Teoria Evol PDF

Department of Biological Sciences, Columbia University, New York, NY, USA
Explanations in evolutionary theory

1
W. J. Bock
Abstract
The theory of biological evolution is dened in many ways, leading to considerable confusion in its application and testing against objective
empirical observations. Evolutionary change is usually dened as genetic which would exclude both cultural and template evolution; hence the
qualifying adjective genetic should not be included in the denition of biological evolution. Darwins theory, always described by him in the
singular, is actually a bundle of ve independent theories about evolution as advocated by Mayr. Furthermore only one of these theories, that of
common descent, is historical, and the other four evolution as such, gradualism, processes of phyletic evolution and of speciation, and causes of
evolution are nomological. Hence not all evolutionary theory is historical. Biological comparisons can be divided into horizontal and vertical
ones and valid conclusions from one type of comparisons cannot be automatically extrapolated to the other. All phyletic evolutionary change, no
matter how extensive it may be, never crosses species taxa boundaries; hence it is not possible to distinguish trans-specic evolution ( evolution
beyond or above the level of the species) from evolution within the species level. Macroevolution does not dier from microevolution except in the
scale of the overall change; no special causes or processes of macroevolution exist.
Key words: Denitions of evolution Darwins ve evolutionary theories nomological and historical theories of evolution nomological-
deductive and historical-narrative explanations functional explanations horizontal versus vertical comparisons in biology species limits
macroevolution
Introduction
Countless people have followed the recent debates and trial in
Harrisburg, Pennsylvania, USA on the teaching of biological
evolution versus intelligent design in public schools in the
United States as well as the earlier one in the state of Arkansas,
USA on teaching biological evolution versus scientic crea-
tionism. In both cases the courts declared against the teaching
of intelligent design and scientic creationism as being
religious, not scientic, and hence violating the separation of
church and state in the United States Constitution. Most
interesting in these two trials was the spectrum of meanings
given to the Theory of (Biological) Evolution which varied so
much that one could be surprised that any decisions could be
reached at all in these trials. Most likely these court decisions
were reached, not on the basis of understanding what is meant
by biological evolutionary theory, but on a conclusion that
intelligent design and scientic creationism are not scientic
theories (Brockman 2006).
A major foundation for this diversity in the meanings of the
theory of evolution goes right back to 1859 and the publication
of Darwins On the Origin of Species, namely that:
(1) Darwin always referred to his ideas as my theory in the
singular;
(2) the impression that all aspects of evolutionary theory are
historical;
(3) the failure to distinguish between horizontal and vertical
comparisons in biology.
Over the decades as more and more was learned about
evolution, many evolutionists stated that Biological Evolution
was no longer a theory, but was factual or a fact ( an
objective empirical observation) which confused the issue even
more as a sharp dierence exists between scientic theories and
objective empirical observations. In most of the statements
that evolution is a scientic fact, the author actually meant
that historical evolutionary theory ( the general notion that
living organisms descended with modications from a com-
mon ancestor) is so exceedingly well tested ( well corrobor-
ated, Popper 1959; 1968: 3234) that it can for all intense
purposes be accepted as factual. However, it is still better to
state that historical evolutionary theory is an exceedingly well
corroborated theory and that massive counter tests supported
by strong empirical objective observations are needed to
disprove it. Facts, as used in science, are quite dierent from
theories and the two are best kept strictly separated.
The major themes to be addressed in this essay have been
nicely summarized a quarter-century ago by Mayr (1982: 8) in
the introductory chapter of his The Growth of Biological
Thought where he wrote: As a consequence, some exceedingly
confused accounts of the history of biology have been
published by authors who did not understand that there are
two biologies, that of functional and that of evolutionary
causations. Similarly, anyone who writes about Darwins
theory of evolution in the singular, without segregating the
theories of gradual evolution, common descent, speciation,
and the mechanism of natural selection, will be quite unable to
discuss the subject competently. Today the problems are
much the same as when Mayr penned this passage in the late
1970s with the addition that most scholars do not comprehend
the distinction between nomological-deductive (N-DE) and
historical-narrative explanations (H-NEs) and hence that
major dierences exist between nomological and historical
theories of evolution. Mayr hinted at this last diculty when
he wrote (Mayr 1982: 27): Every evolutionist who has had a
discussion with lay people has been asked: Has evolution been
proven? or How do you prove that man descended from
apes? But he did not emphasize that these represent quite
dierent scientic questions and require dierent modes of
explanation, the rst being nomological-deductive and the
second historical-narrative. Later in this introductory chapter,
Mayr argued (pp. 7176) for the importance of historical
narratives and for the need of a philosophy of biology separate
1
It is with great pleasure that I dedicate this paper to my teacher,
mentor and friend, Ernst Mayr (19042005) whose writings on evo-
lutionary biology and the philosophy of biology formed the founda-
tion for my thinking on the ideas presented herein.
2007 The Author
Journal compilation 2007 Blackwell Verlag, Berlin
Accepted on 10 December 2006
J Zool Syst Evol Res doi: 10.1111/j.1439-0469.2007.00412.x
J Zool Syst Evol Res (2007) 45(2), 89103
from that of the physical sciences. Yet he never pushed these
ideas to the logical conclusion that a distinction must be made
between nomological and historical theories of evolution, most
likely because of his strong aversion to law-like statements in
science and hence to nomological explanations. Possibly Mayr
equated functional explanations with nomological-deductive
and evolutionary explanations with historical-narrative which
is largely true although this relationship does not hold quite so
simply (Bock 2004a). Or possibly because Mayr did very little
work in functional biology, he was not suciently versed in the
nuances of nomological-deductive explanations (N-DEs) and
overlooked their role in evolutionary biology. Or possibly, he
restricted his view of biology to evolutionary explanations and
considered all other aspects of biological study ( strictly
functional explanations) as physical sciences.
Because Darwin always referred to his ideas as my theory
(Mayr 1982: 8; 1985: 757) always in the singular almost all
workers accepted the existence of only single theory of
biological evolution which was regarded as being strictly
historical. Most specialists and laity alike have been and still
are mainly interested in the historical evolutionary theory, that
is, in the history of life, or in modern terms The Tree of Life.
This is nicely shown by the history of evolutionary studies in
which biologists accepted rapidly the theory of common
descent ( historical evolutionary theory), but ignored or
rejected Darwins theory of natural selection as well as
gradualism and speciation ( nomological historical theory;
see Mayr 1985). Many biologists, then as well as now, do not
care about nomological theories of evolution and the relation-
ship between them and historical evolutionary theory. And if
both nomological ( process) and historical ( pattern)
evolutionary theories are mentioned, emphasis is always
placed on the latter with little consideration on how the two
are connected. Even Popper (1977) focused so strongly on the
historical aspect of evolutionary theory that he concluded that
all evolutionary biology is historical and hence according to his
approach to science, evolutionary theory is not scientic (see
Hull 1999, for a detailed analysis). Caplan (1977, 1978, 1979),
on the other hand, provided a convincing argument that
[nomological] evolutionary theory is not circular and is
deductive. A close examination of his papers reveals that
Caplans position applies only to some of the set of Darwins
theories about organic evolution (Mayr 1985) and in particular
to the one dealing with evolutionary mechanisms (theory D,
see below). Caplans analysis denitely does not apply to the
last of these theories (theory E, or common descent) which is
the aspect of evolutionary theory foremost in most peoples
mind when they use the term Darwinism or evolutionary
theory. Futumya in his text books (1998:1112; 2005:1115)
implies that a distinction exists between nomological and
historical evolutionary theories, but does not develop these
ideas fully.
In this essay four major points have been addressed, namely:
(1) Examining the eect of horizontal versus vertical compar-
isons in biology on evolutionary concepts; (2) providing a
denition of evolution which will serve as the foundation on
which to examine the scope of evolutionary theories and to
show which of these theories are nomological and which are
historical; (3) arguing that Darwin proposed a set of at least
ve dierent evolutionary theories, not just one, as advocated
by Mayr (1985) which can still be used today to characterize
the major, independent evolutionary theories; and (4) showing
which of these evolutionary theories are nomological and
which are historical, following Bock (2004a), and what is the
relationship between the two types of evolutionary theory.
Horizontal and vertical comparisons
Before proceeding into the main discussion on evolutionary
theory, the consequences of horizontal and vertical compar-
isons in biology (Bock 1989a) on evolutionary thinking have to
be considered. In the middle of the 19th century almost all
biologists believed that only single type of comparisons
existed that is, all comparisons between all organisms are
the same because at that time no general concept had yet been
advocated of organisms changing over historical time. This
belief was excusable in early Darwinian thinking but not
thereafter, although most biologists still accept today that a
single type of comparison exists and that conclusions can be
readily extrapolated from any comparison to another.
It must be stressed that the idea of horizontal versus vertical
comparisons is absolutely dierent from the concepts of
horizontal versus vertical evolution. The latter set of concepts
implies an absolute dierence in the time needed for each type
of evolutionary change, but these terms refer to speciation (or
splitting of phyletic lineages horizontal) versus phyletic
evolution ( vertical); both types of evolutionary change
require time, albeit dierent amounts. Use of the terms
horizontal versus vertical evolution is misleading to the extent
of being wrong and is best not used.
Horizontal comparisons are those between individual organ-
isms of the same species taxon or between members of dierent
species taxa ( members of dierent phyletic lineages) whether
or not the organisms exist at the same time level. Most
biological comparisons are horizontal. In horizontal compar-
isons, not only can homologous features be compared but also
nonhomologous ones depending on the nature of the compar-
ison. Dierences observed in horizontal comparisons can never
be evolutionary modications because they do not represent
changes between dierent stages within a phyletic lineage (see
Bock 1979, 1986, 1995, 2004b, for the distinction between the
species concept and the phyletic lineage concept, and between
the species concept, the species category, and the species taxa.).
Because, interspecic horizontal dierences are between dif-
ferent species between organisms of dierent phyletic
lineages great care must be used when extrapolating results
of horizontal comparisons to represent phyletic evolutionary
changes (Bock 1979).
Vertical comparisons are those between members of the
same phyletic lineage existing at dierent times, that is,
between ancestral and descendent groups. Because vertical
comparisons are between organisms in dierent time slices of
the same phyletic lineage, these comparisons are never
between dierent species. The features being compared
vertically are generally homologous ones, although it might
be possible to have useful vertical comparisons between
nonhomologous features. Dierences noted between homol-
ogous features in vertical comparisons represent evolutionary
modications. Empirical vertical comparisons are dicult to
make because one generally does not know whether the
organisms being compared are members of the same phyletic
lineage even when dealing with fossils. Hence such compar-
isons are largely hypotheses and are indicative of whether the
organisms are believed to be members of the same phyletic
lineage at dierent points in time which are almost always
theoretical.
90 Bock
2007 The Author J Zool Syst Evol Res (2007) 45(2), 89103
Horizontal species boundaries are between species taxa
existing at the same time and same place (synchronous and
sympatric) and represent the barriers to gene ow between
species; these boundaries are considered to be unbridgeable,
although in reality they may be somewhat leaky but not to the
extent of resulting in denite gene ow between the two
species.
Vertical species boundaries are postulated to be similar to
horizontal boundaries between species taxa along the same
phyletic lineage, but their exact nature has always been left
rather vague. Most workers advocating vertical species
boundaries have assumed that they are the same as the
unbridgeable horizontal boundaries ( barriers to gene ow),
and hence special evolutionary processes are required for this
trans-specic evolutionary change which are dierent than
those existing for evolution within the species limits. Yet if
phyletic evolutionary change is gradual, then such vertical
species boundaries cannot exist, and that there is no dierence
between evolution within the species limits and evolution
beyond the limits of the species ( trans-specic evolution).
Hence, horizontal comparisons are between members of the
same species taxon or between members of dierent species
taxa ( dierent phyletic lineages), while vertical comparisons
are between members of the same phyletic lineage at dierent
times. Valid conclusions reached on the basis of a horizontal
comparison cannot automatically be extrapolated to a vertical
comparison and vice versa. In the following discussion about
species, I follow the biological species concept (Mayr 1942,
1963) and not the evolutionary or the phylogenetic species
concepts (see Wa gele 2000 for a good modern discussion of
these concepts). The biological species concept is character-
ized, quite correctly by Mayr as non-dimensional, as distin-
guished from the multi-dimensional species taxon. The
evolutionary and phylogenetic species concepts do not distin-
guish between the species concept and the phyletic lineage
concept, causing confusion. Most philosophers of biology do
not make a distinction between the species concept and the
phyletic lineage concept, often leading to confusion in their
analyses. Furthermore, distinctions must be made between the
species concept, the species category and the species taxon
(Mayr 1963; Bock 1995, 2004b), to avoid the ambiguity arising
when using only the term species.
For Darwin and other biologists of his time, dierent species
taxa of organisms were considered to be separated from one
another by unbridgeable gaps a horizontal comparison. That
is, denite boundaries or limits existed around each species
taxon. A major problem facing Darwin and other early
evolutionists was how this boundary can be overcome in the
evolution of one species taxon to another. The concept of the
horizontal barrier between species taxa had been extended
automatically to the idea of a vertical boundary separating an
ancestral species taxon from a descendent one a vertical
comparison. This is what Darwin had in mind when he
concluded that with sucient (phyletic) evolutionary change, a
new species taxon would arise. Or to put this another way, a
foremost question for biologists at that time was how much
evolutionary change was necessary before a variety of a species
taxon (a particular breed, such as the breeds of the domesti-
cated dog, or a subspecies or geographic race), reached the
status of a new species taxon this question still exists for
many workers. Darwin failed to realize that two distinct
evolutionary processes existed phyletic evolution and speci-
ation and concentrated on phyletic evolution which led to his
famous disagreement with Moritz Wagner (see Mayr 1982:
562566). Wagner had stressed the role of geographic isolation
in evolution and especially in the origin ( multiplication) of
new species taxa. It is impossible to be precise on how most
evolutionists (including Darwin and Wagner) viewed the origin
of new species taxa until the period of the Evolutionary
Synthesis (19371948), because of vagueness in stated posi-
tions on both sides.
Yet it was clear well before 1859 biologists realized sharp
and denite boundaries existed between sympatric and syn-
chronous species taxa. In modern terms, these horizontal,
unbridgeable boundaries around species taxa are formed by
intrinsic genetic isolating mechanisms (Bock 1995, 2004b),
preventing the ow of genetic information from species to
species. With the acceptance of evolutionary ideas and the
realization that species taxa had a history, biologists automat-
ically extended this horizontal boundary around species taxa
to a vertical boundary between species in the same phyletic
lineage. This extension was to be expected at that time because
Darwin and most other biologists did not appreciate the
distinction between the species and varieties (including sub-
species or geographical races) within species. By 1860 concepts
about geographic races were developing in the United States
and Imperial Russia (Haer 1986, 1992, 1994), mainly in
ornithology, but these ideas were rejected by most systematists
and evolutionists in the United Kingdom until after the end of
the nineteenth century. Further Darwin and most other
evolutionists did not understand the role of external barriers
in the speciation process and believed that new species arose by
continuous phyletic evolution.
Although in 1859 biologists automatically extended this
horizontal boundary around species taxa to a vertical bound-
ary between ancestral and descendent species, no factual
evidence exists for this vertical boundary. Nor did any need
exist to continue the idea of vertical species taxa boundaries as
evolutionary theory became better understood. A major source
of this problem came from the exceedingly clever and catchy
title of Darwins book On the Origin of Species which stuck in
the minds of biologists, and was used in much later books such
as Dobzhanskys 1937 Genetics and the Origin of Species and
Mayrs 1942 Systematics and the Origin of Species. These
latter authors borrowed the title of Darwins book to claim
proper kinship with it, but the phrase the origin of species in
these later titles should be read as evolution. Yet titles, e.g.,
Systematics and Evolution, would not have been as attractive
as Systematics and the Origin of Species. And it is most
dicult to think of a better title than the one chosen by
Darwin for his 1859 volume although species taxa do not
originate as implied in the title of his book and the text.
The concept of a vertical boundary as well as a horizontal
boundary between species taxa remains until today in the
thinking of almost all evolutionists and lay-persons. Until the
middle of the twentieth century, this notion still implied, at
least for some evolutionists (e.g., Schindewolf 1950; Herre
1951), that a distinct jump existed a macroevolutionary
change from one species taxon to a descendent one, the
so-called evolution beyond the level of a species or trans-
specic evolution. And they argued that a special, and usually
unknown or mysterious, macroevolutionary mechanism exists
that cannot be reduced to the known mechanisms of micro-
evolutionary change evolution within the bounds of the
species taxon (but see Bock 1979 for a reductionistic
approach). Even after the period of the evolutionary synthesis,
Explanations in evolutionary theory 91
some books on evolutionary theory still used the term trans-
specic evolution or evolution above the species level in their
title (Rensch 1947), or as chapter or section heading, although
without necessarily implying any special meaning to this term.
Most anti-evolutionists are mainly concerned with macro-
evolution which they consider being change from an ancestral
species taxon to a descendent one and which violates their
belief that all kinds of organisms were created independently
from one another. Most have correspondently little interest in
microevolution which is regarded as small modications
within the limits of a species taxon, but never exceeding these
limits. Some anti-evolutionists accept the numerous factual
examples of change within species taxa, such as the develop-
ment of tolerance to pesticides by many species of insects and
the development of tolerance to antibiotics by bacteria. For
the anti-evolutionists, these examples do not matter as they
represent evolution within the bounds of a species which
dier from macroevolution ( trans-specic evolution). Based
on the generally accepted belief by many evolutionists of
vertical boundaries between species taxa, the anti-evolutionist
can point out quite properly that evolution within the bounds
of a species cannot be extended automatically to trans-specic
evolution.
Accepting the concept of gradual evolutionary change, the
unbridgeable horizontal boundaries present between species
taxa cannot be extended to vertical boundaries. In a phyletic
lineage, no species boundaries exist between a population
existing at a particular time and one existing at a later time, no
matter how much phyletic evolutionary change occurs (Bock
1979). Speciation exists as a distinct evolutionary process, but
one species taxon does not evolve by phyletic evolution into
another species taxon. Rather in phyletic evolution, the
attributes of the species taxon can change and change and
change, but no matter how much these attributes modify, no
species boundary is passed. A distinction simply does not exist
between evolution within the bounds of a species and
evolution beyond the species limit.
In speciation, two or more species taxa evolve from an
ancestral species taxon. When tracing the phyletic lineage from
the ancestral species taxon to each of the descendent species
taxa, no species boundaries exist as evolution is a gradual
change. This leads to a peculiar conundrum that no boundary
exists between an ancestral species taxon X and a descendent
species taxon A and no boundary exists between the same
ancestral species taxon X and a second descendent species
taxon B, but a denite and non-bridgeable horizontal bound-
ary exists between species taxa A and B. This is not the
equivalent as saying that species X and A are the same and
that species X and B are the same, but only that these two pairs
of species taxa are not separated by species boundaries. Yet
species taxa A and B are separated from one another by a
denite, non-bridgeable horizontal boundary. Because of the
phyletic evolutionary change that has taken place in one, and
usually in both, of the lineages from X to A and from X to B,
characteristics of the ancestral species taxon X diers from
those in both species taxa A and B, yet no non-arbitrary
boundaries exist between X and A or between X and B. During
the phyletic evolution leading from X to A and from X to B,
no species boundaries had been transgressed.
Species taxa terminate when they become extinct that
particular phyletic lineage becomes extinct but they do not
have beginnings or births. It is not possible to speak of the age
of a species taxon such as the human species being 1.3 million
years old or whatever age is placed on the human species.
Because species taxa do not have beginnings, all existing
species taxa at any point in time are equally old. The notion
that a particular species has a certain age is used to indicate
only that the particular phenotypic characteristics of that
species appeared at that time.
Denitions of evolution
Before it is possible to talk about theories of biological
evolution, it is necessary to have a denition of this term. For
convenience of comparison, evolution will be dened as:
Evolution is change in organisms over time with the minimum
time being one generation. Hence evolutionary change is
observed between organisms of one generation and their
descendants. This denition does not include any condition
that the changes have to be genetic or hereditary. This is a
variational denition of evolution as pointed out by Lewontin
(1983: 63; see also Mayr 1988: 1516, 1991: 4344, 1997: 176)
which diers sharply from transformational evolution. I would
like to compare this denition with a series of other denitions
and comments taken from a number of well known sources
that would be utilized by many workers interested in the
meaning of biological evolution. These references are not
meant to be complete or to represent the full span of possible
denitions of evolution. Many important textbooks on evolu-
tion such as those by Ehrlich and Holm (1963), and by
Maynard Smith (1975), do not provide a denition of
biological evolution, although the former provides clear
denitions of phyletic evolution (p. 327) and of speciation
(p. 330) in its glossary.
Dobzhansky (1937: 11; 1951:16) in his epic book heralding
the beginning of the modern evolutionary synthesis stated that:
As evolution is a change in the genetic composition of
populations, the mechanisms of evolution constitute problems
of population genetics. His wording in the 1951 edition is
basically the same. He elaborated (Dobzhansky 1951:11) that:
The theory of evolution asserts that (1) the beings now living
have descended from dierent beings which lived in the past;
(2) the evolutionary changes were more or less gradual, so that
if we could assemble all of the individuals which have ever
inhabited the earth, a fairly continuous array of forms would
emerge; (3) the changes were predominately divergent, so that
the ancestors of the now living forms were on the whole less
dierent from each other than those forms themselves are; (4)
all of these changes have arisen from causes which now
continue to operate, and which therefore can be studied
experimentally. The rst and, in part, the third of these
statements are clearly historical evolutionary theory (see
below) which the second, fourth and, in part, the third are
nomological evolutionary theory (see below).
Mayr (2001: 286) in his glossary provided the denition that:
Evolution: The gradual process by which the living world has
been developed following the origin of life. And in the text of
his book, he (Mayr 2001: 76) wrote: Evolution is best
understood as the genetic turnover of the individuals of every
population from generation to generation. (italics his). This is
the standard genetic denition of evolution which excludes at
least one important form of evolutionary change that of
cultural evolution. Finally Mayr (2001: 264) stated: It is very
questionable whether the term evolutionary theory should be
used any longer. That evolution has occurred and takes place
all the time is a fact so overwhelmingly established that it has
92 Bock
become irrational to call it a theory. To be sure, there are
particular evolutionary theories such as those of common
descent, origin of life, gradualism, speciation and natural
selection, but scientic arguments about conicting theories
concerning these topics do not in any way aect the basic
conclusion that evolution as such is a fact. It has taken place
ever since the origin of life. Earlier Mayr (1991: 112) wrote:
Weismanns attitude toward evolution as such was close to
that of the modern evolutionist, for whom evolution is not a
theory but an accepted fact. Hence for Mayr, evolution is a
fact ( an objective empirical observation).
Gerd von Wahlert (2002), following Professor Vernadsky of
Leningrad, USSR who developed the concept of the bio-
sphere in the 1920s (see also Hutchinson 1965: 1), dened
evolution as development ( change) in the characteristics of
the biosphere over time. Whether this denition diers from
that of Mayr (above) depends on whether the term living
world possesses the same meaning as biosphere; I suspect not
quite. The latter includes not only living organisms but all
environmental aspects (including all physical factors) of the
Earth necessary to maintain these organisms. At least some of
these physical factors, such as the generation of an atmosphere
rich on oxygen, appear to be the consequence of the action of
living organisms. The important aspect of von Wahlerts
denition of evolution is that it includes a clear mention of the
external environment of living organisms which is an integral
factor of evolutionary causes and which is lacking in most
other denitions. The biosphere, in von Wahlerts approach,
includes not only the living organisms, but the physical
characteristics of the Earth. Therefore his expression history
of the biosphere includes transformational evolution of the
earth which is change over time of the same object and
biological ( variational) evolution which is the observed
change in characteristics of living organisms from one genera-
tion to a descendant one. No reason exists why these two types
of change cannot be included in a single denition of
evolution, but one must be careful in any further development
of causes and processes involved in the historical modication
of the biosphere. Von Wahlerts denition does not include
any claim that evolution is factual or that it has to be genetic.
In his well-known textbook on evolution, Futuyma (1979
503; 1986: 551; 1998, Glossary; 2005: 547) provided the
complex statement: Evolution in a broad sense, the origin of
entities possessing dierent states of one or more character-
istics, and changes in their proportions over time. Organic
evolution, or biological evolution, is a change over time of the
proportions of individual organisms diering genetically in
one or more traits; such as changes transpire by the origin and
subsequent alteration of the frequencies of alleles or genotypes
from generation to generation within populations, by the
alterations of the proportions of genetically dierentiated
populations of a species, or by changes in the numbers of
species with dierent characteristics, thereby altering the
frequency of one or more traits within a higher taxon. His
genetic denition of evolution also excludes cultural evolution.
In the very beginning of the text Futuyma (1986: 13) stated:
As is indicated above, evolutionary biology consists of two
principal endeavors: inferring the history of evolution and
elucidating its mechanisms. which clearly appreciates both the
historical and the nomological theories of evolution. And he
wrote further (1979: 14; 1986: 16): Evolution, a fact, rather
than a hypothesis, is the central unifying concept of biology.
and hence agreed with Mayr on this point. In the next edition
of this textbook, Futuyma (1998: 11) retained this position and
wrote that: In the light of the preceding discussion, evolution
[? historical change] is a scientic fact. But it is explained by
evolutionary theory. He repeated this position (Futuyma
2005: 13), saying that: Given these denitions, evolution is a
fact. But the fact of evolution is explained by evolutionary
theory. (italics his). He added the explanation (Futuyma 2005:
13) that: What we call facts are hypotheses that have acquired
so much supporting evidence that we act as if they were true.
These are what Popper and many other philosophers of science
consider as very well corroborated theories and are still best
called theories to distinguish them from objective empirical
observations which can be considered as facts within the limits
of observational abilities and underlying theories.
In another well-known text, Strickberger (2000: 640) dened
evolution in his glossary as: Evolution Genetic changes in
populations of organisms through time that lead to dierences
among them. Again cultural evolution is excluded. Further
Strickberger (2000: 636) dened: Darwinism the concept,
proposed by Charles Darwin, that biological evolution has led
to many dierent highly adapted species through natural
selection acting on hereditary variations in populations.
Scotts (2004: 2345) position on evolutionary ideas are
presented in her Chapter 2 Evolution. where she wrote under
the major heading EVOLUTION BROAD AND NAR-
ROW: In biology, evolution is the inference that living things
share common ancestors and have, in Darwins words,
descended with modication from these ancestors. The main
but not the only mechanism of biological evolution is
natural selection. (pp. 2324). And just below, she stated that
Biological evolution is dened as the descent of living things
from ancestors from which they dier. (p. 27).
Under the major heading BIOLOGICAL EVOLUTION,
she wrote: Descent connotes heredity, and indeed members of
species pass genes from generation to generation. (p. 27), and
at the end of this section, that: Evolutionary biologists are
concerned both with the history of life the tracing of lifes
genealogy and the processes and mechanisms that produced
the tree of life. This distinction between the pattern of
evolution and the process of evolution is relevant to the
evaluation of some of the criticisms of evolution that will
emerge later in this book. First lets look briey at the history
of life.
Although Scott made a distinction between discovering the
process of evolutionary change and working out the history of
life, and does not claim that evolution is a fact, the order in
which she presented her ideas is unclear.
Denton (1986: 3668) in his Chapter 2 on THE THEORY
OF EVOLUTION did not provide a clear denition of
evolution but he was primarily concerned with historical
evolutionary theory ( the history of life) and with diculties
of major evolutionary change in the same way as was Behle
(1996). Denton was concerned almost exclusively to historical
evolutionary theory. And although he did not use this term, he
advocated intelligent design as his primary disagreement in
dealing with macroevolution.
A committee was established in the 1990s by the American
Society of Naturalists and published its report under the title
Evolution. Science and Society (Meager and Futuyma 2001),
which is well worth reading as it covers thoroughly a broad
spectrum of aspects of evolutionary biology, its teaching and
its signicance to many aspects of human society. I would like
to present several quotes from this report.
Under, II. WHAT IS EVOLUTION? they wrote (p. 3):
Biological evolution consists of change in the hereditary
characteristics of groups of organisms over the course of
generations. Groups of organisms, termed populations and
species, are formed by the division of ancestral populations or
species, and the descendant groups then change independently.
Hence, from a long-term perspective, evolution is the descent,
with modication, of dierent lineages from common ances-
tors. Thus, the history of evolution has two major compo-
nents: the branching of lineages, and changes within lineages
(including extinction). Initially similar species become ever
more dierent, so that over the course of sucient time, they
may come to dier profoundly.
And later in the same section (p. 4) they continued:
Evolutionary theory is a body of statements about the
processes of evolution that are believed to have caused the
history of evolutionary events. Biological (or organic) evolu-
tion occurs as the consequence of several fundamental
processes. These processes are both random and nonrandom.
(italics theirs).
And they claried further (p. 5), saying: It is important to
distinguish between the history of evolution and the process
held to explain this history. Most biologists regard the history
of evolution the proposition that all species have descended,
with modication, from common ancestors as a fact that is,
a claim supported by such overwhelming evidence that it is
accepted as true. The body of principles that describe the
causal processes of evolution, such as mutation, genetic drift,
and natural selection, constitutes the theory of evolution.
Next under, III. WHAT ARE THE GOALS OF EVOLU-
TIONARY BIOLOGY? they wrote (p. 5): Evolutionary
biology is the discipline that describes the history of life and
investigates the processes that account for this history.
Evolutionary biology has two encompassing goals:
(1) To discover the history of life on earth: that is, (1) to
determine the ancestor-descendant relationships among all
species that have ever lived their phylogeny; (2) to determine
the times at which they originated and became extinct; and (3)
to determine the origin of and the rate and course of change in
their characteristics.
(2) To understand the causal processes of evolution: that is, to
understand (1) the origins of hereditary variations; (2) how
various processes act to aect the fate of those variations; (3)
the relative importance of the many co-acting processes of
change; (4) how rapidly changes occur; (5) how processes such
as mutation, natural selection, and genetic drift have given rise
to the diverse molecular, anatomical, behavioral, and other
characteristics of dierent organisms; and (6) how populations
become dierent species. Virtually all of biology bears on this
vast project of understanding the causes of evolution, and
reciprocally, understanding the processes of evolution informs
every area of biology. (italics theirs).
After pursuing the comments in the scientic literature, it is
always useful to consult dictionaries; as for example, referring
to the Concise Oxford English Dictionary (Pearsall 2002) and
the Websters Third International Dictionary of the English
Language. Unabridged (Gove 1963). Other dictionaries would
provide much the same denitions.
First in Pearsall (2002), the following denitions are oered:
evolution the process by which dierent kinds of
living organisms are believed to have developed from
earlier forms, especially by natural selection. (p. 494)
Darwinism the theory of the evolution of species,
advanced by the English natural historian Charles
Darwin (180982). (p. 364)
phylogeny another term for phylogenesis. (p. 1078)
phylogenesis the evolutionary development and
diversication of a species or group of organisms.
(p. 1078)
Next Gove (1963) presents the following denitions:
evolution the development of a race, species or
other group Phylogeny
the process by which through a series of changes or
steps any living organism or group of organisms has
acquired the morphological or physiological charac-
ters that distinguish it.
the theory that the various types of animals and
plants have their origin in other preexisting types, the
distinguishable dierences being as a result of mod-
ications in successive generations. (p.789)
phylogeny the racial history of a specied kind of
organism.
the evolution of a race or generically related group
of organisms (such as a species, family or order) as
distinguished from the development of the individual
organism. (p. 1706)
Many evolutionists and philosophers writing texts on
evolution or philosophical analyses on evolution never dene
just what they mean by biological evolution. Dennett (1995:
21) wrote: Let me lay my cards on the table. If I were to give
an award for the single best idea anyone ever had, Id give it to
Darwin, ahead of Newton and Einstein and everyone else. As
far as I can determine Dennett does not provide a denition of
evolution. He did say (1995: 39) that: Darwins project in
Origin can be divided in two: to prove that modern species
were revised descendants of earlier species species had
evolved and to show how this process of descent with
modications had occurred. (italics his). Although numerous
aspects of evolutionary theory are discussed in the excellent
volume The Darwinian Heritage (Kohn 1985), none of the
many authors presented a denition of evolution.
With just this diversity of denitions, of lack of denitions
of evolution and whether evolution is a theory or a fact, it is a
bit surprising that any decisions could have been reached in the
two most recent court cases in the USA on the teaching of the
theory of evolution as advocated by most evolutionary
biologists versus alternative approaches, such as scientic
creationism and intelligent design.
Two points should be mentioned about this series of
denitions of evolution. Although several authors (Dobzhan-
sky, Futuyma, Scott, Meager and Futuyma, and Dennett)
distinguished between the evolutionary history of organisms
and the mechanisms (causes and processes) of evolution by
which these organisms changed over time, they almost always
placed discussion of the mechanisms after presentation of the
history of the organisms, rather than the more logical reverse
order. The major reason delaying Darwin until 1859 on the
publication of his ideas about transformation of organisms
was that he believed he could not present a valid case for the
94 Bock
evolutionary history of organisms without rst providing a
strongly supported set of mechanisms. He was completely
correct nomological theory before historical theory. Darwin
was well aware of the criticism and abuse piled upon the head
of the then unknown author of Vestiges of a natural history of
creation (Chambers 1844), largely because of the unrealistic
method Chambers suggested for modication of organisms
over time (see Secord 2000). Darwin resolved that he would
not fall into the same pit; he wanted to present a solidly
supported mechanism of natural selection when he advocated
his novel concept of evolution of living organisms.
Second, some of these workers used the expression the
theory of evolution to denote the set of mechanisms of
evolutionary change and considered the evolutionary history
of organisms so well supported that it is factual, not a theory.
(Here one could point out the position of most physicists
towards the end of the nineteenth century on the absolute
truthfulness of Newtons laws of motion.) These workers
overlooked the point that almost all anti-evolutionists are
uninterested in evolutionary mechanisms or may even accept
them for change within the limits of a species. The anti-
evolutionists major dispute is with evolutionary history of
organisms which they argue is a theory only and not a very
well supported one at that. They certainly do not consider the
evolutionary history of organisms so well supported that it can
be regarded as factual in contrast to the discussion in Meagher
and Futuyma (2001: 43).
Having presented these other denitions of evolution, I will
reiterate the one advocated above:
Evolution: change in organisms over time with the minimum
time being one generation. Hence evolutionary change is that
observed between organisms of one generation and their
descendants. This is variational or Darwinian evolution
(Lewontin 1983: 63; Mayr 1988: 1516, 1991: 4344, 1997:
176). It diers sharply from transformational evolution, i.e.,
change of the same object, such a star or the earth, over time.
Hence change in the characteristics of an individual organism
during its life is not an evolutionary change, but an onto-
genetic modication ( transformational evolution; Lewontin
1983; Mayr 1988). Aside from one very vague point in
common that change over time occurs, there is nothing similar
in transformational and variational evolution the evolution
of the planet earth and the evolution of life on earth dier
completely. In a similar vein, there in nothing causally similar
between ontogeny (embryologic development during the life of
an individual organism transformational evolution) and
Haeckelian phylogeny ( variational evolution), although
the latter is the basis for the former (the foundation for Mayrs
concept of dual causation in biology, Mayr 2004: 30, and
elsewhere).
Note that the denition of evolution advocated herein does
not include any statement of heredity or genetic change.
Several types of variational evolution exist depending on how
information is transmitted from one generation to the next.
These are:
(1) Genetic evolution in which transmission of information
from generation to generation is via genes or other elements in
the gametes and which is the form of biological evolution
proposed by Darwin and specied in almost all denitions of
biological evolution ever since.
(2) Cultural evolution in which transmission of information
from generation to generation is via learning by the individual
from adults (e.g., its parents) or from other individuals which
are usually conspecic, but do not have to be conspecic; this
transmission is non-genetic and depends only on the ability of
organisms to learn the phenotypic attribute (such as song or
the trail from the summer to the winter ranges) that can be
further transferred by learning, from generation to generation.
Cultural evolution is far commoner than most workers
consider although forgotten about in almost all denitions of
biological evolution. Modications occur when there are
errors in the learning process.
(3) Template (or perhaps epigenetic) evolution in which trans-
mission from generation to generation is by non-learning the
copying of a phenotypic characteristic in the ospring from a
template in the phenotype of the previous generation. Mod-
ications occur if there are accidental changes in the pheno-
typic feature of the parental generation which are hence copied
by the ospring when the new phenotype is developed as
shown by the formation of the teeth (notches) around the
mouth of the protozoa, Diugia corona, in the sandy shell
formed around the body of this animal (Jennings 1937;
Nanney 1968). Template evolution is perhaps quite rare and
is overlooked in almost all discussions of evolution.
Perhaps one could also include in this classication of types
of biological evolution modications of the phenotype that
occur because of changes in the external environment and/or
the internal interactions between parts of the body. The ability
for these modications in the phenotype is genetic, although
the phenotypic change is not so and will reverse in the next
generation if the environmental/internal interaction reverses.
These modications have been listed under a series of terms
from physiological adaptation to somatic modications (Bock
and von Wahlert 1965). These changes can take place during
ontogenetic development and/or during the adult stage, and
may be reversible during the life of the organism; the
modications are determined by alterations in the external
environment or in the internal interactions between parts of
the body (such as muscles aecting the shape of the bony
skeleton); no modications occur in the genetic basis for the
observed phenotypic changes. Although this type of evolu-
tionary change is exceedingly common, I do not include it in
the above list as types of evolutionary change as it does not
depend on a dierent form of information transmission from
an organism in one generation to the next. Unfortunately until
recently there has been little consideration of this non-genetic
evolutionary change (Bock and von Wahlert 1965; West-
Everard 2003; Starck 2005) in spite of its extreme commonness
in evolutionary change.
Evolution (here I will restrict myself, only for simplicity, to
genetic evolution and to sexually reproducing organisms) can
be subdivided into two major processes, which are:
(A) Phyletic evolution or anagenesis is evolutionary change
in a single phyletic lineage. Phyletic evolution occurs without
any speciation; hence no matter how much modication occurs
in phyletic evolution, no species boundaries are crossed. The
major causes of phyletic evolution are: (1) the generation of
individual phenotypic variation; and (2) Selective demands
arising from the external environment. I use selective demands
arising from the external environment because the term
natural selection is usually dened by evolutionists as an
outcome, not a cause (Bock 1993). Although Darwin used
natural selection as a cause in most of his 1859 book, his
clearest denition of this term (Darwin 1859: 61, see below) is
denitely an outcome denition which was adopted by R.A.
Fisher (1930) and J.B.S. Haldane (1932) see also Huxley
(1942) who stated in his Preface that his treatment of
evolutionary mechanisms are based strongly on the earlier
books of Fisher and Haldane). Most evolutionists (e.g.,
Futuyma 1986: 554; 1998: Glossary; 2005: 550) cite a denition
of natural selection which is basically nonrandom dierential
reproduction of genotypes (an outcome) that is adopted from
the analyses of Fisher (1930) and Haldane (1932) and then use
this term as a cause, leading to considerable confusion (Bock
1993: 1115). Other workers (e.g., Lewontin 1970: 1) labeled
natural selection as a motive force (? cause) and still others,
such as Endler (1986: 4) stated that: Natural selection can be
dened as a process in which... (italics his). Interestingly the
three conditions given by these two workers for natural
selection are basically the same.
(B) Speciation or cladogenesis is the multiplication of species
from an ancestral species (Mayr 1942, 1963, 2001) and can be
thought as the splitting of a single phyletic lineage into two or
more. (Speciation occurs only in sexually reproducing organ-
isms and hence could be considered as a narrower process than
cladogenesis which is general to all organisms.) Complete
speciation always involves phyletic evolution in at least one of
the phyletic lineages, and generally in both. No special causes
exist for speciation other than those operating for phyletic
evolution. But a denite initial condition is needed in the form
of an external barrier separating two populations of the
original species during which time intrinsic isolating mecha-
nisms for genetic isolation evolve by phyletic evolution.
Darwins ve theories
Darwin always referred to his ideas published in his On the
Origin of Species as my theory, always in the singular which has
been a primary source of confusion ever since. Mayr (1982: 8)
noted in the introductory chapter of his The Growth of
Biological Thought: Similarly, anyone whom writes about
Darwins theory of evolution in the singular, without segre-
gating the theories of gradual evolution, common descent,
speciation, and the mechanism of natural selection, will be
quite unable to discuss the subject competently. And in a most
important, but apparently little known paper, Ernst Mayr
(1985; see also 2004) elaborated on this point and demonstra-
ted that Darwins theory of evolution as originally advocated
in his On the Origin of Species (1859) was actually a bundle of
ve separate but interrelated theories. Mayr showed that these
theories in various combinations, but not all, were dieren-
tially advocated by diverse biologists before Darwin or by
most of his contemporaries. Only after the evolutionary
synthesis of 19371948 were all of these theories accepted by
most evolutionists. But even after the evolutionary synthesis,
almost all biologists and philosophers still considered evolu-
tion as an undivided, historical theory.
The ve separate theories found in Darwins 1859 book can
still characterize the major areas within evolutionary biology
today; they are:
(1) Evolutionas suchis the theory that states that all populations
of organisms are changing over time, with the minimum time
period being one generation (Mayr 1985: 757758).
(2) Gradualism is the idea that evolutionary change takes place
in steps of the magnitude seen between parents and ospring
and never in large sudden saltations or jumps. Evolutionary
jumps do not take place between species or taxa of higher
levels such as expressed in the idea that the rst bird hatched
from a reptilian egg (Mayr 1985: 761764).
(3) Multiplication of species states that there is splitting of
phylogenetic lineages in addition to transformational change
within lineages. Hence evolutionary change includes two
processes phyletic evolution or transformation and speci-
ation. Although, Darwin appreciated the need for speciation in
his general ideas about evolution, he didnt provide any clear
discussion on how speciation took place or how it diered
from phyletic evolution (Mayr 1985: 764767).
(4) Natural selection is Darwins mechanism for phyletic
transformation. Today this would be expressed as the complete
causes, initial and boundary conditions, or mechanisms of
evolutionary change, regardless of the diverse causes that
dierent evolutionists would include (Mayr 1985: 767771).
Here one must be careful because Darwin used the term natural
selection interchangeably as a cause of evolution, as the process
of evolution and as the outcome. (see Darwin 1859: 61, where
he wrote that: Owing to this struggle for life, any variation,
however slight and from whatever cause proceeding, if it be in
any degree protable to an individual of any species, in its
innitely complex relations to other organic beings and to
external nature, will tend to the preservation of that individual,
and will generally be inherited by its ospring. The ospring,
also, will thus have a better chance of surviving, for, of the
many individuals of any species which are periodically born,
but a small number can survive. I have called this principle. by
which each slight variation, if useful, is preserved by the termof
Natural Selection, in order to mark its relation to mans power
of selection.) This is clearly an outcome denition of natural
selection and is the one advocated by Fisher (1930) and by
Haldane (1932) in their analyses, and has been broadly
accepted by population geneticists and evolutionists.
(5) Common descent which implies that all species or popu-
lations of organisms have descended with modication from
common ancestors; this descent includes both modication
and branching (Mayr 1985: 758671). Darwinian common
descent ( historical evolution although Darwin did not use
this term in 1859) is equivalent to Haeckelian phylogeny
(Haeckel, 1866). Hennigian phylogeny is equal only to the
branching aspect of Haeckelian phylogeny (Mayr and Bock
2002). Common descent is expressed in Darwinian classica-
tions, Haeckelian dendrograms and Hennigian cladograms,
which are all clearly theoretical statements. In recent years,
common descent has been labeled by some workers as The
Tree of Life.
Because most biologists and lay-persons are mostly interes-
ted in the last of these ve theories which is clearly historical
and because most people still consider evolution to be a single
theory, the overwhelming conclusion by most biologists and
philosophers of science is that the theory of evolution is only
historical as did Mayr (2004: 3233) and Ghiselin (2005: 133)
who stated that: Evolutionary biology is an historical science.
Unfortunately this is not completely true. This belief is the
source for much of the erroneous analysis in the philosophy of
evolutionary biology. Mayr (2004: 33) mentioned historical
narratives, but did not provide details for this approach.
Evolutionary explanations
What scientists do is to provide explanations of phenomena,
but not any type of explanations. The approach used by
scientists is to formulate theoretical statements that might
apply to the phenomenon, generate deductions from these
theories, and nally test these deductions against objective
96 Bock
empirical observations. What is absolutely essential in the
scientic methodology is not that empirical observations are
made, but that these observations are objective as opposed to
subjective hence the term objective science. Objective
empirical observations in the philosophy of science means
that the same observations can be made by any person having
the abilities to do so. Abilities means having the proper sense
organs and training. People with certain types of color vision
deciencies cannot make the necessary observations depending
on color in many elds of science. Subjective observations are
ones that only certain persons can make but not everyone.
In an early paper commenting on whether only the choice of
words separated the ideas of G.G. Simpson and H.O.
Schindewolf, von Wahlert and I argued that evolutionary
theory had to be considered both as nomological ( causes,
mechanisms) and as historical that a major dierence existed
between these two approaches (Bock and von Wahlert 1963). I
continued to muse on this distinction between approaches to
evolutionary biology and published my ideas in a series of
papers (Bock 1973, 1978, 1988, 1991, 1994, 1999, 2000a,b,
2004a), but they were either not fully formed or they were
published in rather specialized symposium volumes. Herein I
would like to describe and contrast two major explanatory
systems in science and to show their relationship to one
another. These systems are N-D Es and H-N Es. Although the
latter are probably general for science, they appear to be of
most signicance in elds such as biology, geology and
astronomy. Almost all philosophers of science have concen-
trated exclusively on N-D Es as I know of no general treatment
of H-N Es. An additional system of explanations exists in
biology the dichotomy of Functional Explanations versus
Evolutionary Explanations (see below). Hence it is essential
not only to characterize carefully the properties of N-D Es and
H-N Es, but to show which of these are functional and which
are evolutionary explanations.
Explanations in biology deal with phenotypic attributes of
organisms and their genetic basis, including interactions
between phenotypic attributes found in the same individual
organisms, correlations between phenotypic attributes and
diverse aspects of the external environment, and the relation-
ships among the features in dierent organisms, be they
conspecic or of dierent species. Hence, given any phenotypic
attribute, a complete ( full) explanation includes resolving:
(1) all existing physical-chemical properties (form, function,
biological role, etc.) which are functional explanations; (2) its
ontogenetic development resulting from interactions between
the existing genotype and the external environment (pro-
grammed systems, Mayr 1974, 1988, 1997) which are also
functional explanations; and (3) its evolutionary origin and,
therewith, the evolutionary origin of the genotype (constituting
the programmed systems), which constitutes evolutionary
explanations (Bock and von Wahlert 1963). Although the
topic of full versus partial explanations in biology is an
important one, I will not consider it further herein except to
say that partial biological explanations can be most important
and are all that can be achieved and/or desired in most cases.
Mayr (1961, 2004) has argued strongly that the formation of
all biological phenotypic attributes depends on two dierent
set of causes working simultaneously which he has termed
proximal and ultimate causes and which form the basis of his
seminal conclusions about dual causation in biology and the
autonomy of biology from the physical sciences. His proximal
and ultimate causes are better called functional ( proximal)
and evolutionary (genetic; ultimate) causes with the latter
result from the evolutionary history of the organism.
Throughout, I will use functional explanations in the general
sense of functional analyses in biology (Mayr 1982), not in the
sense of functional explanations in philosophy (see Nagel
1961) as the latter do not appear to be of any value in scientic
explanations and are best omitted from scientic explanations.
In addition, almost all discussions of function in biology by
philosophers of science are presented in a historical evolution-
ary sense which places them in sharp contrast to explanations
by most functional biologists; this distinction should be noted
carefully when comparing the conclusions of papers on
function and functional explanations by most biologists versus
those by most philosophers. A further problem is that many
biologists in the mid-nineteenth century and some even today,
equated functional explanations with teleological explana-
tions. When many biologists realized that Darwinian evolution
eliminated teleology from biology, they also eliminated
discussions about functional properties of biological features
(as well as functional explanations) from their evolutionary
analyses a good example of throwing the baby out with the
bath water. This thinking resulted in serious problems for
evolutionary biology as most, if not all, evolutionary expla-
nations depend on well established functional explanations
and on a full understanding of the interactions of organismic
features with the external environment (Hutchinson 1965;
Bock 2002, 2003; von Wahlert 2002).
Nomological-deductive explanations
Nomological-deductive explanations are the standard form of
explanation in science covering law explanation and deal
with general explanations of a class of phenomena, asking how
has each occurred? This is performed by formulating a
deduction from an appropriate set of laws (be they causes,
processes, or outcomes) and a set of initial and boundary
conditions (observations of the conditions in which the
phenomenon exists), both of which form the explanatory
sentence, or explanans, and from these reach a particular
conclusion or deduction, the explanandum. (Hempel and
Oppenheim 1948; Hempel 1965: 335338). The deduction is
then compared with objective empirical observations factual
observations of the phenomenon. If the observed phenomenon
agrees with the deduction (the degree of error is dependent on
what would be considered admissible), the explanation is
accepted. If an explanandum, resulting from the conjunction
of the set of facts invoked (initial and boundary conditions)
and the set of general laws, disagrees with objective empirical
observations, then that N-D E is not valid (has been falsied),
and one must search for the reasons underlying this falsica-
tion. Falsication means only that the explanandum does not
agree with independent, objective, empirical observations. If
the explanation is not accepted, it is necessary to investigate
the source of the error be it the set of law-like statements,
or the set of initial and boundary conditions used to formulate
the deduction, or the objective empirical observations used to
test the deduction. Falsication of an explanation does not
automatically imply that the general laws used in the explan-
ation are in error, although this is a possibility. Possibly, the
initial or boundary conditions used in the empirical test were
wrong, or the empirical observations were incorrect.
Because they are general, one nomological scientic theory
can be used to test another, and two or more nomological
theories can be included under the umbrella of an over-
arching theory. And as will be discussed below, nomological
scientic theories are required in the testing of any historical
scientic theories.
Nomological-deductive explanations answer the question:
how has a particular phenomenon [explanandum] occurred?
N-D Es apply to universals (non-limited number of phenom-
ena), do not depend on the past history of the objects or the
phenomena being explained, and their premises (the nomo-
logical statements) are assumed to be always true. In saying
that N-D Es apply to universals, these explanations are not
temporally-spatially restricted within the proper region of the
phenomena, which for biology is the earth and more specif-
ically the surface ( the upper part of the crust) of the earth.
Examples of N-D Es include clarication of oceanic tides using
gravity and of phyletic evolution evoking natural selection
(nonrandom, dierential survival and reproduction of organ-
isms). Explanation of how a certain feature of an organism is
an adaptation to selective demands arising from the external
environment is a N-D E, not a H-N E; the origin of the
adaptation, which is an entirely dierent question, is a
historical-narrative explanation (see below).
For almost all philosophers of science (e.g., Popper 1959;
Nagel 1961) scientic methods apply only to N-D E. This is
interesting because Poppers approach has been accepted by
many systematists (e.g. cladists) as the foundation for their
analyses of the relationships among organisms; this cannot be
not correct because any explanations concerning the tree of
life, such as phylogenetic classication, deal with singulars (the
existing phylogeny of organisms) and not with universals (see
below, and Bock 2000a, 2004a).
Many philosophers of science and many biologists have
rejected the idea that law-like statements exist in science (e.g.,
Mayr 1997: 6063; but see Elgin 2006 for an argument for laws
in biology), sometimes substituting a vague idea of concepts in
place of law-like statements. The reasons for this rejection of
law-like statements by biologists and philosophers of biology is
not entirely clear, but may stem from the conclusion that
variation of phenomenon in biological organisms precludes the
existence of laws in biology rather than that this variation
arises from the nature of the initial and boundary conditions
which generally vary, often considerably, among biological
objects such as individual members of the same species and
will, of course, result in variation in the deductions. In this
connection, determinism is often raised as a critical factor
without the realization that the scientist must set the accept-
able limits of determinism in every case. Further, many of
these workers accept that only a single type of explanations
exists in science and hence confuse H-N Es with N-D Es. If this
is done, then the conclusion follows that no laws exist in
biological explanations, which include historical narratives,
because historical laws do not exist. At this point the argument
becomes circular.
Historical-narrative explanations
Contrary to the beliefs of many philosophers, not all science is
nomological. Some sciences, such as biology and geology, are
very largely historical. But little or no mention of these
explanations is found in the literature of the philosophy of
science (e.g., Feigl and Brodbeck 1953; Boyd, Gasper and
Trout 1991; Cornwell 2004). Indeed, considering numbers of
scientists and amounts of funding (which includes medicine
and agriculture as well as biology, geology, and some
astronomy), sciences with a historical aspect are now and
have been for a long time in the large majority. Why
philosophers failed to realize the importance of historical
science may be because the philosophy of science is little more
than one and a half centuries old and has been concentrated
very largely on physics as the ideal science. And physicists have
developed their science as a strictly non-historical inquiry,
which is certainly a completely valid thing to do and is
certainly the prerogative of physicists. But others do not have
to accept physics as the science on which to base all philosophy
of science. In view of the overwhelming acceptance that there
are no historical laws, the major diculty for philosophers in
dealing with these historical aspects of science would be how to
extend the nomological-deductive approach to them (see
Hempel 2001, especially chapters 14 and 15). This was clearly
the reason why Popper (1977; see Hull 1999 for an excellent
analysis of this matter) said, or implied, that evolutionary
theory was not scientic. The diculties of dealing with N-D
versus H-N Explanations and assigning dierent explanations
in evolutionary biology to one or the other of these two types
are well illustrate in the interesting analysis of adaptation by
Amundson (1996).
Historical-Narrative Explanations provide an understand-
ing of the existing attributes of a particular set of objects or
phenomena at specied points in time; these explanations
absolutely depend on the past history of these objects and to be
scientic they must use pertinent N-D Es. The latter point is
essential! Any explanation of historical events that is not based
on pertinent N-D Es is not scientic. This includes not only
approaches such as scientic creationism and intelligent
design, but also claims by some scientists of the existence of
pure order and/or design in nature (including some approaches
to biological classication, see Brower 2000), self-determinism
and structuralism. Phenomena and objects explained by a
HN E are singulars, not universals, and have denite spatial-
temporal positions. HN Es are considered in a non-deductive
and probabilistic basis with the hope of reaching the most
reasonable and probable explanation for the objects studied.
Several aspects of H-N Es are stressed, the, rst being the
most important:
(1) HN Es must be based on pertinent and well-tested N-D
Es, and these N-D Es, together with the pertinent empirical
observations testing them, form part of the chain of arguments
used in testing the H-N E. If no N-D Es exist or if the N-D Es
are poorly tested, then that H-N E lies outside of science.
(2) These explanations are historical in character, which means
that earlier events aect later events earlier events form the
initial conditions for explaining later events. Great care must
be given to formulating the analysis within the presumed
correct chronological order of events and changes.
(3) H-N Es must be tested against objective empirical
observations, which may involve a chain of arguments,
including the underlying N-D Es and the objective empirical
observations used to test these N-D Es.
(4) Acceptance of a particular H-N E is always given on a
probability basis. This is necessary as these explanations
frequently employ some conicting N-D Es and because of
generally considerable uncertainty over the initial and bound-
ary conditions involved in the explanation.
(5) H-N Es are not universal as are N-D Es, in that a successful
H-N explanation for one phenomenon (e.g. origin of
mammalian homoiothermy) need not hold for a similar
98 Bock
phenomenon (e.g. origin of avian homoiothermy) even if both
H-N Es include a number of the same N-D Es.
(6) Because of their complexity, the possible confusion between
competing explanations and the diculty in identifying valid
conrming or falsifying tests, H-N Es must be stated clearly
and in the presumed proper chronological order. Failure to do
this may preclude meaningful tests or appraisal of rival H-N
Es.
(7) Generally the more precisely a H-N E is stated, the more
dicult it is to test and support it. The H-N E that humans
have evolved from a greater chimpanzee-like (Pan troglodytes)
ancestor is more dicult to support than that humans evolved
from an anthropoid ancestor which is more dicult to support
than that of humans evolved from a primate ancestor, etc.
Historical-narrative explanations in biology include the
evolution, phylogeny and classication of organisms or the
evolutionary history of their genetic characteristics or of their
phenotypic attributes that is, anything related to the history
of life, such as historical biogeography. All full explanations in
biology would include a historical-narrative portion which is
why full biological explanations are so dicult to formulate
and test.
Because they deal with singular events, particular historical
scientic theories cannot be used to test any other scientic
theories, be they nomological or historical.
Both nomological-deductive and H-N Es are scientic under
the criterion of demarcation for scientic explanations advo-
cated by almost all philosophers of science in that they are
both available for testing against objective, empirical obser-
vations. N-D and H-N Explanations dier in the many ways of
how they are expressed, tested, and used to test other
theoretical statements, and must not be confused. The accu-
racy of most tests of H-N Es may be weak, and a distinction
must be made between valid tests and weak or unconvincing
tests (see Bock 1989b: 339342, discussing the concept of
homology). Many of the tests available for HN Es are valid,
but are relatively poor or non-robust and should not be
rejected as invalid tests. A robust, valid test is one that has a
high ability to distinguish between correct and an incorrect
hypotheses.
Being theoretical scientic statements, H-N Es are available
to tests against empirical observations, but such tests are often
dicult and inconclusive. Generally H-N Es are not tested by
falsication (in spite of numerous statements in the literature)
but usually by conrmation with the addition of more and
more corroborating support. This procedure is closely akin, if
not identical, to induction in the strict sense of that concept.
Objections cannot be raised to inductive testing of H-N Es
because they are theoretical statements about a singular,
containing a nite number of objects, in contrast to N-D Es
which cover universals or an unlimited number of objects.
Testing of H-N Es depends on argument chains involving
pertinent N-D Es and often on a large number of background
assumptions (hypotheses, many being initial and boundary
conditions), and they must be nally tested against objective
empirical observations. One should proceed to the empirical
observations as directly as possible, although the argument
chain is often complex. The empirical observations and their
roles as tests, whether falsifying or conrming, should be
designated clearly.
What makes H-N Es scientic is point 1 (above) that H-N
Es must be based on pertinent and well-tested N-D Es,
and these N-D Es, together with the pertinent empirical
observations, form part of the chain of arguments used in
testing the H-N E. If no N-D Es exist or if these N-D Es have
not been well corroborated, then that H-N E lies outside of
science. The argument that scientic order exists in nature by
itself and in the absence of any N-DE is simply invalid as a
scientic statement.
Theories of evolution
As some sciences contain very dierent types of explanations,
it seems reasonable to propose that in such sciences as biology,
geology and astronomy, two dierent types of theory exist
these being nomological theories and historical theories.
Therefore we should speak of nomological evolutionary theory
and historical evolutionary theory. Further, for historical
evolutionary theory, it is best to consider both general theory
and special theories. One can discuss general historical
evolutionary theory such as the Haeckelian phylogeny of
organisms and special theories such as the evolutionary history
of birds, of insects, of aquatic carnivores. etc. In geology, the
movement of continental plates over the earths surface would
be a general historical theory and the splitting, including the
time, of North and South America from Europe and Africa to
form the Atlantic Ocean would be a special theory.
Lets reconsider the ve theories of Darwin as outlined by
Mayr (1985). The rst four theories are clearly nomological
evolutionary theories, and are:
(1) Evolution as such is the theory that states that all
populations of organisms are changing over time, with the
minimum time period being one generation.
(2) Evolutionary change is gradualism in that it takes place in
steps of the magnitude seen between parents and ospring and
never in large sudden saltations or jumps.
(3) Evolutionary change includes two processes, namely
phyletic evolution or transformation and speciation. Multipli-
cation of species occurs by a splitting of phylogenetic lineages
as well as phyletic change within at lease one of the two
lineages.
(4) Evolutionary change takes place as the result of a small
number of causes, of which the most important are the origin
of (genetically based) phenotypically varying individuals in the
population and the action on these individuals of selective
demands arising from the external environment. Natural
selection was Darwins term for the overall mechanism of
phyletic transformation.
Recall that there are no special nomological causes restricted
to speciation. Although the process of speciation is nomolog-
ical, it depends on the nomological causes of phyletic evolution
plus the important initial condition of an external barrier
separating two populations for a suciently long period
during which Intrinsic Isolating Mechanisms for Genetic
Isolation evolve (Mayr 1963; Bock 1995, 2004b).
These four theories are all nomological albeit of dierent
types. Some could be further subdivided, especially 3 and 4,
but I wished to keep this list the same as the ve theories of
Darwin from Mayr (1985).
These theories are law-like because they apply to all living
organisms on the Earth although they have been tested only
for a very small number of species. To my knowledge, none of
these nomological theories of evolution failed testing against
the appropriate objective, empirical observations. In a reason-
ably large number of cases, direct observations have been
made of all of these aspects of nomological evolutionary
change, including phyletic evolution in domesticated animals
and plants, development of tolerance to pesticides by many
insects, and development of tolerance to antibiotics by many
bacteria. Phyletic evolution has been observed in the House
Sparrow, Passer domesticus, after its introduction into North
America. This species has spread rapidly over the continent
and dierentiated into numerous local populations or races
(Johnston and Selander 1964; Selander and Johnston 1967).
Speciation has been observed by the re-creation of naturally
occurring as well as novel polyploid species of plants [Appa-
lachian ferns, Asplenium, Wagner 1954; Spartina townsendii
(the only naturally occurring species of plants whose origin has
been observed in the wild), Huskins 1931; the articial radish-
cabbage hybrid Raphanobrassica Stebbins 1950; and many
others, Grant 1963, 1981]. Moreover some breeds of domes-
ticated animals that have become so dierent that they would
act as dierent species under natural conditions. If it was
possible to release a population of giant Irish Wolfhounds and
a population of dwarf Chihuahuas (these breeds were chosen
as the largest and smallest breeds of dogs, but the same would
happen with many other combinations of dog breeds) in a
region where both could survive and reproduce, one will nd
only Irish Wolfhounds and Chihuahuas generation after
generation responding to one another as good species. The
large dierence in size between these two groups of dogs serves
as the intrinsic isolating mechanism for genetic isolation
between these two taxa regardless of whether they are
technically placed in the same species, Canis familiaris, and
regardless of the fact that, with series of intermediate steps
involving dierent breeds of dogs, genetic material could be
transferred from Irish Wolfhounds to Chihuahuas. Moreover
large breeds of dogs and Northern Hemisphere wolves, Canis
lupus, readily interbreed with viable and fertile ospring
although the large and toy breeds of dogs cannot because of
their size dierence, The same would be true if populations of
dwarf horses and the largest dwarf horses were released in a
region where both would survive and breed. If one wishes to
speak of any aspect of evolutionary theory being factual, it is
these direct observations of dierent aspects of nomological
evolutionary theory, not the history of life which remains
theoretical, but exceeding well corroborated with very exten-
sive testing without any falsication.
Only the last of Darwins ve theories as listed by Mayr is a
historical evolutionary theory. This is:
(5) Common descent which implies that all species or popu-
lations of organisms have descended with modication from
common ancestors; this descent includes both modication
and branching. Darwinian common descent is equivalent to
Haeckelian phylogeny. Hennigian phylogeny is equal only to
the branching aspect of Haeckelian phylogeny (Mayr and
Bock 2002). Common descent is expressed in Darwinian
classications, Haeckel dendrograms, Hennigian cladograms,
and the tree of life, all of which are theoretical statements.
Historical biogeography and any other biological theory,
that ts the characteristics of H-N Es outlined above, would
also fall into the class of historical theories. Most important is
that the historical evolutionary theory of common descent, as
well as any other historical biological theory, depends rmly
on a set of appropriate nomological theories to be scientic. In
the case of the historical evolutionary theory of common
descent, it is based on the preceding four nomological
evolutionary theories. Nomological evolutionary theory is
primary and historical evolutionary theory is secondary, and
must be considered and discussed in that order. It is incorrect to
speak rst of the pattern of evolutionary change ( historical
theory) and then the process of evolutionary change
( nomological theory).
The fth of Darwins theories that of common descent is
a general theory and deductions from it can be tested against a
number of objective empirical observations. If an evolutionary
history of living organisms (both fossil and recent) is obtained
from the nomological theory of evolution, then one can reach
three further deductions, namely:
(1) Groups of related organisms will be found non-stochasti-
cally over the surface of the Earthdepending ontheir degree
of evolutionary relationships and their abilities to disperse.
(2) The origins of groups of organisms will be found
chronologically in the fossil record depending on their
ancestral-descendent relationships.
(3) Vestigial structures will be found in descendent groups
with the homologous structure existing well developed in
ancestral groups.
These deductions had been tested successfully against a
exceedingly large number of empirical observations of the
spatial and chronological distributions of organisms and the
occurrence of well developed features in ancestral groups
versus vestigial features in descendent groups. Hence one can
conclude that general historical evolutionary theory is
extremely well corroborated, and that ultra-massive counter
observations would be needed to disprove Darwins historical
theory of common descent. But this well-tested and supported
historical evolutionary theory is still a theory as noted by
almost all philosophers of science and is not a fact or factual as
many evolutionists like to characterize it. The general histor-
ical theory of evolution, although exceedingly well corrobor-
ated, cannot be used for testing any special historical
evolutionary theories and especially for testing any nomolog-
ical theories because historical theories cover singular events;
hence they cannot be used to test other historical events or to
test nomological theories which cover general events.
In addition to the general historical theory of evolution,
endless special theories exist which deal with the evolutionary
history and classication of all groups of organisms at all
hierarchal levels from subspecies to kingdoms. Hence the
theory of whether the Rodentia and the Lagomorpha are
closely related to one another within the Eutheria forming the
taxon Glires, or the theory of whether pinnipeds (seals and
other aquatic carnivores) had a single or a double origin from
the terrestrial carnivores, or whether birds descended from an
early group within the archosaurian reptiles (Martin 2004) or
from a later and more specialized group within the theropod
dinosaurs (Sereno 2004) or perhaps that some of the late
dinosaurs, the maniraptoran radiation, are actually not
dinosaurs but ightless descendants of an early bird such as
Archaeopteryx (Paul 2001, 2002; Martin 2004) are all special
historical evolutionary theories.
Special historical evolutionary theories deal with singular
events and each must be tested independently against objective
empirical observations, which generally include an argument
chain of well corroborated nomological theories and the
observations supporting them. To be scientic, historical
evolutionary theories must also be tested against relevant,
well corroborated nomological evolutionary theories. Any
historical evolutionary theory must also be tested using other
well-tested, non-evolutionary nomological theories. A histor-
ical evolutionary theory on the origin of avian ight must also
100 Bock
be tested with nomological aerodynamic theories as well as
those concerning functional properties of vertebrate muscle-
bone systems, metabolism, respiration, etc. If, nally, a well
corroborated and convincing historical theory has been
reached to explain the evolution of avian ight, this historical
theory works for birds only and cannot be applied to explain
the evolution of ight in bats or in pterosaurs, which are
dierent singular events and may well have very dierent
historical explanations (Bock and Bu hler 1995).
Functional explanations
Although this essay deals with diverse forms of evolution-
ary explanations, a brief word should be said about
functional explanations. Two dierent, but interrelated sys-
tems of explanations exist in biology, which are (1) the
dichotomy of N-D E versus H-DEs and (2) the dichotomy of
Functional Explanations versus Evolutionary Explanations.
The latter system stems from the useful division of biology
into the major areas of functional and evolutionary biology
as noted by Mayr in his Growth of Biological Thought (1982).
He did not use the terms Functional and Evolutionary
Explanations, but his division of biology into these two
major areas can be taken as the basis to consider this
dichotomy of explanations. It should be noted that the use of
functional explanation in biology is sharply dierent from the
use of this term by many philosophers (Nagel 1961); I will
use it strictly in the biological meaning. It should also be
noted that many or most philosophers of biology use the
term function in a historical evolutionary sense which I do
not advocate because I believe that it leads only to confusion.
These two systems of explanations in biology do not have a
simple relationship to one another. All functional explanations
are N-D Es and all H-N Es are evolutionary, but evolutionary
explanations can be either N-DEs or H-N Es. Hence it is
essential not only to characterize carefully the properties of
N-D Es and H-N Es, but to show which parts of biology, and
especially of evolutionary biology, are nomological-deductive
and which are historical-narrative. Nomological evolutionary
theory is primary to historical evolutionary theory. Moreover,
although I cannot provide a complete argument at this time,
functional explanations appear to be primary to evolutionary
explanations. Part of this problem is that good denitions of
the areas of functional and of evolutionary explanations in
biology are not available. These two areas of biology cannot be
characterized simply by the commonly used distinction of
How questions versus Why questions which is too vague to
be of real use.
Conclusions
In discussions inside and outside of science, it is not possible to
talk simply about the theory of evolution and about the proofs
supporting this theory. Nor is it possible to take the position
that the theory of evolution has been so well corroborated and
supported that it can be considered to be factual. Such
approaches simply lead to confusion and failure to reach any
real agreement. Most denitions of biological evolution are at
the same time not suciently broad in that they exclude
several valid forms of change (social evolution and template
evolution) and include an unnecessary criterion that evolu-
tionary change has to be genetic. Further, the theory of
evolution as proposed originally by Darwin and still advocated
today by most people is not a single theory but a bundle of ve
independent theories as originally mentioned by Mayr in 1982
and further elaborated by him in 1985 and repeated in at least
four of his later books (Mayr 1991: 3539; 1997: 177189;
2001: 8687; 2004: 97115). These theories are not of the same
nature in that four are clearly nomological and one common
descent is denitely historical. Philosophers of science have
concentrated almost exclusively on nomological science which
has further confused matters in that biologists have tried to
force all evolutionary explanations into this mold or that,
using the opposite tack, some philosophers of science under
the belief that evolution is only historical, have declared
evolution theory not scientic. Nomological and historical
scientic theories are markedly dierent in their expression,
what they cover and how they are tested; all most be tested
against objective empirical observations. None can be proved
and none are factual.
Historical evolutionary theories must be based on the
appropriate nomological evolutionary theory to be scientic.
Classications, phylogenies, dendrograms, clarications and
all other explanations of the evolutionary history of groups of
organisms and of their particular attributes, including distri-
bution both in time and space, are historical theories; these
explanations must be presented and tested in a very dierent
way than the standard nomological theories in science. As for
all historical theories, evolutionary historical theories and
explanations must be presented and tested realistically with a
sensible estimate of the degree of condence that one can have
in the particular explanation. The testing of historical theories
against empirical explanations is a far more dicult task than
realized by most biologists, and more historical theories enjoy
a far lower degree of condence than most biologists estimate
(Bock 2004a).
Dialogues between people accepting evolution and those
objecting to evolutionary explanations are almost entirely
limited to historical evolutionary theory. Most anti-evolution-
ists simply do not concern themselves with nomological
evolutionary theory. Many are quite comfortable with possible
evolutionary change within the species limit with their objec-
tions being largely to the idea of evolution beyond the species.
No resolution is possible using many of the traditional
arguments supporting evolution theory as advanced by most
biologists, and quite possibility no resolution will ever be
possible. But if evolutionary biologists and philosophers of
science present clearly and completely evolutionary theory in
all of its fascinating diversity, then at least we know where the
arguments are and which views are based on science and which
on belief.
Acknowledgements
First and foremost, I would like to thank Professor Ernst Mayr who
for over 50 years encouraged me to think critically and not to accept
automatically statements in the literature, including his, no matter how
broadly they are accepted by other workers. I would also like to thank
the late Professor Ernest Nagel, Hans Winkler, Professors Frederick E.
Warburton, Gerd von Wahlert and Arthur L. Caplan with whom I had
numerous discussions on evolution theory and the philosophy of
science. And I thank Gerd von Wahlert for translating the abstract
into German. Lastly I would like to express my appreciation to the
many students in my courses in Introduction to Evolutionary Biology
and in the Philosophy and History of Evolutionary Biology for their
discussions and often embarrassing questions which led to a cleared
understanding of the ideas presented herein.
Zusammenfassung
Erklarungen zur Evolutionstheorie
Die Evolutionstheorie wird auf vielfache Weise deniert, was zu einem
erheblichen Durcheinander bei ihrer U
berpru fung bei Anwendung auf

empirische Beobachtungen fu hrt. Evolutive A
nderungen werden u bli-

cherweise als genetisch deniert, was sowohl die kulturelle wie die
epigenetische Evolution ausschliet; also sollte das qualizierende
Adjektiv ,genetisch in der Denition der biologischen Evolution nicht
verwendet werden. Darwins Theorie, die von ihm immer im Singular
beschrieben wird, ist tatsa chlich ein Bu ndel von fu nf unabha ngigen
Theorien, wie das Mayr vorgetragen hat. U
berdies ist nur eine dieser

Theorien, na mlich die der gemeinsamen Abstammung, historisch, und
die anderen vier sind nomologische Aussagen [u ber Evolution selbst,
u ber den Gradualismus, u ber den Verlauf der phyletischen Evolution
und der Artbildung und u ber Evolutionsursachen]. Also ist nicht die
Theorie der Evolution insgesamt historisch. Bei biologischen Verglei-
chen muss zwischen horizontal und vertikal unterschieden werden, und
gu ltige Schlu sse bei einemTyp von Vergleich ko nnen nicht automatisch
auf einen des anderen Typs u bertragen werden. Keine evolutive
A
nderung, wie drastisch sie auch sei, u berschreitet die Grenze eines
Taxons; deshalb ist es unmo glich, eine ,transspezische Evolution
(Evolution u ber die Artgrenze hinweg) von der innerhalb einer Art zu
unterscheiden. Makroevolution unterscheidet sich nicht von der
Mikroevolution mit Ausnahme des Ausmaes der A
nderungen: Es gibt
keine eigenen Ursachen oder Ablaufsformen fu r die Makroevolution.
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