2. Examples of leaf metamorphosis (heterophylly): (a) Common Sowthistle and (b) Common Ragweed. In each example, the leaves on the left are from the base of the
plant while those on the right are from the apex. (a) from leaf pressings made by author; (b) courtesy of C. Holdrege.
737
a static representation or an abstraction but a dynamic movement,
a continuum, a living potential that belongs to the whole phenom-
enon of the plant as much as do the individual organs. Again, this is
not to be confused with a ‘‘general plan common to all organs’’
(Bortoft, 2012, p. 58). Rather, it is an expression of the plant’s
becoming, its temporal dimension. Accordingly, the leaves can be
conceived as frozen moments in the development of the plant, vis-
ible steps that enable the continuum to come to expression in an
observer’s consciousness. Although the continuum is not sensibly
perceived, it is, as Goethe maintained, a cognitive experience, for
if the leaves are mixed randomly, a student, never having seen
the plant, is able to order them correctly with relative ease by
weighing the differences against the similarities and thus ‘‘seeing’’
the movement in his or her mind’s eye. Furthermore, if an early
and later leaf of the sequence were removed and juxtaposed, their
relatedness would be questionable (for example, the third leaf
from the left and last leaf of each sequence of Fig. 2). But, as differ-
ent as the two leaves are morphologically, in the ‘‘context of move-
ment’’ (Brady, 1998) their relatedness is immediately
comprehensible.
The next step in apprehending the plant in its temporal devel-
opment is to observe how the floral organs are further metamor-
phic elements of the ‘‘leaf,’’ the ideal transformative impulse (and
not an actual leaf; see Richards, 2002, p. 396). In The Metamorphosis
of Plants (1790), Goethe carefully describes the transition to calyx,
petals, anthers, etc. as the plant continues its development beyond
the vegetative stage: ‘‘serial homology’’ in contemporary terminol-
ogy. Compared to tracing the foliar metamorphosis, a further cog-
nitive penetration of the phenomena is needed to identify the
unifying thread that weaves though the floral parts. In some cases,
however, the transition between foliar and floral organs is patently
obvious as seen, for example, in the transition between leaf and
bract in the neotropical heliconia (Fig. 3). In addition, after
200 years, Goethe’s proposal has been acknowledged and corrobo-
rated by genetic research that has identified how homeotic varia-
tion in plants can induce the development of different floral
organs (e.g., Dornelas & Dornelas, 2005; Friedman, 2009; Smyth,
2005; Theißen & Saedler, 2001; Weigel & Meyerowitz, 1994).
738 M.F. Riegner / Studies in History and Philosophy of Biological and Biomedical Sciences 44 (2013) 735–744
Fig. 3. Heliconia latispatha showing transition (‘‘metamorphosis’’) between leaf and
bract.
The previous observations need to be extended to yet another
level. As becomes evident through a contemplation of Fig. 2, by
engaging the cultivated imagination, one can quite readily and
objectively envision potential leaves within the gaps of the leaf se-
quence; again, a student can easily sketch a potential leaf any-
where in the sequence. It is quite a cognitive leap, however, to
grasp the lawful nature of ‘‘plantness’’ so that one can pictorially
envision non-existent, yet conceivably possible, whole plants in a
similar way that one can invent undrawn triangles. This ap-
proaches Goethe’s cognitive experience of the Urpflanze, or, in
approximate modern terminology, his experience of the theoretical
morphospace of plants, the ‘‘occupiable’’ morphospace of Arthur
(1997; for definition of theoretical morphospace, see McGhee,
1999, 2007). In this regard, one must take Goethe’s words literally:
With this model and the key to it, it will be possible to go on for-
ever inventing plants and know that their existence is logical;
that is to say, if they do not actually exist, they could, for they
are not the shadow phantoms of vain imagination, but possess
an inner necessity and truth. The same law will be applicable to
all other living organisms. (quoted from Goethe’s Italian Journey
in Brady, 1987, p. 268)
Clearly, more is demanded of our cognitive abilities to reach the
experience of the archetypal Plant than is needed to cognize a miss-
ing leaf in the sequence, but in each case it is the ability to appre-
hend dynamically the precise boundaries and possibilities of an
organic integration that is prerequisite. In this sense, Goethe’s no-
tion of the archetype anticipated Cuvier’s principle of the correla-
tion of parts (presented in 1798), which also acknowledged the
principle of organic integration. But because Cuvier’s organic vision
was grounded in a static typology, it failed to incorporate the pos-
sibility for transmutation (Farber, 1976), that is, for evolution.
3. Between totipotentialism and constraint
The typological thinking implicit in Goethe’s dynamic concep-
tion of the archetype is paradoxical. Regarding the leaf metamor-
phosis example (Fig. 2), on the one hand there exists an infinite
number of potential leaf shapes between any two actual leaves;
this follows as a property of a continuum. In fact, every organism,
through its ontogenetic trajectory, moves through a continuum of
infinite forms. I will call this property totipotentialism (as distinct
from the more narrowly defined concept of totipotency, which re-
fers specifically to the fate of undifferentiated cells during morpho-
genesis). On the other hand, the shapes of possible leaves are
bounded and restricted, not just by the two actual bordering leaves
surrounding each gap in the sequence but also by the unified
movement itself; not any random leaf will fit in the series. This
is constraint. Thus the living organism exists somewhere in the dy-
namic tension between totipotentialism and constraint, which can
also be thought of as the parameters of theoretical morphospace.
Note that, contrary to the standard representation of typology
(e.g., Mayr, 1982; Sober, 1994), Goethe’s dynamic typological think-
ing actually embraces the uniqueness of individual organisms or parts
thereof, and thus the distinction between this form of typology and
population thinking begins to dissolve (see Levit & Meister, 2006).
In fact, it is only through the identification and comparison of vari-
ations on multiple levels of biological organization that Goethe’s
dynamic archetype comes to presence through at least partial dis-
closure. Evidently, striving to understand the ‘‘great malleability of
nature’’ is a central theme in Goethe’s scientific works (Tantillo,
2002, p. 104).
But there’s more. According to Bortoft (2012), to enter more
fully into Goethe’s way of seeing, one must understand what is im-
plied by the act of distinguishing. ‘‘Distinguishing is a dual move-
ment of thinking which goes in opposite directions at once: in
one direction it differences [read as verb], whereas in the other
direction it relates. So the act of distinction ‘differences/relates’—
not differences and relates, because this would be two movements,
whereas there is one movement which is dual’’ (p. 22). Further-
more, our attention is usually drawn to what is distinguished and
consequently the act of distinguishing goes unnoticed, so we over-
look the unity of phenomena and instead focus on the differences
and thus see only separation; this is characteristic of the analytical
mode of thinking. However, if we shift our awareness to the act of
distinguishing, or the ‘‘coming-into-being of distinction’’ (p. 22),
we begin to grasp Goethe’s way of seeing, which is characteristic
of the holistic mode of thinking. This shift of awareness is critical
to understanding the dynamic typological thinking implicit in Goe-
the’s phenomenology, but it is also an elusive and difficult cogni-
tive activity to experience—and, to be fully grasped, it must be
experienced. What is necessary, as Bortoft (2012, p. 27) maintains,
is a shift from thinking ‘‘downstream,’’ from what is already distin-
guished, to ‘‘upstream,’’ to ‘‘the primary act of distinction.’’
Keeping in mind this notion of ‘‘coming-into-being,’’ we return
to a central theme in Goethe’s study of morphology: metamorpho-
sis. In this sense, my earlier description of metamorphosis was
somewhat incomplete but necessary to draw attention to a feature
of Goethe’s thinking that is typically overlooked: that is, the need
to shift thinking from ‘‘downstream,’’ from what is finished, to ‘‘up-
stream,’’ to what is nascent. This is expressed unambiguously by
Bortoft:
The metamorphosis is in the earlier embryonic stage of the
coming-into-being of the organs, and not in the later adult stage
of organs that are already finished. Goethe’s way of thinking is
intrinsically dynamic: it goes back ‘‘upstream’’ into the coming-
into-being of the organs, instead of beginning ‘‘downstream’’
with the organs that are already formed. Metamorphosis is only
to be found in the coming-into-being, and the failure to realize
this leads us to look in the wrong direction by trying to under-
stand metamorphosis in a downstream way. This is the source
of much of the misunderstanding about Goethe’s work. (Bortoft,
2012, p. 66)
In fact, Goethe’s own words point to this distinction between think-
ing upstream, which here is implied by ‘‘reason,’’ and downstream,
here implied by ‘‘practical understanding’’:
Reason is applied to what is developing, practical understand-
ing to what is developed. The former does not ask, What is
the purpose? and the latter does not ask, What is the source?
Reason takes pleasure in development; practical understanding
tries to hold things fast so that it can use them. (quoted in
Miller, 1995, p. 308)
This point, that development is central to Goethe’s view of nature, is
wholly consistent with contemporary evo–devo’s platform that
ontogenies, not adult stages, undergo evolutionary metamorphosis
 M.F. Riegner / Studies in History and Philosophy of Biological and Biomedical Sciences 44 (2013) 735–744
(Amundson, 1998) and that ‘‘[f]orm always has a developmental
origin’’ (West-Eberhard, 2003, p. 201). Moreover, the formulation
of a unified theory of evo–devo itself assumes ‘‘that development
will find a more inclusive place in the study of evolutionary biol-
ogy . . .’’ (Hall & Olson, 2003, p. xv). And finally, the developmental
emphasis of dynamic typological thinking illuminates the historical,
and somewhat contemporary, divide between Darwin—especially
the neo-Darwinians—and the earlier embryologists and morpholo-
gists, poignantly articulated by Amundson (2005): whereas the
morphologists were concerned primarily with the explanation of
the origin of form, Darwin’s goal was the explanation of change, with
little interest in understanding how form arises. With the advent of
evo–devo, this schism is being bridged, and, if his contribution is
understood correctly, it is incumbent upon us to include Goethe
in this dialogue.
4. The nested hierarchical structure of types, convergent
evolution, and deep homology
The notion of the dynamic archetype described above is in need
of still further elaboration. Returning to the earlier example of tri-
angles, one can slightly reduce the specificity that unites the ob-
jects to now embrace the category ‘‘polygon.’’ Consequently,
triangles are now joined by rectangles, trapezoids, hexagons, etc.
into a relatively broader, more inclusive class of geometric con-
structs. Accordingly, the category of triangle becomes nested into
a less specific category, that is, the category of polygon. In contrast,
we could make the triangle category more specified by adding the
requirement of possessing a right angle; consequently, inclusion
would be restricted to only right triangles of which there still exists
a potentially infinite number.
For a biological example, consider cats (family Felidae). If we
were to emulate Goethe’s cognitive experience, but this time direc-
ted toward grasping the ‘‘archetypal Cat’’ in the mind’s eye, after
careful empirical study we would be able to recognize in each of
the 37 living species of felids the lawful integration of organic fea-
tures that constitute the expression of the dynamic type in each
species’ configuration. As disparate as are a tiger, a mountain lion,
and an ocelot, for example, they are but variations on a theme, the
One form expressed in the many (Bortoft, 1996), or the One form
coming-into-being as ‘‘self-differencing’’ (Bortoft, 2012), just as
are the leaves along the stem of an annual plant (Fig. 2). But the
family Felidae is nested within a higher recognized category, that
of the order Carnivora, which itself is nested within the class Mam-
malia, and so on. (As with right triangles mentioned above, we
could also consider a more restrictive category, such as the genus
Panthera, for example.) Thus, archetypes, in a Goethean sense, are
construed as nested hierarchical identities, or as subtypes nested
within types, and it’s the observer’s intentional focus that circum-
scribes the taxonomic level under investigation. The structure of
Goethe’s dynamic type, therefore, meshes seamlessly with the no-
tion of a hierarchical structure of animal (and plant) body plans
central to evo–devo: ‘‘A hierarchical or nested view of Bau-
pläne . . . is . . . both appropriate and essential for any investigation
into the origin of body plans and the systematic organization and
evolution of structures and organisms’’ (Hall, 1996, p. 226).
But there is yet another level of complexity and interrelated-
ness, the expression of which is observed in so-called convergent
evolution. Traditionally, convergence is understood to result when
two unrelated lineages evolve similar adaptations independently in
response to similar selection regimes, for example similar environ-
ments. This scenario presupposes that the same randomly gener-
ated variations must be present coincidentally in each of the two
independent and isolated lineages in order for natural selection
to operate congruently, an assumption that has rarely been vali-
739
dated or even tested. Can such an improbable scenario be the cau-
sal factor behind the striking similarity between, for example, New
World hummingbirds and Old World sunbirds, placental mammals
and Australian marsupials, the black-and-white color pattern of
giant pandas, indri lemurs, and orcas, or the stem-succulent mor-
photype of cacti and euphorbs? From the perspective of dynamic
typological thinking—where no morphotype exists in isolation—
these similarities are not problematic in that just as types and
subtypes are in a sense nested hierarchically, so too can their qual-
itative features be construed to intergrade horizontally, that is,
across diverse phylogenetically unrelated taxa (note that the limi-
tations of language constrain such descriptions to physical terms).
Thus, the black-and-white pattern of the aforementioned mam-
mals has most likely not arisen randomly in each respective
lineage but is an expression of a pattern-generating mechanism
shared among otherwise diverse species. For example, Mundy
et al. (2004) identified a shared genetic mechanism that underlies
a similar plumage pattern found in distantly related birds (e.g.,
Snow Geese and Arctic Skuas), while Riegner (2008), in a broad
study across the entire class of birds (Aves), found repeating plum-
age-pattern elements (e.g., streaks and bars), intercorrelated with
other features, distributed with regularity and predictability across
a wide array of unrelated species (discussed below). And these
shared color-pattern motifs need not be limited to members of
the same vertebrate class. For example, just as complex patterns
of spots and stripes are typical in species of wild cats (e.g., tiger,
jaguar, and ocelot), which are carnivorous, so too are they common
among birds of prey (e.g., hawks, falcons, and owls), which also are
carnivorous. Thus, the expression of these similar pattern elements
in unrelated taxa may, in fact, stem from a homologous pattern-
generating mechanism. Standard homology, however, is insuffi-
cient to explain these shared patterns among very distantly related
taxa but must be extended to a deeper hierarchical level of homol-
ogy, that of ‘‘deep’’ homology (Wake, 2003) or ‘‘underlying’’ homol-
ogy (Rutishauser & Moline, 2005). Of the many remarkable
discoveries of evo–devo in the past few decades, perhaps the most
surprising has been the uncovering of shared developmental path-
ways between vastly different organisms separated by millions of
years of evolution. For example, the formation of eyes in flies and
vertebrates is attributed to a shared developmental pathway
(Carroll, 2005), as is pelvic bone reduction in both stickleback fish
and manatees (Shapiro, Bell, & Kingsley, 2006), while a stunning
recent discovery has demonstrated deep homology of the arthro-
pod central complex (i.e., nervous system) and vertebrate basal
ganglia (Strausfeld & Hirth, 2013). Again, from a Goethean typolog-
ical perspective, these discoveries of profound relatedness among
markedly diverse animals are consistent with the notion of the
One ideal organism—at the most inclusive hierarchical level of the
animal archetype—self-differencing into the multifarious species
that populate the earth.
5. Archetypes and patterns in morphospace
The foundation of a Goethean study of any class of phenomena
requires painstakingly detailed observations followed by morpho-
logical comparisons and the consequent identification of patterns
of interrelationships. Undoubtedly the most extensive application
of a Goethean approach to any class of phenomena is that of evo-
lutionary morphologist Wolfgang Schad who has demonstrated
what can be gained by applying a Goethean-derived dynamic
typology to the class of mammals (Schad, 1977, 2012; see also
Riegner, 1998). Schad’s approach has inspired recent research on
a number of taxa, including rather unusual groups such as dino-
saurs (Lockley, 2008) and early vertebrates (Kümmell, 2011). Here,
taking some cues from Schad’s work, and drawing on a broad study
740 M.F. Riegner / Studies in History and Philosophy of Biological and Biomedical Sciences 44 (2013) 735–744

of avian morphology (Riegner, 2008), I briefly outline how one may together, correlations become evident, which can be integrated
address the question: What is the archetype—or, in other words, into a conceptual model (Fig. 4). Thus, small birds (e.g., passerines,
the parameters of theoretical morphospace—of birds? such as warblers, sparrows) tend to have a countershaded or
Clearly, the world of birds presents a dizzying array of color pat- streaked (i.e., longitudinally striped) plumage, a long tail, and are
terns, sizes, morphologies, ecologies, behaviors, etc. Approached hatched in an altricial condition. Larger birds (e.g., nonpasserines,
from a Goethean perspective, one must entertain the notion that such as ostrich, geese, pheasants), in contrast, tend to have a uni-
each detail holds significance and that all are intercorrelated on formly colored plumage or have bold color patches and/or bars
various levels (i.e., nested hierarchies) of biological organization. (i.e., transverse stripes), a relatively shorter tail and accentuated
In order to establish a context for defining the archetype—i.e., the- anterior (e.g., long neck and/or casque, wattles, dewlap), and are
oretical morphospace—of birds, it is instructive to begin by com- often hatched in a precocial condition. Intermediate-sized birds
paring morphologically widely divergent taxa. Besides occupying (e.g., birds of prey) often display complex pattern of streaks, spots,
the opposite extremes of body size in avian morphospace, ostriches or bars (Riegner, 2008). Moreover, the plumage-pattern trajectory
and hummingbirds in many respects are morphological opposites. is recursive, that is, pattern elements recur in modified configura-
For example, whereas the head is relatively small and the neck dis- tions in distantly related species across the diversity of birds (for
proportionately long in the ostrich, the opposite is the case in the discussion of recursion, see Bird, 2003). For example, the Common
hummingbird (Table 1). A comparison of these avian morphotypes Raven (Corvus corax) is in the passerine taxon but clearly not
highlights Goethe’s principle of compensation or, in contemporary countershaded or streaked but uniformly dark as is typical for large
terminology, developmental trade-offs, both within and between species. However, this species is among the largest of passerines, so
organisms, as noted in his observation that ‘‘the neck and extrem- in that taxonomic context the recurrence of uniform coloration is
ities are favored in the giraffe at the expense of the body, but the consistent with the overall pattern. Similarly, plovers are generally
reverse is the case in the mole’’ (quoted in Miller, 1995, p. 121). small-bodied birds, but within this taxon the smallest species (e.g.,
In other words, an organism cannot be the best of all possible genus Charadrius, Ringed and Semipalmated Plovers) are typically
worlds: an exaggeration in one feature can arise only because an- countershaded, occasionally with broken bands below, while the
other is de-emphasized. For example, the loss of hind limbs in larger species (e.g., genus Pluvialis, European and American Gold-
whales has presumably permitted the remarkable over-develop- en-Plovers) exhibit solid black undersides in the breeding plumage,
ment and lengthening of the jaws (which, in a sense, are the that is, reverse countershading in relatively robust birds, which
‘‘limbs’’ of the head). In contrast, the huge hypermorphic hind again is a recurrent plumage pattern typical of the largest of birds
limbs of T. rex were only permissible due to the stunted, vestigial within a given taxonomic context (see Fig. 4).
forelimbs of this Cretaceous dinosaur (a similar morphological pat- Clearly, the model depicted in Fig. 4 does not define the com-
tern is seen in the modern-day kangaroo mice and rats, family Het- plete multidimensional theoretical morphospace of birds, and it
eromyidae). On a physiological level, a comparison of relative brain certainly does not represent an archetype in the Goethean sense
size and gut length in howler monkeys—which eat mainly difficult-
to-digest but easily accessed leaves—and in spider monkeys—
which eat easy-to-digest but more difficult to acquire fruits—indi-
cates that the former has a relatively smaller brain but larger gut
than the latter (Allman, 2000). Emlen (2000) has identified a devel-
opmental trade-off (i.e., an inverse relationship) between the
length of ‘‘horns’’ and the size of compound eyes in Onthophagus
dung beetles, and many more examples have been documented
in other animals as well as in plants (see, for example, West-Eber-
hard, 2003, especially chap. 16). Accordingly, by considering Goe-
the’s principle of compensation, the limits and possibilities of
potential morphologies can be grasped objectively through careful
study of any class of biological phenomena. These considerations,
in turn, point to complex patterns of compensatory developmental
trajectories or, in other words, dissociated heterochronies (McNa-
mara, 1997), a discussion of which would fall outside the scope
of this article.
In order to continue articulating an outline of the theoretical
morphospace of birds, it is necessary to examine various character-
istics and search for correlated trajectories; in other words, it needs
to be derived empirically. For example, Riegner (2008) considered
plumage pattern, body size, morphological accentuation (i.e., ante-
rior—posterior), and developmental mode (i.e., altricial—precocial)
across a wide array of birds. When these features are considered

Table 1
Morphological comparisons between ostrich and hummingbird.

Relative sizes Ostrich Hummingbird

Head: body size Small Large
Bill: head size Short Long
Neck: trunk length Long Short Fig. 4. Approximation of avian morphospace. The trajectory of plumage patterns is
Trunk: body length Short Long correlated with various morphological features and is recursive at finer levels of
Wing: body length Short Long taxonomic comparison. Plumage patterns are, from left to right: countershaded;
Leg: body length Long Short streaked; spotted, drab, or blended; barred; bold separation of black and white or
Foot: leg length Short Long uniform dark; and reverse countershaded. Figure reproduced from Riegner (2008)
and used with permission.
 M.F. Riegner / Studies in History and Philosophy of Biological and Biomedical Sciences 44 (2013) 735–744
(which, recall, can never be visually represented), but it does
identify the dynamic interrelationships of selected features that
are grounded in empirical observation and, taken together, can
subsequently serve as a conceptual guide to make sense of other-
wise diverse, and mistakenly random, phenomena. Moreover,
based on just a few given characteristics, the model can be used
to predict what a possible species may look like or, at the very
least, what forms are not permissible in avian morphospace, such
as an adult duck the size of a finch or vice versa. And finally, it is
notable that the color pattern—body size relationship seen in birds
is also apparent in other animal groups, such as mammals. For
example, small mammals (e.g., rodents, such as deer mice) tend
to be countershaded, large species (e.g., hoofed mammals, such
as bison) are often uniformly colored, and intermediate-sized
mammals (e.g., carnivores, especially cats) display complex pat-
terns of spots and/or stripes (Riegner, 1998; Schad, 1977, 2012).
Furthermore, when rodents have stripes they are aligned horizon-
tally (e.g., thirteen-lined ground squirrel), and when hoofed mam-
mals have stripes they are aligned vertically (e.g., zebras). Among
cats, in smaller species (e.g., ocelot) the striping pattern is horizon-
tal (as in rodents), while in larger species (e.g., tiger) the stripes are
vertical (as in hoofed mammals); intermediate-sized cats (e.g.,
leopard) display a spotted pattern that is neither longitudinally
nor transversely orientated. Thus, while Lewens (2012) argues
against the reliance on essentialism to explain the ‘‘stripyness’’ of
tigers, from a Goethean perspective the more interesting, and
revealing, questions are: why do tigers specifically have vertical
stripes, or why do mountain lions altogether lack stripes? These
questions can be addressed when one grasps the dynamic nature
of the mammalian type and associated subtypes (Riegner, 1998;
Schad 1977, 2012).
6. Theoretical morphospace and recursion
Whether one observes the changing leaf shapes in a graded
series, the morphological variations among wild cat species, or
the different kinds of teeth in the mammalian jaw, dynamic typo-
logical thinking shows that each grouping discloses a single ideal
form in multiple guises, that is, ‘‘multiplicity in unity’’ (the epis-
temological inversion of ‘‘unity in multiplicity;’’ see Bortoft,
1996). From a traditional perspective, this is the notion of homol-
ogy, including serial or ‘‘iterative’’ homology (Roth, 1991). Dy-
namic typological thinking, however, reaches further in that the
variation among these manifestations is not considered random
or accidental but obeys a strict, inherent lawfulness. The effort
to discover these ‘‘inherent laws of form’’ is what lies at the heart
of Goethe’s science of morphology and his notion of the dynamic
archetype.
If practiced diligently, dynamic typological thinking can answer
the call to develop a theoretical morphology in which ‘‘[t]he ulti-
mate triumph. . .would be an understanding of biological diversity,
framed in terms of the boundaries between the possible and the
impossible. It should integrate across all levels of structure, from
organic molecules to entire and seemingly complex functioning
organisms, where as yet undiscovered laws of structural conso-
nance may exist’’ (Hickman, 1993, p. 170). In other words, ques-
tions can be addressed that seek to understand ‘‘the significance
of the spatial distribution and density of forms within morpho-
space’’ (McGhee, 2001, p. 172) based on the distribution of obser-
vable morphotypes (Gould, 2002) or ‘‘occupied’’ morphospace
(Arthur, 1997). As discussed earlier, Goethe’s Urpflanze can be con-
strued as defining the theoretical morphospace of plants, not as an
abstraction but as an objective cognitive experience. Thus, if rightly
understood, the thinking implicit in Goethe’s notion of the arche-
type can be applied to any class of biological phenomena to invite
741
a more thorough apprehension of the organizing principle, the dy-
namic activity that unites the disparate parts.
The varying but related forms of, for example, sequential leaves
up a stem or vertebrae along a spinal column, or the set of primary
feathers of a bird’s wing, display recursive properties in that the
morphological elements reiterate in modified configurations both
within a given organism and between species. Indeed, the evolu-
tionary process itself seems to be recursive, generating fractal-like
variation, a characteristic of well-ordered chaotic systems (Bird,
2003). Apparently, even our thinking activity exhibits recursive
properties (Corballis, 2007). In addition, gene regulatory networks
are described not only as structured hierarchically (Erwin & David-
son, 2009; Schoch, 2010) but as highly recursive (Davidson & Er-
win, 2006). Accordingly, the principles and applications of chaos
theory and fractal geometry (e.g., Barnsley, 2006) evidently inter-
sect those of dynamic typology, and there is presumably much to
learn here. Dynamic typological thinking, in fact, is well suited to
trace the unifying thread that weaves through the fractal variation
encountered among natural phenomena.
Chaotic, fractal, and recursive properties are evident not only
among serially homologous structures within an organism (e.g.,
leaf sequence) or in comparisons of morphological elements
among unrelated organisms (e.g., avian plumage-pattern ele-
ments) but also by noting variations in phenotypic expression of
a given organism under varying environmental contexts. This latter
phenomenon is addressed by epigenetics (Jablonka & Lamb, 2005)
and, more specifically, by phenotypic plasticity (Pigliucci & Pres-
ton, 2004). Yet again, over 200 years ago Goethe’s astute observa-
tions described these phenomena: ‘‘. . . a plant growing in low-
lying, damp spots will . . . develop smoother and less refined leaves
than it will when transplanted to higher areas, where it will pro-
duce rough, hairy, more finely detailed leaves’’ (Goethe, 1790, p.
19). These observations revisit the notion of the dynamic relation-
ship between totipotentialism and constraint. For example, it is
clear that, when an acorn germinates, an oak—and nothing else—
will subsequently grow; that is constraint. But the particular form
a given oak takes, from a potentially infinite number of possibili-
ties, will depend on how the environmental conditions—soil chem-
istry, slope, sun exposure, wind, snow loading, animal damage,
etc.—influence the expression of the genotype, the outcome of
which cannot be precisely predicted. Similarly, we know that a ti-
ger will have stripes, but the exact distribution of those stripes
across the tiger’s body, like our fingerprints, is indeterminate, most
likely due to immeasurable developmental perturbations of initial
conditions.
7. Is the idea of the archetype a mental abstraction, an actual
organizing principle, or neither?
Adhering to Goethe’s experience of dynamic typological think-
ing as described above, it is a worthwhile exercise to ask whether
the archetype, as so construed, is merely a mental abstraction
added to the phenomena, or whether it has any claim to reality
drawn out of the phenomena independent of a human mind to
apprehend its characteristics. In other words, is it an organizing
principle that plays a role in morphogenesis, development, and
even organic evolution?
This, evidently, is an ontological question, which I will not pre-
sume to answer except by exploring what I think would have
approximated Goethe’s interpretation. First, recalling the leaf
metamorphosis example above (Fig. 2), what exists between the
sensibly perceived elements—that is, between the leaves—what
moves between them, is as crucial to Goethean phenomenology
as the elements themselves. In other words, the complete plant
phenomenon includes not only all the morphological structures
742 M.F. Riegner / Studies in History and Philosophy of Biological and Biomedical Sciences 44 (2013) 735–744
but also the dynamic movement—that is, the set of objective, rela-
tional ideas—which links together each of the separate parts (note
that the parts only appear as separate in a spatial dimension).
Furthermore, as I hope to have demonstrated, the dynamic move-
ment of the developing plant, its coming-into-presence, is not
merely a subjective mental representation, nor an abstract gener-
alization, but an objective cognitive experience based on the tangi-
ble existence of the actual leaves and floral morphology. We
apprehend the dynamic Idea of the plant by delving into the details
of its parts and thereby accessing the intensive dimension of the
phenomenon (Bortoft, 1996, 1998). Accordingly, Goethe may have
proposed that this dynamic cognitive activity is in resonance with
the transformative principle itself, which shapes the developing
organism and ‘‘moves it’’ through its various ontogenetic stages.
As Bortoft (1996, pp. 240–241) explains, ‘‘The organizing principle
of the phenomenon itself, which is its intrinsic necessity, comes
into expression in the activity of thinking when this consists in try-
ing to think the phenomenon concretely. What is experienced is
not a representation of the organizing principle, a copy of it ‘in
the mind,’ but the organizing principle itself acting in thinking.’’
In this regard, Goethe (1790) referred to ‘‘sap’’ as becoming
more purified as it rises through the plant, and the image of this
is observed in the sequential refinement of the leaves and then flo-
ral parts traced up the stem. But here again, as in his designation of
‘‘leaf’’ as the unifying principle behind metamorphosis, ‘‘sap’’ may
have been his expression for the dynamic biological activity that
continually shapes and reshapes matter in a living organism. This
is not a vitalistic notion, as some have mistakenly asserted, but
points to an organizing principle, which to this day remains elusive
but central to the biological sciences. What, for example, drives the
processes of growth and differentiation in the embryo? Why is the
eight-cell stage of development not ‘‘content’’ to remain as such
but continues dividing, growing, and differentiating? These ques-
tions reside at the frontier of our understanding of morphogenesis,
and, I maintain, is what Goethe’s dynamic typology attempts to
explore.
8. Is dynamic typological thinking compatible with traditional
principles of evolution?
Regarding descent with modification, although Goethe’s arche-
type does not necessarily imply ancestral forms, it does not ex-
clude them. After all, there’s no reason why a temporal series of
fossil forms cannot be arranged sequentially just as individual leaf
shapes in the graded series and, through interpolation, lend them-
selves to a determination of what the missing—i.e., potential—
forms may have been. Indeed, chapter nine of D’Arcy Thompson’s
classic text, On Growth and Form (1961), demonstrates how this
is undertaken through mathematical applications, which, however,
should not be confused with engaging the faculty of dynamic cog-
nition described here. Thompson’s analytical methods remain
external to the phenomena whereas a cognitive apprehension of
the type internalizes the phenomena; superficially, the results
may appear similar or even identical, but the two procedures have
different implications for our understanding of evolutionary
dynamics.
Additionally, dynamic typological thinking does not reject the
central tenets of neo-Darwinism, that is, random mutation and
natural selection. However, while the premise of non-directional
variation is only weakly supported by the view expressed here,
natural selection remains intact, though shifted to a level of sec-
ondary importance. In other words, ‘‘the variation presented to
selection is nonrandom’’ (Raff, 1996, p. 428) and thus results in
‘‘developmental bias’’ (Arthur, 2004), which would ultimately ‘‘bias
a lineage’s access to morphospace’’ (Sterelny, 2000, p. S385). Con-
sequently, natural selection should be considered not creative but
only eliminative. Thus, dynamic typological thinking (and evo–
devo) does not deny random mutation and natural selection but
does not view organic evolution as fully, or even satisfactorily, ex-
plained by these mechanisms.
The evidence for gene regulatory networks lends further sup-
port to Goethe’s notion of the dynamic archetype. In fact, the par-
allels between the two are remarkable. As seen in the earlier
example of the leaf sequence (Fig. 2) in which the intervals be-
tween the leaves are crucial, it has become apparent that, although
the genes themselves influence a given phenotype, it is the net-
work of interactions between the genes, the relationships between
and among them, that ultimately translates into the phenotype
(Davidson & Erwin, 2006). This discovery moves well beyond plei-
otropy and epistasis, and is clearly a paradigm shift that is begin-
ning to alter the way we understand gene expression and the
origin of biological form. Whether this shift in understanding will
ultimately lean closer to the ideas discussed here is an open
question.
And finally, it is notable that recent publications by philoso-
phers of science have revisited typology and essentialism and have
not only identified flaws within the ‘‘essentialism story’’ (Amund-
son, 2005; Winsor, 2006) or the ‘‘received view’’ (Wilkins, 2009)
promulgated by a generation of evolutionary biologists, but also
have proposed convincing arguments for reconsidering and even
‘‘resurrecting’’ at least some form of biological essentialism (e.g.,
Devitt, 2008; Lewens, 2009b; Love, 2009; Okasha, 2002; Walsh,
2006; Wilkins, 2010). Note, however, that these proposals admit-
tedly differ somewhat from those made in this paper and that none
considers the dynamic typology implicit in Goethe’s way of under-
standing nature.
9. Setting the record straight
Historically, uncritical thinking has amalgamated what I have
called static and dynamic typological thinking. Consequently, Goe-
the, and most likely others (even Plato; see Bortoft, 2012), was mis-
takenly accused of promoting a world view that has no place in
modern evolutionary biology. Moreover, the notion that ideas
inhere in nature—a central position of idealist morphology
(Richards, 2002)—has been rejected with an almost knee-jerk reac-
tion by some vocal neo-Darwinians because, from their perspec-
tive, ideas have no inherent reality. But if that were the case, it
follows that, because their criticism is itself an idea, it too would
need to be discounted. Evidently, we cannot escape from ideation,
and thus the idealist morphologists were perhaps on to something.
Richards (2002, p. 192) summarizes this eloquently: ‘‘It hardly
seems easier to believe the world is really a ball of mathematical
strings that reveals itself to our consciousness as natural objects
of ordinary experience than to believe it is an organic structure
of ideas that reveals itself in comparable fashion. Idealism cannot
be defeated, only forgotten.’’
There is a relatively recent and explicit literature that points to
the unfair assessment of Goethe’s way of science, but it has lan-
guished below the radar of evolutionary biologists. Most notable
are the epistemological contributions by philosopher of science
Brady (1987, 1998) and physicist Bortoft (1996, 1998, 2012). In
addition, there are numerous studies that either apply Goethean
methodology to a plethora of natural phenomena or explore its
epistemology: besides Schad (1977, 2012), these include Amrine
et al. (1987), Bockemühl (1998), Brook (1998), Seamon and Zajonc
(1998), Lockley (1999, 2008), Steigerwald (2002), Verhulst (2003),
Dornelas and Dornelas (2005), Ebach (2005), Holdrege (2005,
2013), Holdrege and Talbott (2008), Steiner (2008), Williams and
Ebach (2008; especially chap. 3 and epilogue), Skaftnesmo
(2009), and Suchantke (2009).
 M.F. Riegner / Studies in History and Philosophy of Biological and Biomedical Sciences 44 (2013) 735–744
Whether or not one endorses the epistemological approach im-
plicit in dynamic typology, as exemplified in a Goethean way of
science, is contingent upon one’s understanding of this methodol-
ogy. However, if one criticizes and offhandedly rejects such an ap-
proach, one must at least be aware of what it is that one is
criticizing and ultimately discarding. As I’ve tried to show, this
has not been the case historically due to the confounding of static
and dynamic typological thinking. As mentioned previously, this
confusion stems from the overlooked requirement to engage our
cognitive activity in a way in which we are unaccustomed. Such
an engagement requires more than a conceptual change or concep-
tual reorganization (sensu Chi & Roscoe, 2002) and even more than
a paradigm shift (sensu Kuhn, 1970), in which only the content of
thinking changes. Rather, it necessitates a conscious transformation
of the cognitive activity itself, and therefore a transformation of the
scientist him- or herself. This is the piece that has perennially
eluded critics of Goethe’s way of science and the dynamic typolog-
ical thinking it exemplifies. As Brady (1998, p. 84) points out, ‘‘The
writers of scientific histories are unable to perceive what they can-
not conceive.’’ Interestingly, Goethe himself alluded to this pitfall:
There is a delicate empiricism which makes itself utterly iden-
tical with the object, thereby becoming true theory. But this
enhancement of our mental powers belongs to a highly evolved
age. (quoted in Miller, 1995, p. 307)
As I’ve attempted to show, the confluence of modern evolutionary
developmental biology with Goethe’s dynamic typology holds
promise in deepening our understanding of the origin, diversity,
and limitations of biological form and how those forms manifest
and change over time. In this sense, Goethe should be seen not sim-
ply as a forerunner of contemporary evolutionary developmental
biology but as providing a narrative that can contribute to the fur-
ther development of a new archetypal biology.
Acknowledgments
I am grateful to the late Ron Brady for many hours of engaging
conversations in his office at Ramapo College in 1987–88; I dedi-
cate this article to his memory. I also thank Malte Ebach, Craig
Holdrege, Trond Skaftnesmo, and John Wilkins for suggestions that
improved an earlier draft of the manuscript, and Greg Radick and
two anonymous reviewers for very helpful comments that guided
me in reshaping a more recent draft; of course, any misinterpreta-
tions rest on my own shoulders. Prescott College deserves my grat-
itude for awarding me a sabbatical leave in spring 2010, which
afforded me the time to gather my thoughts on this subject. And
finally, I thank the many students over the years in my class Form
and Pattern in Nature for helping me refine my own understanding
of the ideas expressed herein.
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