Palaeogeography, Palaeoclimatology, Palaeoecology 176 (2001) 1^10

www.elsevier.com/locate/palaeo

Palaeotrends

The structure of species, outcomes of speciation and the

`species problem': ideas for paleobiology
William Miller III
Geology Department, Humboldt State University, Arcata, CA 95521, USA
Received 25 July 2001; received in revised form 23 August 2001; accepted 23 August 2001

Abstract
Paleobiologists reviewing the recent literature on species concepts are likely to come away with the impression that
the fundamental unit of biodiversity is impossible to define in terms of a universal concept that applies to all forms of
cellular life and the varied outcomes of species-level evolution. Operational concepts are likely to differ from specialty to
specialty in biology, but a general theory of species has been available for nearly 50 yr in the form of G.G. Simpson's
evolutionary species concept. In addition to this, a general picture of species structure has been accepted for at least as
long, consisting of demes or habitat clusters that bud off other demes/clusters, split, fuse and often go extinct. More
recently, it has become obvious that the outcomes or products of species-level evolution may include any of the
following: speciation with morphologic differentiation (producing the so-called `good species' of applied biology and
probably the stable entities resolved in patterns of stasis in the fossil record); speciation without pronounced phenotypic
differentiation (cryptic species); or acquisition of diagnosable characters without either complete reproductive isolation
or pronounced morphologic change. The view that species are lineages containing networks of demes (or habitat
clusters of uniparental organisms), each with its own biologic tendencies and role ^ with a unique place within a clade
composed of similar historical entities, each having a certain kind of internal structure and resulting from a different
evolutionary outcome ^ is the ultimate concept many of the operational approaches seem to be aiming to uncover.
Considering species structure, speciation outcomes and the evolutionary species concept together brings the nature of
species into a much sharper focus. 2001 Elsevier Science B.V. All rights reserved.
Keywords: species-level evolution; species concepts; internal structure; evolutionary outcomes; historical entities; ontology

``If we accept the assumption of most systematists and evolutionists that species are real things in
nature, and if the sets of species specied by dierent concepts do not overlap, then it is reasonable to
conclude that real entities of the world are being
confused.''
Cracraft, 2000

``The problem is that no single denition of the

species category has proved optimal for all of its
dierent uses. Consequently, although one denition or class of denitions has often come to be
favored for a certain period of time or by a certain
group of biologists, none of them has enjoyed universal endorsement within biology as a whole. This
situation has come to be known as `the species
problem'.''
de Queiroz, 1998

E-mail address: wm1@axe.humboldt.edu (W. Miller III).

0031-0182 / 01 / $ ^ see front matter 2001 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 1 - 0 1 8 2 ( 0 1 ) 0 0 3 4 6 - 7

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``Theory neutral concepts are impossible, but

even if they were possible, they would be undesirable.''
Hull, 1997
1. Introduction
Sampling the vast literature on species concepts
probably leaves most readers with the unsettling
feeling that one of the central issues or core concepts of biology ^ the nature of species as the
fundamental units of biodiversity and evolutionary patterns ^ is becoming hopelessly confused by
a seemingly endless proliferation of ideas. The
more pragmatic readers have always investigated
the various proposals and, like discriminating
shoppers, selected the versions that best suited
their favorite group of organisms or research programs. My reaction to the so-called `species problem' is entirely dierent: I think the problem has
been solved, or nearly so, and that the ongoing
debate is really about ironing out details.
There are three reasons for my reaction. First,
we have had a realistic picture of the internal
organization of species for at least 50 yr (really
since Wright's (e.g., 1932) early contributions in
population genetics): we know that species usually consist of multiple populations geographically
and temporally separated from one another; that
these subdivisions bud o from other populations,
divide, fuse or go extinct; and therefore the structure of species is like the bundle of bers making
up a hemp rope (G.G. Simpson's (1953, gure 48)
classic line-drawing is the basic model). Second,
the `conceptual space' of possible species concepts
already has been rather thoroughly explored. The
major categories of concept include those that
stress reproductive isolation, recognition or cohesion (integration of the genotype) ; those that view
species as historical entities or lineages that can be
objectively identied as being more or less related
to other such entities (phylogenetic units); and
those that stress some form of compositional
(phenotypic, genetic) similarity or separation
without paying too much attention to evolution
theory (Hull, 1997). Any future proposal of a new
concept is likely to be a variation or expansion of

one of these three major themes. And third, the

outcomes or products of species-level evolution
are fairly well known, and include speciation
with or without pronounced morphologic dierentiation (Vrba, 1980).
In this essay I will argue that a universal concept of species, one that could potentially include
all kinds of cellular life and evolutionary products, is within view and depends on the three developments mentioned above. This is not to say
that all the problems associated with the species
category have been settled. It is still unclear, in
some important practical cases, how to accommodate uniparental or asexual organisms, the outcomes of hybridization (two problems that have
always dogged the neobiology community), and
the results of apparently transspecic anagenetic
evolution (a problem close to home for paleobiologists). I will begin with a brief review of the large
amount of recent work on species concepts, then
illustrate what I call the structure of species, relate
this to outcomes or products of species-level evolution, and end by claiming that the most realistic
species theory ^ not the most eectual or universal operational concept (see the recent discussions
by de Queiroz (1999) and Hey (2001)) ^ has been
available for 50 yr.
2. Progress in the conceptualization of species
To begin to understand what the phylogenetic
systematists and evolution theorists are discussing
these days, paleobiologists will need to do some
homework. In order to come up to full speed
without getting lost in the scattered literature, I
recommend the comprehensive (and surprisingly
readable) summary of species concepts by Mayden (1997) as the starting place. Mayden reviewed
25 more or less dierent concepts, attempted to
synonymize some of the concepts, and used direct
quotes of the denitions proposed by various authors, producing a handy identication manual of
species concepts. The briefer reviews by Harrison
(1998) and de Queiroz (1998) also are useful.
After a sti dose of denitions (Table 1 lists the
practical concepts most often mentioned in summaries), I recommend some critical reection in

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Table 1
Denitions or characterizations of the most prominent operational species concepts related mostly to sexually reproducing organisms (based on Mayden, 1997; Harrison, 1998; de Queiroz, 1998; Wheeler and Meier, 2000)
Concepts related to reproductive isolation, internal cohesion or species recognition (e.g., isolation and maintance of reproductive
networks)
Reproductive isolation
`I dene biological species as groups of interbreeding natural populations that are reproductively isolated from other such groups.
Alternatively, one can say that a biological species is a reproductively cohesive assemblage of populations.'
(Mayr, 2000, p.17)
Cohesion
`The cohesion concept species is the most inclusive population of individuals having the potential for phenotypic cohesion
through intristic mechanismsT'
(Templeton, 1989, p.12)
Recognition
`A species is that most inclusive population of inidvidual, biparental organisms which share a common fertilization system.'
(Paterson, 1993, p.105)
Concepts depending on the properties of identied lineages (e.g., the various concepts within phylogenetic systematics)
Hennigian
`Species are reproductively isolated natural populations or groups of natural populations. They originate via the dissolution of
the stem species in a speciation event and cease to exist either through extinction or speciation.'
(Meier and Willmann, 2000, p.31)
Diagnosable
`We dene species as the smallest aggregation of (sexual) populations or (asexual) lineages diagnosable by a unique combination
of character states.'
(Wheeler and Platnick, 2000, p.58)
Monophyletic
`A species is the least inclusive taxon recognized in a formal phylogenetic classication. As with all hierarchical levels of taxa in
such a classication, organisms are grouped into species because of evidence of monophyly.'
(Mishler and Theriot, 2000, p.46-47)
Concepts based largely on compositional properties of specimens
Morphologic
A group of organisms delineated from other such groups based on possession of `essential morphologic attributes'.
(Mayden, 1997, p.403)
Phenetic
`Operationally, where variation in a set of characters is less within a group than between groups the entitiy is recognized as a
distinct taxon.'
(Mayden, 1997, p.404)
Genetic
`This concept is similar to the morphological species concept except that the method used to delineate species is a measure of genetic dierencesT independence is assessed using methods varying from chromatography, to protein electrophoresis, to sequencing.'
(Mayden, 1997, p.399)

the form of two recent essays by Hull (1997;

1999). Stocked with possible concepts and knowing something about what the philosophers of biology are saying to the apparent inability of biologists to settle on a single concept for the basic
unit of biodiversity, it would be time to plunge
into the freshest controversies surrounding these
issues by reading the extended debate by some of
the most visible participants, edited by Wheeler
and Meier (2000).
Going that far, readers are likely to make some

of the same generalizations that I have made in

the following list:
b Concepts stressing reproductive isolation, cohesion or recognition are now seen by an increasing number of workers as secondary attributes of
potentially more robust concepts, or simply to be
limited to certain kinds of (bisexual, biparental)
organisms.
b Most recent concepts view species as real
things in nature, not subjectively delineated clusters of specimens, with beginnings (speciation),

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Fig. 1. The possible internal structures of species-lineages. In general, species consist of demes or habitat clusters, and comprise
clades. A) A lineage consisting of a bundle of demes that oscillate through the phenotypic/habitat space of an established species
but does not ll up the space, as in the other models. A variation of this structure involving the splitting and fusion of the bundles of lineages could be used to represent geographic races or population groups. B, D: Reminiscent of Simpson's (1953, gure
48) line-drawing of species structure, in which demes explore most of the phenotypic/habitat possibilities of a species or dene
those possibilities through time. The component demes bud o from other demes, fuse or go extinct. B) consists of many demes
packed into the space of the species, so that phenotypic distance between neighboring demes is not pronounced; D) features
demes having greater phenotypic separation (dierent traits associated with dierent habitats). C) represents a single, well-mixed
population lling essentially the entire phenotypic/habitat space of an established species. Lineages that `escape' from the boundaries of adjacent cylinders and fuse could be used to represent hybrids. The phylogenetic tree to the right gives a sense of the
upper level of organization (species-lineages 1^3 within a clade). Each of the cylinders represents an idealized internal structure.
Small crosses are demic or habitat cluster extinctions. Speciation associated with pronounced phenotypic dierentiation (suggesting adaptive species-level evolution) is indicated with sm; speciation without this kind of dierentiation is indicated with s. Species-lineage boundaries are idealized as circular cylinders to emphasize prevalence of stasis; boundaries could be the result of external controls or internal constraints.

individual histories and ends of some kind (extinction).

b Many workers, but not all, can visualize species as the level of biologic organization at the
transition from dominance of tokogenesis to the
dominance of phylogenesis (i.e., species undergo
mostly cladogenesis, but their component populations have reticulated patterns of relationship
through time).
b Patterns of descent are paramount; phenetic
patterns are not always the most reliable sources
of information in phylogenetic reconstructions.
b Most of the noise in the debate about what
species are comes from inside the phylogenetic
systematists' camp and involves operational issues
such as whether or not ancestral species survive
speciation events, the signicance and interpretation of apomorphy and monophyly, and whether

or not theory should color the application of analytic methods.

b There appears to be no consensus about
whether a universal (or monistic) species concept
is available, or if pluralism is the best bet
(although most participants in the debate probably hold out hope for a universal theory of species).
Having said all of this, most applied biologists
(neoecologists and paleoecologists, biogeographers, biostratigraphers, conservation biologists)
would still say that a `good species' ought to be
reproductively isolated from other species and
that morphologic (really compositional) dierences can be used to diagnose or recognize dierent
species. And most theorists would agree that asexual or agamospecies, hybrid-forming species complexes or syngameons, and `chronospecies' (mor-

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Fig. 2. Possible outcomes of species-level evolution. The matrix shown in (A) is based on Vrba's diagram (Vrba, 1980, gure 5),
and scores morphologic dierentiation against speciation. Most paleobiologists would agree that a `good' species ought to be differentiated and isolated (black square), and presume that cryptic species (gray square) are insignicant. A more comprehensive
picture is shown in (B) in which the additional complication of diagnosable characters has been added. In this view of outcomes,
the black cube is equivalent to isolated, dierentiated species-lineages produced when speciation and adaptive phenotypic transformation are associated (sm in Fig. 1). Many other possibilities are possible, including isolation without dierentiation or acquisition of new character states without complete isolation (gray cubes; represented by s in Fig. 1).

phologically dierentiated anagenetic species) still

are the sources of unresolved problems.
At this point I can oer two pieces of advice.
Paleobiologists obviously need to test the waters
and determine if their notion of species needs adjustment to accommodate the recent thinking outlined above. More importantly, generalizations
about causation will always contain inherent
weaknesses if the issue of ontology is never adequately addressed. What follows is an attempt to
clarify the ontology^causation linkage.
3. The structure of species and species-level
evolution
Simpson ^ a paleontologist with the time dimension clearly in mind ^ portrayed the internal
structure of a species-lineage (Simpson, 1953, gure 48) as a bundle of anastomosing bers or
strands (representing demes drawn through
time), which in turn consist of trellis-like networks
of sexually reproducing individual organisms. The
purpose was to illustrate species as historical entities, as opposed to non-dimensional clusters of
similar specimens or isolated groups of organisms,

and to emphasize ``Tthat joining and anastomosis

in phylogeny concern (major features of evolution) only to the extent that they are part of the
intimate mechanism of evolution which does rise
to levels where joining no longer occurs'' (p. 378).
I have used the same approach in Fig. 1 to attempt to illustrate the possible range of internal
structures of species-lineages, which is based on
the same kind of hierarchical thinking. By comparison, the lineage structure diagrams used in
recent papers by phylogenetic systematists resemble more closely the structural model illustrated
by Hennig (1966, gures 4 and 6) in leaving out
the strands of demic systems and only showing
networks of individuals within the lineages.
Nevertheless, the general purpose is the same: to
begin to understand speciation one must understand what species are, and to do that one must
be able to visualize their internal structure.
Each of the lineage segments in Fig. 1 represents possible demic or population system structure within an established biparental species over
perhaps 103 to 104 generations (or if the species is
uniparental, possibly 10 to 102 iterations of habitat separation). The individual strands are demes
separated in terms of phenotypic variation that

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corresponds to habitat separation. Boundaries of

the species-lineages are idealized ; they could have
been depicted as meandering instead of straightsided cylinders to emphasize the moving average
of phenotypic properties of the species, or they
could have been drawn having alternating wide
and narrow sections indicating changes in the
maximum variation in phenotypic properties or
habitat variation through time. I have used the
vertical boundaries to emphasize overall specieslineage stability, which is realistic considering the
prevalence of stasis in the fossil record (Jackson
and Cheetham, 1999; Jablonski, 2000; Benton
and Pearson, 2001). Demic systems bud o from
established systems; drift or undergo directional,
stabilizing or disruptive selection; fuse with other
demes; or experience local extinction. Demes consisting of biparental organisms are in potential
reproductive connection within the cylinders, but
have potentially severed the connection or at least
have acquired a unique diagnosable character
when they cross the boundary (begin to`escape'
from the cylinder or domain of the established
species). For uniparental organisms, the representation of `demes' as separate strands also represents spatial or habitat separation. Fusion would
occur when these separate clusters are reunited in
a mixed habitat cluster or possibly owing to `lateral' genetic exchanges (Templeton, 1989 Margulis, 1993; Sapp, 1994); extinction would occur
when a local cluster or variant is eliminated.
Crossing the boundary would again represent incipient speciation by acquisition of new characters
and potentially permanent habitat separation.
Several species structures are possible. In Fig. 1,
structures B and D, consisting of reticulated
demes and many instances of local extinction of
populations/variants, are probably typical for animal and plant species deployed in multiple populations. The internal structure shown in A, involving a sinuous, vine-like pattern of demes,
might result from environmental pacing or developmental changes in regional ecosystems, with the
domain of the species-lineage remaining essentially xed (dened by the potential range of phenotypes of population systems). Structure C is
supposed to represent one large `deme' lling
the potential phenotypic/habitat space of an es-

tablished species without signicant internal discontinuity. This might occur in some pelagic species (but see the recent review by Norris (2000)
suggesting previously undetected population
structure and dierentiation in pelagic systems)
or in well-mixed cultures of micro-organisms.
Demes of biparental organisms or habitat clusters of phenotypically identical uniparental clones
are mainly conned to cylinders because the cross
sections represent the range of (externally or internally controlled) conditions in which survival
and tness are supported (see the discussion of
`cohesion mechanisms' in Templeton, 1989);
crossing the boundary amounts to relinquishing
a stable association with environmental and ecologic factors or exceeding some form of cohesion
(small arrows marked with s). A thorough discussion of processes occurring at the boundary ^
where speciation begins ^ can be found in Futuyma's (1986) excellent book. Rarely, and perhaps
only when new adaptive opportunities occur, will
these excursions result in a speciation event that
coincides with morphologic (adaptive) dierentiation (marked with sm in the phylogenetic tree). In
other words, the escape of population systems
from established species, or the `attempt' to escape, could be a fairly common phenomenon
(Williams, 1992), but the successful establishment
of isolated, dierentiated new species-lineages
would have to coincide with the provision of environmental or ecologic opportunity (Eldredge,
1995; Miller, in press). The resulting entities are
expected to become the essentially static specieslineages emphasized in the theory of punctuated
equilibria (Eldredge and Gould, 1972; Gould and
Eldredge, 1993).
The possible outcomes of species-level evolution were depicted by Vrba (1980, gure 5), in
terms of speciation with or without morphologic
dierentiation (Fig. 2A). These model outcomes
have been discussed recently by Eldredge (1989,
1995). I have expanded this generalization to include acquisition of a diagnosable character,
which may or may not coincide with complete
reproductive isolation and pronounced morphologic dierentiation (Fig. 2B). Is there any species
concept that could accommodate all of these potential structures, outcomes and complications?

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4. The `species problem' has been solved ! (around

50 yr ago)
We know that the internal structure of specieslineages must look something like one of the models in Fig. 1, perhaps typically like versions B and
D that recall the diagram used by Simpson (1953)
to help visualize the same pattern. Species usually
consist of multiple demes or habitat clusters of
some sort, which change through time, and in
turn comprise clades of genealogically related lineages (Eldredge, 1985; 1989). Species-level evolution could involve any of the outcomes depicted
in Fig. 2B. At the same time, there is an increasing realization that species concepts based on operational criteria (gene ow, tests of lineage relatedness, purely compositional attributes) are
secondary, empirical approaches ^ they do not
take us all the way to the desired goal of a universal concept, one that encompasses all evolutionary outcomes in all kinds of cellular life.
Many systematists and evolution theorists picture the work on the `species problem' as a long,
hard march that will take us to the ultimate vantagepoint, and there we will see what species
really are like. In fact, the ultimate picture was
painted decades ago in the form of the theoretical
evolutionary species concept (Simpson, 1951;
1961). The expanding work on operational concepts is extremely signicant, but from a theoretical point of view it all amounts to clearing multiple paths toward the same goal (compare this
with the similar assessment oered by de Queiroz
(1999)). Recent advocates (e.g., Wiley and Mayden, 2000) of the evolutionary concept describe it
as if it were a new solution to a old problem when
it has been with us since Simpson was attempting
to draw the broadest outlines of species and speciation. The most widely quoted version is from
his Principles of Animal Taxonomy (Simpson,
1961, p. 153): ``An evolutionary species is a lineage (an ancestral-descendant sequence of populations) evolving separately from others and with its
own unitary evolutionary role and tendencies.''
Critics of the concept point to vagueness of the
terms `role' and `tendency', and declare it useless
as an operational approach. Proponents emphasize that the concept accommodates all kinds of

organisms and provides an `ultimate picture' (not

an operational concept) ^ a conceptual goal or
necessary adjustment in the core concept ^ of species as historical entities (Mayden, 1997; Wiley
and Mayden, 2000).
The possible structure of species, as depicted in
Fig. 1, is clearly compatible with this well-known
denition of species. How could all the possible
outcomes of species-level transformation (Fig. 2B)
be accommodated within the concept? The establishment of a new species probably involves something like the following chain of events: (1) appearance of an innovation (new character state,
either as minor as a new protein structure or as
major as a new anatomic feature, developmental
variant, behavioral pattern or physiologic process)
at the demic level; (2) isolation or separation to
conserve or protect the innovation (incipient speciation) or simply geographic separation to keep
the new character from disappearing owing to
interbreeding; and in some cases (3) signicant
morphologic dierentiation (adaptive transformation) matching a resource opportunity or new
ecologic position. When the sequence includes
these steps, most paleobiologists would agree
that a new species-lineage has been established.
The new species really emerges at the interface
of tokogenetic processes and phylogenesis ^
when the cylinder boundary has been crossed. Appearance of a new character state without protective reproductive isolation (for biparental organisms)/habitat separation (for asexuals) would
leave the new lineage vulnerable when contact is
made with the ancestral lineage (by interbreeding
(biparentals) if isolation has not been achieved;
by competitive exclusion at the initiation of reproductive isolation/habitat separation) (Futuyma,
1986; 1987). After establishment, intraspecic anagenetic evolution might result in the honing of
phenotypic properties to environmental or ecologic windfall, or the new species-lineage might
emerge fully outtted for the opportunity or
role. In any case, if the steps produce a new, independent lineage, Simpson's concept would apply.
The rst two steps described above could occur
essentially simultaneously, or in reversed order. It
is also possible that reproductive isolation could

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occur without transformation of any other character states except those producing or enforcing
isolation. The result could be cryptic species that
resemble very closely ancestral or sister species in
terms of phenotypic properties. Traditional morphologic approaches to species delineation and
identication, in both neo- and paleobiology, are
likely to overlook these products of species-level
evolution (see discussion in Knowlton and Weigt,
1997).
A dierent set of circumstances might produce
a spatially separated lineage having a single diagnosable character, which could be maintained for
long intervals of time without complete reproductive isolation (see Cracraft, 1989). Because this
lineage would have `escaped' the domain of the
ancestral species-lineage (a cylinder boundary in
Fig. 1), it too could be accommodated in the evolutionary species concept. Prolonged separation
would result inevitably in reproductive isolation
in biparentals (Turelli et al., 2001).
Strongly dierentiated new lineages are special
cases of outcome in which a non-trivial economic
opportunity or role has been `discovered' or `captured' by the new species. These are probably the
stable lineages preserved in the fossil record as
patterns of species-lineage stasis. Species never
conquering a resource or capturing a signicant
economic position may remain rare or vanish
soon after they emerge (see Stanley, 1979; Eldredge, 1995; McKinney et al., 1996), but they
would still qualify as evolutionary species. Thus,
cellular life is composed of economically prominent `haves' and the less conspicuous `have-nots',
both qualifying as good evolutionary species. Any
number of operational approaches could be used
to discover them.
5. Conclusions
Debates about what makes a species a species,
both from an operational and a theoretical point
of view, have always been at the heart of understanding the processes and products of speciation :
the issues of ontology and causation ^ as in many
other biologic problems ^ are inseparable. Paleobiologists need to pay close attention to recent de-

velopments surrounding the species problem, and

to re-examine their own ideas about species concepts to incorporate the recent thinking outlined
above. This is especially true in light of the widespread adoption of cladistic methods and the visualization of species as historical entities (lineages).
There are interesting theoretical issues brewing
that relate directly to the problem of ontology.
Not all species are born equal: some speciation
events produce new lineages that become the
dominant players (the source of many local populations) in regional economies, while others produce new species that never conquer a resource or
acquire a prominent ecologic position. The former
are expected to be fully dierentiated in phenotypic characters compared to ancestral species
(i.e., to feature new adaptations) ; the latter may
never achieve such dierentiation. Both, however,
are kinds of species-lineages and are accommodated in the evolutionary species concept. Lineages of organisms that feature a few diagnosable
characters, eectively separated from the ancestral
lineage preventing interbreeding but not achieving
reproductive isolation, also are accommodated in
the concept. The relative frequency of these outcomes in various groups of organisms is not well
documented, although typical occurrence of
strong morphologic dierentiation during speciation as opposed to frequent formation of cryptic
species seems to be a more common event in some
groups than in others (compare the assessments of
late Cenozoic bryozoans in which speciation and
morphologic dierentiation apparently go hand in
hand (Jackson and Cheetham, 1990) and modern
birds with possibly many cryptic species (Graham,
1996)). Uniparental organisms are likewise accommodated if descendent lineages are at least
minimally dierentiated from ancestors and
achieve habitat separation, so that their species
structure resembles that of biparental species.
The signicance of chronospecies (anagenetic lineage segments interpreted as separate species by
some taxonomists) may have been exaggerated in
paleontology; higher resolution records of speciation would probably reveal some form of cladogenesis as the dominant pattern in groups supposed to contain chronospecies (e.g., Norris,
2000).

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Based upon what we now know about the internal structure of species, the hierarchical relationships of genealogic entities, and the possible
outcomes of species-level evolution, a broadly acceptable picture of species is within view, and has,
in fact, been available as a formal denition since
Simpson (1951; 1953; 1961) grappled with these
same issues nearly a half century ago. Viewing the
`species problem' in terms of possible structure
and speciation outcomes brings the picture of species as real things in nature into a much sharper
focus.
Acknowledgements
This essay resulted from involvement in the
Ecological Processes and Evolutionary Rates
Working Group supported by the National Center
for Ecological Analysis and Synthesis (Santa
Barbara, California), a center funded by the
National Science Foundation (Grant #DEB-9421535), the University of California at Santa
Barbara and the State of California. I am grateful
to Mike Benton and Niles Eldredge for their
reviews.
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