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Initial work on this volume was supportedby SSHRCgrant number 410-960497 awarded by the Social Sciencesand Humanities ResearchCouncil of
'
Canadawhile I was at Queens University ; I completedthe volume with the
assistanceof a ResearchBoard Grant Horn the University of Illinois, UrbanaChampaign. I would like to thank Allison Dawe for her researchassistance
with the project in its early stages; Peter Asaro fQr his assistancein its latter
stages; and Unda May for her masterly, efficient, and patient work in formatting
the manuscript.
Although it should almost gQ without saying, I would especially like to
thank all the contributors to the volume, not simply for collectively being
the sinequa non for the volume, but for making the editorial aspectsof the
project less chorelike than they might have been. (SometimesI even had the
illusion that I was having fun.) Apart Horn providing the essaysthemselves,
'
many contributors also read and offered comments on other contributors
essaysand gave generaladvice, as needed, about the constitution of the volume
as a whole.
Contributors
1997) and, with Kim Sterelny, Sexand Death: An Introductionto the Philosophy
of Biology(University of Chicago Press, 1999).
David L. Hull is the Dressier Professor in the HumaniHes in Weinberg
College at Northwestern University, Evanston, Illinois. He is the author of
Darwin and His Critics (Harvard University Press, 1973; reprinted by the
University of Chicago Press, 1983), Philosophyof BiologicalScience(PrenticeHall, 1974), Scienceas a Process(University of Chicago Press, 1988), The
Metaphysicsof Evolution (State University of New York Press, 1989), and
Science
and Selection(Cambridge University Press, 1998). He is editor of the
seriesScience
and Its ConceptualFoundations
at University of Chicago Press.
Frank C. Keil is a professor in the Department of Psychology at Yale University
. He taught for twenty years at Cornell University, is the author of
Semanticand ConceptualDevelopment
: An Ontological Perspective(Harvard
, Kinds, and CognitiveDevelopment
University Press, 1979) and Concepts
( MIT
Press, 1989), and is the coeditor, with Robert Wilson, of TheMIT Encyclopedia
of the CognitiveSciences
(MIT Press, 1999).
Brent D . Mishler is a professor in the Department of Integrative Biology
and Director of the University and Jepson Herbaria at the University of
California, Berkeley. He receivedhis PhiD. from Harvard University in 1984
and was in the faculty at Duke University unHl1993 , when he moved to UC
Berkeley. He is a systemaHst specializing in mossesand has thus grappled
with speciesboth in practice and in theory. He has published extensively on
species concepts as well as on phylogeneHcs at many scales, from moss
speciesto the overall relaHonshipsof the green plants.
David L. Nanney becamea professor in the School of Life Scienceat the
University of Illinois in Urbana-Champaignin 1959. He becamean emeritus
professor in the Department of Ecology, Ethology, and Evolution in 1991.
He receivedhis graduatetraining under Tracy Sonnebornat Indiana University
(PhiD., 1951) and taught at the University of Michigan before moving
to Illinois. His studieshave focusedon the domesticationof the ciliated protozoan
Tetrahymenathermophiiaas a laboratory instrument and have included
.
genetic, developmental, and evoluHonary mechanisms
Daniel C. Richard son is a graduate student in the Department of Psychology
at Cornell University .
Kim Sterelny is an Australian now living in exile in New Zealand. StarHng
as a philosopher of language and mind, he has become increasingly interested
in the philosophy of biology . He retains his original interests, but now
~ th a strongly evolutionary spin. He is coauthor, with Paul Griffiths, of Sex
' and Death: An Introductionto the Philosophyof Biology, due to appearwith the
. University of Chicago Pressin 1999.
Contri~ ors
Robert A . Wilson is an associateprofessor in the Department of Philosophy and a member of the Cognitive ScienceGroup at the BeckmanInstitute
at the University of Illinois, Urbana-Champaign. He taught previously at
'
Queens University , Canada. He is the author of CartesianPsychologyand
of the Mind (Cambridge University
PhysicalMinds: Individualismand the Sciences
Press, 1995), and is the general editor, with Frank Keil, of TheMIT
( MIT Press, 1999).
of the CognitiveSciences
Encyclopedia
Introduction
to the history of biology , they are often able to appeal to that history in
order to enrich our understanding of speciesand the biological world. The
psychologicalperspectiveis most explicit in the essaysby Atran and by Keil
and Richardson, but also underlies central argumentsin severalother papers
(e.g ., by Wilson and by Griffiths). Together, thesetwo featuresof the volume
provide for a broad perspectiveon speciesand on the issuesin the philosophy of biology and in biology proper to which speciesare central.
The papers have been organized into five sections that seemed to me
to represent the most cohesive clusters of views and the most interesting
sequenceof papers to read from beginning to end. Those sections are:
"
Monism , Pluralism, Unity and Diversity " ; " Speciesand Life' s Complications
" "
" "
; Rethinking Natural Kinds ; Speciesin Mind and Culture" ; and
"
SpeciesBegone!" The rest of this introduction mainly provides an overview
of the papers in the order that they appear. There are, of course, other
thematic commonalities, shared perspectives, and oppositions that this
organization (or any single artifactual classificationscheme, suchas a table of
contents) will obscure. One alternative way of thematically locating particular
papersin the volume and of viewing the orientation of the volume as a
whole is to considerthe five themesthat authors were invited to addressand
the pair of themeseachpaper concentrateson most intensely. Those themes,
ranked in order '~ m those that feature in the highest number of papers to
those that feature in the smallestnumber, together with someaccompanying
questions, are:
1
. Given the
Species Realism
3 Pradicallmport
In what ways are answers to the questions asked under the other four
themes important for the practice of evolutionary biology and related
/
Introduction
sciences
? Should we view the resolution of the cluster of issuesoften called
lIthe species problem" as foundational in some way? To what extent is
the speciesproblem (merely) definitional? What is the relationship between
the speciesproblem and empirical practice within the bi.ological(and other)
sciences
? [Hull , de Queiroz, NanneyI SterelnyI Griffiths, Mishler]
4
Historical
Dimensions
In what ways are the views of major historical figures or movementsin evolutionary
biology of significancefor contemporary views of species1Is our
own view of important historical episodes(e.g., formation of the Linnaean
hierarchy, the Modem Synthesis) skewed in important ways1 How can we
shedlight on contemporarydiscussionsby reflecting on the recent history of
evolutionary biology1 [ Nanney, Ereshefsky]
5 Cognitive Underpinnings
To what extent do the literature on children' s naive biology and anthropological work in cross-cultural psychology support nativist and universalist
views of species1What fruitful interplay exists between explorations of the
mental representationof biological knowledge and the philosophy of biology
as it has been traditionally circumscribed1[Atran , Keil and Richardson]
The summariesof the sectionsand essaysindicate that many other issues
are raised in Species
: New InterdisciplinaryEssays
, including the plausibility of
the individuality thesis about species
the
death
of essentialism
,
, the interplay
between ecology and evolution, the relationship betweencommon senseand
scientific taxonomies, and the challenge that recent developmental systems
: New
theory posesto taxonomy in terms of evolutionary homologies. Species
all
.
advances
debate
about
of
these
issues
Between
the
InterdisciplinaryEssays
overviews of contemporary debates and the novel insights provided in
many of the essays, the volume should prove invaluable for professionals
working in the contributing fields and useful for advanced undergraduate
and graduate coursesin either the foundations of evolutionary biology or
the philosophy of biology .
Let me turn more directly to the individual essaysand the sections into
"
which they are organized, beginning with Monism , Pluralism, Unity , and
"
Diversity , containing papers by John Dupre, David Hull , and Kevin de
"
Queiroz. As the title of his essay ( On the Impossibility of a Monistic
"
Account of Species
) suggests, Dupre argues for the rejection of monism
bout
.
He
claims
, moreover, that this conclusionis the proper one to
species
~
draw from the completeassimilationof Darwin' s insights about the organizati9n of the biological world . There are no perfectly sharpboundariesbetween
preexisting natural kinds species that would allow for a monistic account
of species. Rather, what we find when we investigate the biological world is
/
Introduction
/
Introduction
Introdu~
Intr~
tion
'
organization in the biological world that doesnt distinguish the Linnaean
speciesand genus categories; he proposes a domain specific representation
-basedhabits of the
of this category and explores its relationship to essence
mind and the cultural development of various speciesconceptsin Western
science.Atran concludeshis paper with somethoughts about recent views of
pluralism and speciesand about what these views imply about the relation
betweencommon senseand science.
Frank Keil and Daniel Richardson discuss the psychological representation
of speciesand of biological knowledge more generally in their essay,
"
, Stuff, and Patterns of Causation." They argue that the substantial
Species
developmentalliterature on biological knowledge often presentsamisleading
conception of what intuitive or folkbiology must be like in order for
speciesand other biological categoriesto have the distinctive psychological
featuresthat they do, suggesting severalnew lines of empirical research. By
"
"
exploring what has been called psychological essentialism about biological
kinds and its relationship to essentialismin the philosophy of biology , Keil
and Richardson call for more careful empirical examination of the nature
of our mental representationof the biological world and identify a number
"
of cognitive blases that contribute to what they call the vivid illusion of
"
species. They claim that although speciesdo seemto have a distinctive psychological
representation, the specificform that representationtakesremains
an
largely open empirical question.
"
The concluding section- SpeciesBegonel" - contains two essaysthat, in
their own ways, expresssome skepticismabout the specialreality of species
that is the focus of biological and philosophical controversy regarding
"
"
species(as in the speciesproblem ). Both authors feel that speciesare as real
as higher taxa, but no more than the genuses, families, orders, and so on that
' "
those speciesconstitute. Marc Ereshefskys Speciesand the LinnaeanHierarchy
"
offers a review of our current thinking about the speciescategory,
advocating a replacementof the entire Linnaeansystem of classification. Ere
shefsky questionsthe distinctive reality of the speciescategory by pointing
to the problems in drawing the distinction between speciesand higher taxa
and by using the critiques of monistic accounts of speciesthat motivate
pluralism to suggest the heterogeneity of the species category. Because
the point of the Linnaeanhierarchy and the distinctions that it draws (e.g .,
between speciesand higher taxa) has been lost through the Darwinian revolution
, our current taxonomic practice createsproblems that alternative systems
of classificationmay avoid. Ereshefskyconcludesby examining two
such systems, though he acknowledgesthat any changeshould not be made
lightly" .
'
In Getting Rid of Speciesf Brent Mishler explores the application of
, Mishler views the Linnaean
phylogenetics to speciestaxa. Like Ereshefsky
.hierarchy as outdated, and like de Queiroz (1992; cE. de Queiroz, chapter3 in
this volume), he thinks that phylogenetic schemesof classificationare necessary
. Mishler argues that taxa at all levels, including the least inclusive,
/
Introduction
REFERENCFS
-McCord
on moralrt Rlism
Boyd, R. (1988). How to be a moralrealist. In G. Sayre
.
, ed., Essays
.
Ithaca
, N.Y.: CornellUniversityPress
, anti-foundationalism
for naturalkinds. Philosophi
, andthe enthusiasm
Boyd, R. (1991). Realism
calStudies
61, 127- 148.
, M., H. Dawah
, andM Wilson(1997
: Chapman
and
) . The"nitsof biodiomit,v. London
Claridge
Hall.
de Queirol. K. (1992). Phylogenetic
definitionsandtaxonomicphilosophy
. BiologyandPhiloso
.
7
295
313
.
,
phy
de Queirol. K. (1998). The generallineageconceptof &pedes
, &pedesaiteria, andthe process
of speciation
: A conceptualunificationand terminologicalrecommendations
." In Howardand
Berlodter
.
: Philosophical
. Cambridge
of things
Dupre, J. (1993). Thedisorder
founationsfor thedisunityof scimce
.
, Mass.: HarvardUniversityPress
. Philosophy
, M. (19928
). Eliminativepluralism
, 59, 671- 690.
Ereshefsky
of Science
.b). Theunitsof eoolution
: Ess
, M., ed. (1991
. Cambridge
Ereshefsky
,
Ilvs on the Mtureof species
Mass.: MIT Press
.
Ghiselin
, M. (1974). A radicalsolutionto the species
. Systmratic
Zooiogv1.3, 536- 544.
problem
.b.
1991
Reprintedin Ereshefsky
andtheoriginsofspecies
Ghiselin
, M. (1997). Metaphysics
. Albany, N.Y.: SUNYPress
.
.
Berlocher
HowardD . J., and SH
andspeciation
. Oxford:
, eds. (1998). Endless
fomrs: Species
OxfordUnivenity Press
.
HulLD. (1976). Are species
1.5, 174- 191.
reallyindividualsS,vstmratic
Zooiogv
HulL D. (1978). A matterof individuality. Philosophy
45, 335- 360. Reprintedin
of Science
.
b.
1991
Ereshefsky
HulL D. (1997). The idealspeciesde&nition andwhy we can't get it. In Claridge
, Dawah
, and
Wilson
.
Kitcher,P. (1984). Species
. Philosophy
51, 308- 333. Reprintedin Ereshefsky
.b.
1991
of Science
Lambert
, D., and H. Spence
. Baltimore
, eds. (1995). Spttiationandtherecognition
: Johns
concept
.HopkinsUniversityPress
.
Mayden. J. (1997). A hi~
y of speciesconcepts: The denouementin the sagaof the species
. problem. In Claridge, Dawah, and Wilson.
Otte , D., and J. Endier, ed. (1989). Spttiationand its conseq
UtnCtS. Sunderland, Mass.: Sinauer.
/
Introduction
.
andSpeciation
. In E. Vrba (ed.) Species
Paterson
. H. (1985). Therecognitionconceptof species
1992band Paterson
Pretoria
: TransvialMuseumMonographNo. 4. Reprintedin Ereshefsky
1994.
:
. Baltimore
: Colleded
andtherecognition
roritings
Paterson
. H. (1994). Eoolution
of species
concept
.
JohnsHopkinsUniversityPress
62,
. Philosophy
of Science
Stanford
, K. (1995). For pJuralismand againstmonismaboutspedes
70- 91.
. New
: A debate
andphylogenetic
Wheeler
theory
, Q., and R. Meier, eds. (1999). Species
concepts
.
York: ColumbiaUniversityPress
xvii
Intr~
on
On the Impossibility
of a Monistic Account
of Species
John Dupre
of true relationshipand
[ i]f we can onceandfor all lay the bogeyof the existence
- genealogical
relationship
realizethat thereare, not one, but many kinds of relationship
we
shall
and
so
on
,
, cytologicalrelationship
, morphologicalrelationship
releaseourselvesfrom the bondageof the absolutein taxonomyand gain enormously
.
in flexibility and adaptabilityin taxonomicpractice
- J. S. L. Gilmour, "The Development of Taxonomic Theory Since 1851"
, is meantthe actual, or ideal, arrangement
of any seriesof objects
By the classification
the
and
are
like
which
those
separationof thosewhich are unlike; the purpose
of
of this arrangementbeingto facilitate the operationsof the mind in clearly
.
conceivingand retainingin the memory, the charactersof the objectsin question
other
natural
or
series
,
,
Thus theremay be as many classifications
of
of
of any
bodies
, as they havepropertiesor relationsto oneanother, or to other things; or,
again, as thereare modesin which theymay beregardedby the mind.
- T. H. Huxley, Introductionto the Classification
of Animals
Most of the philosophical difficulties that surround the concept of species
can be traced to a failure to assimilatefully the Darwinian revolution. It is
'
widely recognized that Darwin s theory of evolution rendered untenable
the classicalessentialistconception of species. Perfectly sharp discontinuities
between unchanging natural kinds could no longer be expected. The conception
of sorting organisms into speciesas a fundamentally classi6catory
exercisehas neverthelesssurvived. Indeed, the concept of a speciestraditionally
has been the paradigmaticunit of classi6cation. Classi6cationis centrally
concerned with imposing conceptual order on diverse phenomena.
'
Darwin s theory, as the title of his most famouswork indicates, is about the
origins of diversity, though, so it is no surprise that the dominant task in
post-Darwinian taxonomy has been to connect classi6catorysystemsto the
received, Darwinian, account of the origin of diversity . Attractive though
. The patterns of diversity
this task undoubtedly is, it hasproved unsuccessful
that evolution has produced have turned out to be enormously diverse, and
in many casesthe units of evolutionary analysis have proved quite unsuitable
for the basicclassi6catoryaims of taxonomy. Or so I argue.
Why do we classify organisms? A natural and ancient explanation
expressedclearly by , for example, Locke (1689, bk. 3, chap. S, sec. 9) and
/
I. Monism
, Pluralism
, Unity andDiversity
WITHMONISM
TROUBLFS
The potential conflict between two main goals of classificationhas long been
recognized. The first and most traditional goal is to facilitate the communication
of information or to organize the vast quantities of detailed biological
information. From this point of view, a taxonomy should be constructed so
that knowing the taxon to which an organismbelongs should tell us as much
as possible about the properties of that organism. This goal must, of course,
be qualified by pragmatic considerations. Indefinite subdivision of classifications can provide, theoretically, ever more detailed information about the
individuals classified: assignmentto a subspeciesor a geographicalrace will
presumablygive more information than mere assignmentto a species.As the
basal taxonomic unit, the speciesshould be defined, therefore, to classify
organisms at a level at which the gains from finer classificationwould be
outweighed by the costs of learning or transmitting a more complicatedset
of categories. If organismsvaried continuously with no sharp discontinuities,
this balancing of costs and benefits would present a largely indeterminate
problem. By happy chancefor many kinds of organismsthere appear to be
sharp discontinuities at a relatively fine classificatory level that are much
sharperthan any discontinuitiesat any lower level. To the extent that this is
the case, the selection of the appropriate level for assignmentof organisms
to speciesappearsunproblematic.
in recent years, this goal of organizing biological information has been
emphasizedmuch less than a second, that of mapping the currents of the
evolutionary process. A recent anthology of biological and philosophical
essayson the nature of speciescarried the title The Units of Evolutionand the
subtitle Essayson the Nature of Species(Ereshefsky1992). Though the idea
:~
Dupre
/
Momsm, PluralismUnity andDiversity
/
of a MonisticAccountof Species
: OntheImpossibility
Dupre
I. M~ sm
. Pluralism
Unity andDiversity
likely event that the flow is quite spatially limited, the claim that the whole
complex group with its virtually worldwide distribution can be seen as
reproductively connectedis tenuous to say the least.
The conclusion I want to draw at this point is that the BSC will frequently
lead us to distinguish speciesin ways quite far removed from traditional
Linnaean classi6cationand far removed from the optimal organization of
taxonomic information. Moreover, the theoretical motivation for the BSC
seemsseriously de6cient. The sorts of criticisms I have been enumerating
above have led, however, to a decline in the extent to which the BSC is now
acce}?ted, and this decline has been accompaniedby increasing interest in a
rather different approach to evolutionarily centered taxonomy that can be
speciesconcept(PSC).
broadly classi6edunder the heading of the phylogenetic
(The definite article preceding the term should not be taken too seriously
here, as there are severalversions of the generalidea.)
The central idea of all versions of the PSC is that species- and, in fact,
higher taxa as well should be monophyletic. That is, all the membersof a
speciesor of a higher taxon should be descendedfrom a common set of
ancestors. An appropriate set of ancestors is one that constitutes a new
. The
branch of the phylogenetic tree. Sucha group is known as a stemspecies
is
a
taxon
is
whether
of
PSC
versions
between
merely
important distinction
or to
stem
a
of
descendants
to
contain
species
only
particular
required
. The latter position is definitive of
contain all and only such descendants
cladism, whereasthe former, generally describedas evolutionary taxonomy,
requires some further criterion for deciding which are acceptablesubsetsof
.6 Two issuesarise in explicating a more detailed account of the
descendants
PSC. First, what constitutes the division of a lineage into two distinct lineages
? Second, what constitutes
and hencequali6esa group as a stem species
a lineageand its descendantsas a species(or, indeed, as any other taxonomic
rank)?
The traditional answer to the mst question is that a lineage has divided
when two components of it are reproductively isolated from one another,
but the difficulties raisedin connectionwith the BSCsuggestthat this answer
is inadequate. Examplessuch as oaks suggest that reproductive isolation is
not necessaryfor the diVision of a lineage, and worries about the lack of
gene flow within apparently well-de6nedspeciessuggest that it is not sufficient
either. An illuminating diagnosis of the difficulty here is provided by
Templeton (1989), who distinguishes geneticexchangeability, the familiar
graphic
ability to exchange genetic material between organisms, and demo
that
'"
extent
the
to
two
between
exists
which
,
organisms
exchangeability
they share the samefundamentalniche (p. 170). The problem with asexual
taxa and with a variety of taxa for which gene exchangeis limited is that
Tile boundaries de6ned by demographic exchangeability are broader than
those de6ned by genetic exchangeability. Conversely, for casesin which
well-de6ned speciespersist despite gene exchange, the boundaries de6ned
I. MonJsm
. PluralismUnity and Diversity
tween the theoretical account of a species and a practically useful classification would surely be severed.
The question that must be faced, then , is whether from the PSC point of
view the idea that the species is the basal taxonomic unit - where taxonomy
is conceived as providing a practically useful classification - can be maintained
. Abandoning the BSC will take care of species that look unsatis factorily large by allowing a variety of cohesion mechanisms apart from
reproductive isolation , but it will tend to imply the presence of disturbingly
small species. Frequently there are clearly distinguishable groups of organisms
subspecies, varieties , geographical races below the species level .
There is no reason to suppose that these groups are not monophyletic and
no reason to suppose that they are not , at least for the moment , evolving
independently . There is no doubt that such groups are often clearly distinguishable
, and indeed for many purposes classification at this level is the
most important . Stebbins ( 1987, 198 ) notes , for instance, that foresters are
often more concerned with geographic races than species and indeed can be
hampered in their work by the confusing attachment of the same specific
name to trees with quite distinct ecological properties and requirements . A
judge at a dog show is not much concerned with the criteria that identify
something as Canis familiaris . Such groups may go extinct , they may merge
With other subgroups in the species, or they may be destined to evolve
independently into full -blown species or higher taxa. Their evolutionary significance
is thus unknown and unknowable . The same, of course, could be
said of groups recognized as full species, though the second alternative
(merging with other groups ) may be rare.
THECASEFORPLURALISM
An evolutionarily based taxonomy appearsto be faced at this point with
only two possibleoptions. The first is to considerspeciesasby definition the
smallest units of evolution. Leaving aside the insurmountable difficulty of
, my argument so far has been that .this
detecting such units in many cases
will
a
fundamental
classification
that is often much too fine to
option
provide
be useful for many of the purposesfor which taxonomieshave traditionally
been used.7 Mishler and Donoghue (1982) suggestthat this proposal is also
"
conceptuallyconfused. They argue that there are many evolutionary, genealogical units within a given lineage . . . which may be temporally and spatially
"
overlapping (1982, 498). They suggest, therefore, that it is an error to
supposethat there is any such thing as a unique basalevolutionary unit and
that the particular evolutionary unit one needsto distinguish will dependon
'
the kind of enquiry with which one is engaged. If there is no unique basal
unit, then there is no privileged unit and, from an evolutionary point of
view, no theoretical reason to pick out any particular group as the species.
Mishler and Donoghue therefore propose the second option, to " [a]pply
speciesnamesat about the samelevel as we have in the past, and decouple
the basaltaxonomic unit from notions of ' basic' evolutionary units" (p. 497).
This processinvolves seeingspecieson a par with generaand higher taxathat is, as ultimately arbitrary levels of organization, chosenon a variety of
8
pragmatic grounds.
Although Mishler and Donoghue seethe speciesas an ultimately arbitrary
ranking criterion, they do maintain a version of the PSC and, hence, do not
seeit as arbitrary from the point of view of grouping. In fact, they endorse
the strong, cladistic concept of monophyly as a condition on a group constituting
a species(or, for that matter, a taxon at any other level). Their
"
"
"
pluralism, however, entails that comparative biologists must not make
inferencesfrom a speciesname without consulting the systematicliterature
to see what patterns of variation the name purports to represent" (p. 500).
But given this degreeof pluralism, and the rejection of the attempt to equate
the basal taxonomic unit with any purportedly fundamental evolutionary
unit, one may reasonably wonder why it is desirableto insist neverthelesson
the requirementof monophyly . I suspectthat part of the motivation for this
requirementis the idea that there must be someanswer to the question what
a speciesreally is. It was once, no doubt, reasonableto supposethat evolution
had produced real, discrete speciesat approximately the classificatory
level of the familiar Linnaeanspecies.Perhapsthis supposition was an almost
inevitable consequenceof the transition from an essentialist, creationist view
of nature to an evolutionary view. Acceptanceof evolutionary theory would
require that it more or less serve to explain biological phenomenaas theretofore
understood. Nevertheless, a further century of development of the
evolutionary perspectivehas given us a radically different picture of biological
diversity . The sharpnessof differentiation between kinds and the processesby which suchdifferentiation is producedand maintainedhave proved
to be highly diverse. There is no reasonto supposethat evolution has provided
any objectively discoverableand uniquely privileged classificationof
the biological world.
Why , then, should we continue to insist that evolution should provide a
necessarycondition, namely monophyly, on any adequatebiological taxon?
I can think of only three possible answers. First, it might be held that a
better understandingof evolution is so overwhelmingly the most important
biological task that any taxonomy should be directed at improving this
understanding. Second, it might be thought that an evolutionarily basedtaxonomy, despite its problems, would provide the best availabletaxonomy, or
at least a perfectly adequatetaxonomy, for any biological project even far
removed from evolutionary concerns. Or third - and this, I suspect, is the
. most influential motivation it may be held on general methodological
grounds that a central concept such as the speciesmust be provided with a
unitary definition. This third motivation might be grounded either in ageneral
commitment to unification as a scientific desideratumor on the fear that
failure to provide a unified accountof the speciescategory will lead to massive
.confusion as biologists attempt to communicate with one another. I
sm
, PluralismUnity andDiversity
/
: On theImpossibilityof a MonisticAccountof Spedes
Dupre
/
Monism, Pluralism
, Unity andDiversity
pose that the moon was a planet. Not every possible misunderstandingcan
be forestalled.
The danger of confusion is a more plausible concern regarding the idea
that the sameorganismsmight be subject to different principles of classifica
tion for different biological purposes. In one sense, I am happy to agreethat
this type of confusion should be avoided. It would be undesirablefor a particular
, to be variously defined and to have
speciesname, say Mus musculus
extensions
to
the
taxonomic
varying
according
theory espousedby various
authors. We should aim to agree as far as possible which organisms are
house mice. In the concluding section of this paper, I explain how I think
such species names should be understood. If, to recall my hypothetical
exampleabout rats, it proves useful to treat scavengingrats as a basickind in
some ecological model, it would be misguided to insist that scavengingrats
constitute a species. Equally clearly, however, this concessionto standardized terminology does not at all require that all species names be conceived
as answering to the samecriterion of what it is to be a species. The
other consequenceof insisting on an unambiguousinterpretation of particular
speciesnamesis that we Cannotassumea priori that the canonicaltaxonomy incorporating standard speciesnameswill be suitable for all biological
purposes. The question here is, again, an empirical one that depends ultimately
on how orderly biological nature turns out to be. If it should prove
to be disorderly in the relevant sense, then biology would prove to be a
more complicated discipline than is sometimesassumed
. But once again I
cannot seethat any unavoidableconfusion need be introduced.
CONCLUSION: A CASE FOR TAXONOMIC CONSERVATISM
Many taxonomists and almost everyone who usesthe results of taxonomic
work have complained about the genuine confusion causedby changesin
taxonomic nomenclature. Someof thesechangesseementirely gratuitousfor example, changesin the names of taxa grounded in the unearthing of
obscureprior namingsand in appealsto sometimesesotericrules of priority .
Other changesare more theoretically based adjustments of the extent of
particular taxa. Many suchtheoretically motivated changeshave beenalluded
to in this paper. BSC-committed theorists will urge that discoveriesof substantial
gene-flow betweenotherwise apparently good speciesshould lead us
to apply one speciesnameto what were formerly consideredseveralspecies.
Phylogenetic taxonomists will want to amend the extensionsof any higher
taxa that fail their favored tests for monophyly, and strict cladists will promote
the breaking up of prior " species" into various smallerunits when their
favored criteria for lineage splitting demand it .II Less theoretically commi,tted taxonomists may promote the splitting or lumping of higher taxa on
the basis of general principles about the degree of diversity appropriate to
a particular rank.
/
: OntheImpossibilityof a Monistic Account of Species
Dupre
/
I. Monism, Pluralism
, Unity and Diversity
mind in clearly conceiving and retaining in the memory the charactersof the
objects in question. Plainly to the extent that taxonomic namesare undergoing
constant modification, what anyone person " conceivesand retains in
the memory" will be potentially incommunicableto others, and the possibility
of reliably adding further information obtained from the work of others
will be constantly jeopardized. This is not to say that taxonomic revision is
never justified. If a speciesis included in a genus in which it is highly anomalous
, and if that speciesis much more similar to other speciesin someother
genus, then the goals of organizing information will be better served by
reassigningit . It is of coursealso true that monophyletic taxa will tend to be
more homogeneousthan polyphyletic taxa, and that in paraphyletic taxataxa in which some of the descendantsof the common ancestorsof aparticUlar
taxon are excluded- there will be often be a case, on grounds of similarity
, for including the excluded parts of the lineage. My point is just that
theseconsequences
rather than monophyly itself shoUldprovide the motivation
for taxonomic change, and the benefitsof such changemust be weighed
carefully against the potential
. costs. In this weighing process, the presumption
that taxon namesretain constant extension shoUldprobably be kept as
strong as possible to maximize the ability of biologists to maintain reliable
and communicableinformation.
To take perhapsthe most familiar example, it seemsto me that there is no
case at all for revising the class Reptilia (reptiles) to include Aves (birds).
This move is requiredby a strict cladistic conceptof monophyly becauseit is
believed that birds are descendedfrom ancestralreptiles. We cannot exclude
these avian ancestorsfrom the class that includes modem reptiles because
crocodiles, still classedas reptiles, are believed to have diverged from the
main reptilian lineage earlier than birds did. The fact remains, however, that
most zoologists, I suppose, woUld consider crocodilesmuch more like other
reptiles than either is like any bird. The attempt to convince the learned or
the vulgar world that birds are a kind of reptile strikes me as worse than
pointless. It may be said that the only important claim is that Aves shoUldbe
classifiedas a lower-level taxon included within Reptilia, and that this classification has nothing to do with our common usageof the terms reptileand
bird. Although it is certainly the case that scientific taxonomic terms frequently
differ considerably from apparently related vernacular terms, this
differentiation is a sourceof potential confusion that shoUldnot be willfully
exacerbated(seeDupre 1993, ch. 1, and forthcoming). It is also unclearwhat
advantageis to be gained from insisting on sucha revision. All evolutionists,
I suppose, are likely to be familiar with recent thinking on the historical relatio ~ships within the main groups of vertebrates, and if they are not, their
ignoranceis not likely to be relieved by terminological legislation. Similarly,
experts on smaller groups of organisms will presumably be familiar with
current thinking on phylogenetic relationshipswithin those groups. To celebrate
every passing consensuson these matters with a change in taxonomic nomenclatureis an inexcusableimposition of a particular professional
/
Dupre: On the Impossibiliry of a Monistic Account of Speaes
/
I. Monism,Pluralism
, Unity andDiversity
species and had in mind kinds , not things . Arguably , the tension between
these two usages is at the root of the great philosophical perplexity that the
concept of species has generated in this century . In arguing for reversion to
the earlier usage of the term species
, I am at least honoring conventions of
priority . What I am proposing , however , is not much like a Linnaean taxon omy either . As many have observed , Darwin forced us to give up any traditionally
essentialist interpretation of taxonomic categories and even any
objectively determinate taxonomy . But almost a century and a half of biological
work in the Darwinian paradigm have also shown us that evolution
does not reliably produce units of biological organization well - suited to
serve the classificatory purposes for which the concept of species was originally
introduced , so perhaps rather than a reversion to Unnaeus , it would be
better to see my proposal as a quasi- Hegelian synthesis . At any rate, if I
seem to have been implying that Darwin may have been responsible for
introducing some confusion into biology , I am sure no one will take this as
more than a peccadillo in relation to his Unquestion ably positive contributions
.
ACKNOWLEDGMENTS
I would like to thankRobWilsonandChrisHorvathfor valuablecomments
on a draft of this essay.
NOTES
1. It is not entirely clearhow to makethis idea precise. Obviously, not every organismfounds a
. A natural idea is that every organism in
lineage, unlessevery organismis to constitute a species
any way genetically distinct from its parent should found a new lineage. Given, however, the
possibility of the samepoint mutation occurring more than once, it could turn out that a set of
genetically identical organismsmight constitute two or more distinct species.
The proposal also leads to the surprising conclusion that the vast majority of speciesare
asexual. As Hull notes, this conclusionmay mitigate the well-known difficulty in explaining the
origin of sex by showing that sexual reproduction is a much rarer phenomenonthan is often
'
supposed(1989, 109). I should also mention that Hull s proposal is made.in connectionwith the
thesisthat speciesare individuals, and is thus not necessarilyan explicit defenseof the BSC.
2. It appearsthat the sameis probably true for somekinds of flowering plants (seeNiklas 1997,
74 fE.).
3. This claim is perhapsless true now than it was twenty years ago. An influential evolutionary
classificationof bacteria was proposed by Woese (1987); see also Pace (1997). On the other
hand, Gy Uenbergand others (1997) aim explicitly to producea classificationthat is optimal from
an information-theoretic perspective, a goal that there is no reasonto supposewould be met by
any imaginable phylogenetic scheme. Seealso Vandammeand others (1996) for a related proposal
. It is clear, at any rate, that any possiblephylogenetic classificationof bacteria, if it is to be
of ~ y practical use, must de6ne taxa with great clonal diversity. Gordon (1997), for instance,
Collpopulations in feral mice was an increasing
reports that the genotypic diversity of Escherichia
function of the age of the mouse, indicating the development of distinct clones during the lifetime
of the n:' use. I assumeone would not want to think of this developmentas speciation, but
/
: On the Impossibilityof a MonisticAccountof Species
Dupre
given this clonal diversity, it is difficult to seehow any useful taxonomy could avoid being arbitrary
from a phylogenetic perspective. The situation is still worse in view of the partially reticulate
phylogeny consequenton genetic exchangebetweenbacteria.
"
4. An extreme statementof this optimistic view can be found in Ruse(1981, 237) : There are
different ways of breaking organisms into groups and they coincide! The genetic speciesis
the morphological speciesis the reproductively isolated speciesis the group with common
"
ancestors.
'
S. On R. fruticosus
, the common blackberry, Benthamand Hooker (1926, 139) wrote: it varies
considerably. The consequencehas been an excessivemultiplication of supposedspecies. ..
although scarcely any two writers will be found to agree on the charactersand limits to be
"
"
"
"
assignedto them. The same species is describedby Schauer(1982, 346) as aggregate, variable
"
with very numerous miaospecies . More optimistically, The Oxford Book of Wildjlowm
"
(Nicholson, Ary , and Gregory 1960) statesthat [t ]here are severalhundred speciesand hybrids
"
in the Rubusgroup, and only an expert can identify all of them .
6. SeeSober(1992) for a very clear exposition of this distinction. Although the debatehere is a
fundamentalone, it is not of central concernto my essay.
1. SeeDavis (1918, 334- 338) for a discussionof someof the difficulties in subspeci&c classmca
tion of angiosperms.
8. For further elaboration, seeMishler and Brandon(1987) . For more generalargumentsagainst
any fundamentaldistinction between speciesand higher taxa. seeEreshefsky(1991 and chapter
11 in this volume) and Mishler (chpater 12 in this volume).
9. Somemore realistic exampleshave beendiscussedby Kitcher (1984).
10. For referencesto bacterialtaxonomy and brief discussion
, seenote 3.
11. De Queiroz and Gauthier (1990, 1994) claim that taxonomic changesthey advocate will
promote constancy of meaning, or definition, for taxa. Mammalia, for example, should be
defined as the set of descendantsof the most recent common ancestor(i.e., ancestralspecies
) of
monotremesand therians. The extensionof sucha term, however, will be constantly revisablein
the light of changesin opinion about the details of evolutionary history . From the point of view
of the consmnerof taxonomy, at least, I suggest that constancy of extension is surely more
valuablethan constancyof definition.
'
12. Subsequentto Walters paper, the quince appearsto have been reconceivedas Chatnomtles
(though not unanimouslyaccording to the few sourcesI consulted on this matter). This reconception
effects a conjunction with the ornamentalflowering quinces. One might speculatethat
the increasingobscurity of the quinceas a fruit might have exposedit to this annexation, whim
one doubts could have happenedto the apple.
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Ehrlich
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58,
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, highertaxa
, andthe unitsof evolution. Philosophy
Ereshefsky
of Science
84- 101.
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, M., ed. (1992). Theunitsof evoluh
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MIT Press
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. Bacteri
, G. D. (1962). Someremarkson the theoreticalaspects
Floodgate
26, 277- 291.
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2,
Ghiselin
, M. T. (1987
, individuality,andobjectivity. BiologyandPhilosophy
). Species
concepts
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Ghiselin
, M. T. (1997). Metaphysics
Gilmour,J: S. L (1951). Thedevelopment
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, andM. Verlaan(1997). Classi
Gyllenberg
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: Two thousandyearsof stasis
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Hull, D. L (1989). Themetaphysics
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Kitcher,P. (1984). Species
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MilL J. S. (1862). A system
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/
I. Monism, Pluralism, Unity and Diversity
Party Line
DavidL. Hull
In the nineteenth century, one of the hot topics of debate was the plurality
of worlds. Did God create a single Earth inhabited by all and only those
souls that Jesusgave the opportunity to be savedfrom eternal damnation, or
did He createmillions of worlds inhabited by just as many morally responsible
beings? On the first alternative, God would appearto be as profligate as
the most extravagant wastrel. He createdmillions of nebulae, each containing
just as many stars, eachof which might have planets circling it in stately
regularity, but on only one of these planets circling a single star did He
breath soul into a single species.What a waste. But if we assumethat God is
the ProtestantGod of " wastenot , want not," surely He would not have let so
many opportunities slip through his fingers for creating subjectsto worship
him. Not only did Jesuscome down to Earth to be sacrificedfor our sins, but
apparently he also repeatedthis ritual in world after world after world (for a
'
history of this controversy, seeDick s 1982 Plurality of Worlds).
No less a figure than William Whewell entered into this debate- on the
side of the monists. For fear of damaging his hard-eamed reputation as a
sober seeker after truth , Whewell anonymously published The Plurality of
Worlds (1853). To those who complained that God and hence nature did
nothing in vain, Whewell cited all the waste that was already so apparentin
this world:
We reply, that to work in vain, in the senseof producing meansof life which
are not used, embryos which are never vivified , germs which are not developed
; is so far from being contrary to the usualproceedingsof nature, that it
is an operation which is consistently going on, in every part of nature. Of
the vegetable seedswhich are produced, what an infinitely small proportion
ever grow into plants! Of animal ova, how exceedinglyfew becomeanimals,
in proportion to those that do not; and that are wasted, if this be waste!
(p. 249)
A similar question might be asked of God about speciesin general. God
creatednumerousdifferent species
, but did He create a singlesori of species
or many differentsoris of species7Each and every organism belongs to one
speciesand one speciesonly , but are all these speciesof the samesort7 Or
possibly, does anyone organism belong to many different sorts of species7
/
I. Monism
, Pluralism
, Unity andDiversity
REALISM
ANDANTIREALISM
One reasonwhy philosophers
find the monism-pluralismdebateso interesting
is its apparentconnectionto the disputeover realismandantirealism
. Of
the four possiblecombinationsof thesephilosophicalpositions, two seem
, andpluralismcombinedwith
quite natural: monismcombinedwith realism
antirealism
. Monistsarguethat scientistsshouldstrive to find the bestway
to divideup the world andthat sucha bestway doesexist, eventhoughwe
) whetheror not we have
may neverknow for sure(whateverthat means
found it. Yes, scientistsarefallible. Yes, conceptualrevolutionsdo occurin
science
, revolutionsthat reqUireus to startnot all over again, but at leasta
few stepsback. Giventheseassumptions
, the goal of findinga single, maximally
informativeconceptualization
of natureseemsboth desirableandreasonable
. If one is going to be flat-footedand simpleminded
with respectto
onephilosophicalposition, why not two? (Holsinger[1987] interpretsSober
[1984a]asa monist-realist.)
, pluralistsarguethat the differencesof opinion so characteristic
Conversely
of sciencewill continueindefinitelyinto the future. The natureof these
differences
everemerge
, but rarelydoesconsensus
, andwhen
surelychanges
" the" world canbe
it does, it is likely to be short-lived. How come? Because
charaderizedin indefinitelymany ways, dependingon differencesin perspectives
, worldviews, paradigms
, and what haveyou. Eventhoughnot all
of thesewaysareequallyplausible
, acceptable
, or promising
, no oneway is
clearlypreferableto all othersonceandfor all. Onemustkeepan openmind.
If one is going to be sophisticated
and nuancedwith respedto one philonot
two?
Stanford
(
[1995] portrayshimselfasbeinga
sophicalposition, why
antirealist
.
)
pluralist
The other two combinationsof the two philosophicaldistinctionsare
somewhatstrained
. It would seema bit strangeto arguethat one and only
one way existsto divide up the world, but that the groupsof naturalphenomena
arenot "real." They areas real
producedon this conceptualization
asanythingcanget! Of course
, realcanbe definedin sucha way that nothing
couldpossiblybe real, just asphilosophers
havedefinedknowso that no
one ever knowsanythingand law so that no generalizations
ever countas
la.ws
but
this
as
much
fun
as
it
is
to
seems
,
game,
play,
nonproductivein the
. A combinationof pluralismandrealismseemsequallypeculiar(but
extreme
seeDupre1981andKitcher1984a
). Theworld canbe dividedup into kinds
in numerousdifferentways, and the resultsareall equallyreal! Onceagain,
/
Hull: On the Plurality of Species
incommensurable
, but the sort of holistic semanticsthat generatesincommensurability again seemsa high price to pay. (For a discussionof realism
and pluralism with respect to the units of selection controversy, see Sober
1984b; Sterelny and Kitcher 1988; Kitcher, Sterelny, and Waters 1990;
Waters 1991; Soberand Wilson 1994; Shanahan1997.)
A RETURNTO BABEL
Kitcher (1984a, 326) suggeststhat one worry possibly bothering opponents
of pluralism is that it might " engendera return to Babel." Do monists treat
languagemonistically1Do they think that every utterancehas one and only
one meaning1Are pluralists pluralistic when it comesto language1Should I
treat every utterance made by pluralists pluralistically, or is it just possible
that texts do on occasionconstrain interpretations1As far as I know, no one
deniesthe existenceof ambiguity and vaguenessin language. In fact, in the
face of imperfect knowledge, vaguenessmay be necessaryfor communication
(Rosenberg 1975). Again, I find it difficult to tell, but even the most
rabid deconstructivist shies away from anything- literally anything- goes.
Even they seemto assumethat they are saying something or trying to say
. Possibly, they are not saying
something with varying degrees of success
one and only one thing with absolute clarity, but not all interpretations
are equally acceptable- Emily Dickinson as a Marxist feminist. Perhapsour
intended meaning is neither patent nor all there is to the story, but at the
very least I have been taught that we should all aim to present our views as
clearly and unambiguously as possible, and I see no reason to give up the
ghost at this late date. PerhapsI can be a selective pluralist about a halfdozen conceptsat a time, but I cannot treat all of languagepluralistically all
at once, not if I want to say something, not if I want other human beings to
understandme.
REFLEXIVITY
Early on in the prehistory of what has come to be known as the Science
Wars, young sociologists who were in the midst of rediscovering epistemological relativism also stumbled upon reflexivity when an occasionalopponent
askedwhy they were gathering so much evidenceto cast doubt on the
. efficacy of evidence. These sociologists tried a variety of ways to extricate
"
"
themselvesfrom this tension in their position. Some argued that sociolo"
gists, when they are acting as sociologists, must treat the social world as
real, and as something about which we can have sound data, whereas we
should treat the natural world as something problematic- a social construct
rather than as something real" (Collins 1981c, 217; see also Collins 1981a
/
I. Monism
, Pluralism
, Unity andDiversity
CONsm ERAnO NS
PROFFSSIONAL
.
I.n general, people find .it much more plausible and desirableto counselpluralism
with respect to otherpeople' s areasof expertise than their own. As I
have argued above, philosophers find pluralism extremely attractive in
science, -much more so than in philosophy. Scientists in turn do not find
/
Hull: On thePluralityof Species
pluralism all that attractive in their own area of expertise and usually stay
well clear of philosophy. At least sometimes, scientiststhink that they have
the right answer to a particular question. What would sciencebe like in the
absenceof such convictions? Picture hundredsof scientists, eachbeing terribly
considerateof eachother' s hypotheses:
"You think that selection occurs
only at the level of the genetic material?
That may be so, but in addition, individual organismsare the main target of
selection."
"
I agree with everything you say and want only to add that selection
wandersup and down the organizationalhierarchy in biology ."
"
To supplementthe nuancedposition being expressed
, I submit that no such
as
It
selection
exists.
is
two
es
not
one."
,
really
thing
process
" So true and selectionis of
"
,
only peripheralimportancein evolution.
"
"
Speakon, oh wise one.
A preference for monism and pluralism waxes and wanes as various
groups gain and lose power. Right now, advocatesof developmental systems
theory are trying to supplant the current gene-centered world view
(Moss 1992, Griffiths and Gray 1994). None too surprisingly, these developmentalists are urging pluralism. In general, groups who hold minority
opinions at a particular time find pluralism to be the correct philosophical
view, whereasthe groups in power are not nearly so attracted to it . During
the heyday of the biological speciesconcept, Mayr saw no reason to give
ground to his opponents. He insisted that there are basic units in the evolutionary
process, that theseunits are delineatedin terms of reproductive isolation
, and that making these units coincide with the basic units of
classification is both possible and desirible. I predict that if and when
developmentalistssee their views prevailing, they will ceasetheir pleas for
pluralism and becomestaunchmonists. Pluralismlooks good to outsiders
regardlessof whether they belong to different disciplines(e.g., philosophers
looking at science) or to groups currently a minority within a particular
discipline (e.g., developmentaliststrying to muscle evolutionary biologists
out of their positions of power).
ANALYSFSOF CAUSA
nON
/
I. Monism
, Pluralism
, Unity andDiversity
Hull:OnthePlurality
ofSpecies
been funded? If so, would these pluralists be willing to live with the disastrous
results of their decision? If not , why not ? How about today ? Are
poppers still a plausible causative agent for AIDS ?
If the contrast between monism and pluralism is to be of any significance
at all , advocates on both sides of this divide have to admit that at least on
occasion the position they prefer might be wrong . Monists and pluralists
alike have to present a list of criteria to help in deciding when one, two ,
three, or more possible alternatives are justified . The danger is that every
situation , no matter how apparently straightforward , turns out to be hope lessly complex . It also must be noted that the decisions that we make in this
connection are likely to have effects on society . People do not , as a rule , pay
excessive attention to what we philosophers have to say, but sometimes the
distinctions that we make and the positions that we espouse find their way
into the public at large . For example , the tobacco industry for years has
claimed that the scientific data are inadequate to prove that smoking causes
lung cancer. After all , some people smoke three packs of cigarettes a day and
die at age ninety -five in a car accident , whereas others contract lung cancer
at an early age although they have never been exposed to cigarette smoke.
In most scientific contexts , the factors that scientists pick as causes are
rarely necessary conditions ; they are even more rarely sufficient conditions ;
and they are hardly ever both necessary and sufficient conditions . Perhaps
finding necessary and sufficient conditions for the occurrence of natural phenomena
is the ideal, but in most contexts we have to settle for much less.
The issue is statistical correlations . Holding everything else constant , how
'
much does smoking increase one s chances of contracting lung cancer- or
heart disease, for that matter ?
"
If Smoking causes heart disease" turns out not to be true for humankind , I
take it that its truth has been established for the various populations of
Western countries in which it has been systematically investigated . It is now
known , for example , that smoking causes heart disease for the human population
of the United States. There may be inductive hazards in the extrapolation
of this result to other identifiable human populations ; but it seems
unnecessarily cautious to restrict the claim to an as-yet -unidentified subpopulation
of inhabitants of the United States. ( Dupre 1993, 200 )
If we philosophers were to stop here, the effects of our analysis would be,
from my perspective , decidedly beneficial . Smoking is a major cause of both
lung cancer and heart disease, and anyone with a shred of intellectual integrity
has to concur , pluralism be damned.
However , we do not st~ p here. Promiscuous pluralists (not to be confused
'
with Dupre s promiscuous realists ) feel obligated to resist the conclusion that
.
smoking causes lung cancer, even in a statistical sense, because it is insufficiently
nuanced. Eve.n when people do smoke and do contract lung cancer, it
"
"
does not follow that smoking is the cause of lung cancer. All sorts of
alternatives and combinations of alternatives might and probably do playa
role in people coming down with lung cancer. In fact , as the father of statis -
tics (not to mention the synthetic theory of evolution), Ronald Fisher (1958,
108) mused, perhapslung canceris one of the causesof smoking cigarettesf
Causalsituations are extremely complicated. All sorts of supplementalfactors
can playa role, and the reasonsfor selecting one factor and terming it
" the" causewhile
"
demoting all the others to the position of supplemental
"
factors are far &om obvious. This issue arises time and again in science.
Natural selectionis the causeof organic complexity, while every other influence
"
is a constraint." Genesare the causeof various traits, whereaseverything
else is demoted to being part of the "background knowledge." Many
evolutionary biologists claim that gene exchangeis crucial in the individuation
"
"
of speciesas units of evolution, but all other factors only contribute
to the cohesion that is so important in speciesbeing species. On what
grounds are causesassigned to these various categories? This literature
throws up a picture of the empirical world so murky that even the tobacco
industry can hide in the fog.
ANDMUDDLED
METAPHYSICS
SPEaFS
At long last, after all the precedingpreparatory discussion, it is time to turn
to the speciesproblem. Numerous philosophers have urged pluralism with
respectto biological species. As a point of departure, I take Philip Kitcher' s
(1984a, 1984b, 1987, 1989) discussion of species. As is usually the case,
'
deciding what an author s position is can pose seriousproblems. Too many
alternatives present themselves. (In this respect, pluralists are correct more
often than I would prefer.) With respect to his own general philosophical
position, Kitcher (1984a, 308) is not very pluralistic. In his paperson species,
he sets himself the dual tasks of explaining a position about speciesthat he
"
"
terms pluralistic realism and of indicating in a general way why he thinks
"
that this position is true." Kitcher does not say that from someperspectives
and in certain circumstance
, pluralism is preferable, or that from other perspectives
and in other circumstancesmonism is the correct position to hold.
If I read Kitcher correctly, he believes that no form of monism is acceptable
by anyone no matter what. With respect to his own philosophicaloutlook,
Kitcher is inclined to monism. So am I with respectto mine.
Kitcher then turns to the issue of the ontological character of species.
Periodically, authors have tentatively suggestedthat speciesare like individual
organismsmore than they are like universalssuchas triangularity, but
until
Michael Ghiselin (1974) did this position becomewidely discussed.
not
According to Ghiselin and later Hull (1976), if speciesare to fulfill their role
in the evolutionary process, they must be conceivedof as spatiotemporally
10'caIized entities connectedin spaceand time (seealso Mayr 1999; Mishler
and Theriot 1999: and.Wiley and Mayden 1999). At anyone time, species
must exhibit a certain degree of cohesiveness(though the mechanismsproducing
this cohesivenessmight vary ), and through time, they must be connected
can be
.2_They are chunks of the genealogicalnexus. The term species
/
Hull: On thePluralityof Species
that nothing in biology makes sense except in the light of evolution is a bit
of an exaggeration , but not much. In sum, I think that (a) the evolutionary
"
"
perspective in biology is in a significant sense basic to all of biology ; (b)
from the evolutionary perspective , species must be treated as historical entities
'
; and (c) Kitcher s interpretation of species as historically connected sets
'
and Boyd s interpretation of species as homeostatic property cluster kinds
are, to say the least, strained . Even if we accept the alternatives that Kitcher
and Boyd suggest , the distinction between universals and particulars must be
reintroduced by distinguishing between spatiotemporally restricted and connected
sets or kinds and those sets or kinds that lack these restrictions . What
such an exercise accomplish es, I fail to see.
I. Mo m, Pluralism
, Unity andDivenity
if not
pervasive in our conceptions of the world is currently unanalyzed,
be askedof speciesdefinitions in terms
unanalyzable. A question that must
"
"
"
is
similar
similar
How
is
of similarity ,
enough and in what senseof similar
" ? Can one level of
similarity be specified- one level that can be applied
classifiequally acrossall organismsto produce even a minimally acceptable
cation? The answer to this question is, thus far, no. One implication of this
conclusion is that no general purpose classificationof plants and animals is
overallsimilarity (but
possible. Generalpurposeremainsas unanalyzed, as does
seeDupre, chapter 1 in this volume).
The historical alternative avoids the problems that plague speciesdefinitions
in terms of similarity, but it hasproblemsof its own- chiefly difficulties
involved in reconstructing phylogeny, which in turn requires that homologies
be distinguished from homoplasies. Organisms on Earth evolved the
includes lots
way that they did and no other way . Even though" phylogeny
" But
reconstructing
of merger, there is one and only one phylogenetic tree.
the information
of
Most
.
not
if
difficult
be
,
can
impossible
extremely
phylogeny
extant
of
is
world
organisms. Our
that we have of the biological
. The principles
sketchier
knowledge of extinct forms .is even sketchier much
of cladistic analysis were devised to establish transformation series in
which genuine characters(homologies) nest perfectly and hence produce
not nest perfectly is
perfectly monophyletic taxa. Any characterthat does
volume) argues, we
this
in
5
not a genuine character. As Sterelny (chapter
"
must view phenomenologicalspecies identifiable clustersof organisms
"
as fallible cluesto the existenceof evolutionarily linked metapopulations.
With respect to monism versus pluralism, systematistsare put in a bind.
The rules of nomenclaturedo not allow them to be pluralists. They have to
as Kitcher would have
produce one and only one classification not nine
them do. Systematic principles that take history as basic seem appealing
becausethey can promise a single classification. It may not be equally useful
for all sorts of purposes, but it is at least attainablein principle. Possessinga
faults, is better than
single classificationas a referencesystem, no matter its
. The metric
having dozensof alternative and incommensurableclassifications
.
system of measurementis not the only possible system of measurement
all
for
not
is
.
It
on
it
from
purposes.
God did not deliver
equally good
high
different
constructed
systems
very
Given different contingencies, we might have
. In fact, we did. We have two systemsof weight, distance, and so on
'
the metric and the English. I cant speakfor others, but I find the presenceof
thesetwo systemsa persistentirritation . I would hate to think how inconvenient
having a dozen such systemswould be.
The same goes for naming and organizing the elements in the periodic
table. All sorts of different ways of organizing the elementswere proposed.
The end result that any.one who ever took an introductory courseinchemis
- fashion
try stared at for hours on end arosein a hit or miss, highly contingent
of
names
the
. In fact, we are only now getting around to formalizing
the elements with atomic numbers from 101 to 109 (see Pureand Applied
enableus to makethesignificantquestions
throughwhichwe extendsuccessful
" 79
schemata
moretractable
in original). Thus, we canrejectspecies
( , emphasis
definitionsif they areredundant
, boring, or wrongheaded
. A species
division
is redundantif it fails to "makeany significantquestionsmoretractable
." A
"
speciesdivision is boring if it doesnot help us to pursuefurther goals."
Finally, a speciesdivision is wrongheadedif the schemataon which it is
based"involvepresuppositions
we believeareincorrect" (Stanford1995, 80).
In his ensuingdiscussion
, Stanfordclarifieshis criteriaby applyingthemto
. Creationismis wrongheaded
particularcases
. The explanatoryschemata
of
"
the creationists"restuponsubstantiallymistaken
presuppositions(Stanford
1995, 80). For example
, they attempt to argueaway the implicationsof
carbondatingby postulatinga directionalchangein the rate of radioactive
decay.They explainthe patternsto be foundin the fossilrecordin termsof
which organismscould climb, swim, or fly the highest
during the Great
Flood. I needgo on no further. Accordingto Stanford
is boring.
,
pheneticism
'
The "naturaldependencies
identifiedby the pheneticists
OperationalTaxonomicUnits aretrivial andunhelpfulin pursuingany practicalend" (but see
Dupre, chapter1 in this volume). The pheneticists
would find this objection
especiallydamningbecause
they takethe practicalusefulnessof their classi
ficationsto be one of their chiefvirtues. In defenseof the phenetic
species
, even Mayr (1981), the chief opponentof pheneticism
concept
, finds the
establishment
of phenaan importantfirst stepin the recognitionof genuine
.
species
In the discussion
of his examples
, Stanfordmakesit clearthat applications
of his criteriaarehistoricallycontingent
. Rightnow, the creationistandpheneticspeciesconceptscanbe rejected
, but in the pastthey might well have
led to scientificprogress
. For example
, the appealof pheneticism
restedon a
common
if
, not universal
, convictionthat somethingout thereexists
quite
that answersto the name"overallsimilarity." If Atran (1990) is
right, this
modeof perceptionmayhavea significantgeneticbasis.If nothingelse, the
pheneticistsshowedthat, contrary to their own goals, no such thing as
overallsimilarityexists. If suchbright, hardworking
, and creativescientists
in a period of twenty yearsor so could not comeup with
anythingeven
approximatingoverall similarity, it is very likely not to exist in the first
'
place. In Cuviers day; theremight havesomethingto sayfor structuralism
,
but lessso today.
I. Moni ~ , Pluralism
, Unity and Diversity
cepts. To begin, scientists value the universalityof their concepts (but see
Boyd, chapter 6 in this volume). For example, any de6nition of elementmust
apply to all matter, not to just a subset. Physicistswould be lessthan pleased
if their element concept applied only to metals or to nonradioactive substances
. Biologists would like their preferredspeciesde6nition to apply to all
organisms, not just some, but fulfilling this desideratumhas proven to be
very difficult. For example, the biological speciesconcept applies to only
those organismsthat reproducesexually, at least on occasion.
.
Biologists, like all scientists, would prefer that their conceptsbe applicable
that
the
circumstances
,
way
Perhapsthey neednot be totally applicablein all
At the
better.
the
are
the
more
but
,
,
they
applicable
operationists propose
be
never
can
a
that
in
such
least
they
applied
, defining concepts
way
very
runs counter to the testability criterion of science. The testing may be difficult
, indirect, and fallible, but it must be possible. Philosophersare usually
content once we have decidedthat a particular concept in scienceis in principle
applicable, but scientistswant more, much more. They want grouping
criteria- criteria that enable them to decided whether two or more organisms
belong qr do not belong to a speciesin a significant percent of the
cases. They also want ranking criteria- criteria that enable them to decide
whether a taxon is a subspecies
, species, or genus (Mishler and Brandon
1987).
Finally, philosophers of scienceare currently convinced that to be useful,
all scientific conceptsmust be theoreticallysignificant. They must function in
a significant scientific theory. Becauseno scientific concept can be totally
theory free, the issue becomes which theory colors which concepts (see
Dupre, chapter 1, this volume). Next we must rank these theories according
'
to how fundamentalthey are. To use the traditional example, Newton s
theory of universal gravitation (once fixed up) is more fundamental than
'
Kepler s laws of planetary motion (once fixed up). On this view, those concepts
required by the most fundamental theories take precedenceto those
is
conceptsrequired by less fundamental theories. And if theory reduction
the
to
reduced
can
be
possible, all of these various upper level theories
lower-level theories. With respect to the connection between process
theories and the patterns discerniblein nature, many scientistsdisagreewith
have
philosophers about the primacy of theories. Perhaps philosophers
have
not.
scientists
They
worked their way free of inductivism, but many
insist that all scientific investigations must begin with direct, theory-free
observation and proceed as cautiously as possible, avoiding idle speculation
(seethe papersin Claridge, Dawah, and Wilson 1997).
In my paper, I grouped the speciesconcepts that I evaluated into three
families. The first family includes speciesconcepts that determine species
statusin terms of some.form of similarity- for example, traditional morphotogical species concepts, the phenetic species concept, as well as certain
, all of these speciesconcepts
molecular concepts. Until the past few decades
were typo logical in the sensethat a single list of dtaracters was developed
ofSpecies
Hull:OnthePlurality
/
I. Monism, Pluralism
, Unity and Diversity
data, I could not produce the result I had anticipated. All of the
species
conceptsI evaluatedscoredabout the same! One reasonfor this outcome is
that the most easily applied conceptstend to be those with the least theoretical
commitment, whereasthose conceptsthat produce theoretically significant
speciestend to be the most difficult to apply. Universality, in its
turn, does not covary with either theoretical significanceor easeof
application
. Some theoretically committed species concepts, such as the monophyletic speciesconcepts, apply to all organismsand are moderately easy to
apply.
The grudging conclusion of my paper (Hull 1997) is that none of the
speciesconcepts that I evaluated are all that superior to the others- that
is, if universality, applicability, and theoretical significance are
weighted
equally. However, in this samevolume, Mayden (1997) set himself the same
task, but cameto quite a different conclusion. Mayden's goal was to find the
primary speciesconcept. The differencesbetween our papers is instructive.
First, Mayden recognizes twenty -two different species
concepts, and he
combines as single concepts several formulations that others take to be
separateand distinct speciesconcepts. For example, the two formulations of
the phylogenetic speciesconcept that I classifiedas two
separatespecies
conceptsare classifiedby Mayden as a single speciesconcept. Next , he evaluates
these twenty -two conceptsaccording to their " convenience, accuracy,
"
precision, and the successfulrecovery of naturalbiological diversity (Mayden
1997, 381).
Finally, Mayden evaluateshis speciesconceptson severaladditional criteria
that are a good deal more specific. To serve as the primary speciesconcept
, a concept must be theoretically significant and include sexual, asexual,
and hybrid species
; it must be a nonrelational lineage concept that treats
as
individuals
rather than as classes
species
; and it must place no constraints
on necessaryattributes that a speciesmust possessin order to be validated.
As the primary speciesconcept, it need not be operational becauseother,
secondary speciesconcepts provide the operational basis for this primary
speciesconcept. For the primary speciesconcept, theoretical significanceis
of primary importance. Only after a speciesconcept passesthis test do the
other criteria come into play. The only speciesconcept that fulfills all of
these criteria is the evolutionary species concept as reworked by Wiley
(1981) and by Wiley and Mayden (1999).
The main reasonwhy Mayden and I cameto very different conclusionsin
evaluating various speciesdefinitions is that I combined theoretical significance
and operationality. A speciesconcept might score quite highly on
theoretical significance, but if it was not very operational, it endedup with a
'
mediocrecumulative score. Mayden, to the contrary, took theoretical significance
as necessaryand then rankedtheoretically significant speciesaccording
to his other criteria, including operationality. A secondreasonwhy Mayden' s
conclusion is so different Horn mine is that he included more substantive
criteria. Yes, I think that any adequatespeciesconcept must treat speciesas
I. Mo
m, Pluralism
, Unity and Diversity
CONCLUSION
"
"
Postmodemistshave made positivists all-purpose whipping boys, usually
parodying their views in the process. Other authorshave also joined in these
parodies. For example, just about everyone claims that attempting to demarcate
sciencefrom nonscienceor pseudoscienceis terribly wrongheaded, but
? Someof it is very bad science;
then what do we do about creation science
some of it is not scienceat all (Reisch 1998). Has philosophy really become
'
so sophisticatedand nuancedthat we cant distinguish between scienceand
? My fellow philosophersare likely to respond that courts of
creation science
law are not graduate seminars, and they insist on limiting themselvesto
is irrelevant.
graduateseminars. The rest of
' society
In deciding which speciesconceptsto take seriously, we seeminextricably
caught up in the issue of what counts as genuine scienceand what not.
Kitcher rejectsthe creationist and pheneticspeciesconceptsfor philosophical
reasons. The problem with pheneticism is that it comes into conflict with
his philosophical monism of the moment- namely, that no such things as
theory-free observations, let alone concepts, exist. Hence, any attempt to
deAne the species category in a theoretically neutral way is impossible.
Kitcher is putting his bet on this philosophicalposition prevailing for awhile.
'
I share Kitcher s prediction. Not only will philosophers continue to value
theoretical significancehighly , but I am betting that an increasingnumbersof
systematistswill come to sharethis conviction as phylogenetic cladists win
out over their pattern cladist brethren.
But theoretical significanceonly narrows the number of philosophically
acceptablespecies concepts. For those of us who are more monistically
inclined, traditional philosophical criteria alone are not sufficient forevaluating
.
speciesconcepts. On this score, thereis more to sciencethan philosophy
exists
Right now, an extremely powerful, well articulated theory actually
in biology - evolutionary theory. This theory places constraints on both
other
speciesdefinitions and traditional philosophical desiderata. Of course,
articulating
in
little
a
are
headway
theories are possible. Some scientists
making
alternative ways of viewing the living world , but until the promise of
thesealternativesis realized, we cannot treat them on a par with evolutionary
th.eory. In connection with our understanding of the evolutionary process,
Sober (1984a, 335) is ilguessingthat [the] species-are-individuals perspective
' s conviction but
,
':Villwinll (but seeWheeler and Platnick 1999). I shareSober
.someday, way down the road, this perspectiveon speciesmay be overturned.
The possibility of future alternatives is not, however, a sufficiently strong
reasonfor acceptingthe pluralist philosophicalmonism of the moment.
/
Hull: On thePluralityof Species
/
I. Monism
. Pluralism
. Unity andDiversity
ACKNOWLEDGMENTS
I wish to thank John Dupre , Marc Ereshefsky, Kim Sterelny , M . H . V . Van
Regenmortal , Ed Wiley , Rob Wilson , and an anonymous referee for reading
and commenting on this paper .
NOTFS
1. As Dupre remarkedin responseto the examplegiven, alwaysusing virus-impenetrable
of the
rubberswoulddefeatthe AIDSepidemicin two ways: it wouldpreventthe transmission
would
also
the
but
in
the
it
virus to new hostsvia sexualintercourse
,
process
preclude birth
suchprophylacticswould alsobe spermimpenetrable
of new humanbeingsbecause
, a cure
.
decidedlyworsethanthedisease
2. Kitcher(1987, 187) enlistsMishlerandDonoghue(1982) asfellow pluralists
, but they protest
. "Kitcher'S (1984a
, 1984b
) brandof pluralismimpliesthat thereare many possibleand
complex
for a given situation(say, the Drosophila
mtllmogaster
pennissiblespeciesclassi6cations
. In contrast
, Mishlerand
), dependingon the needsandinterestof particularsystematists
'
-purposeclassi6cation
Donoghues (1982) brandof pluralismimpliesthata singleoptimalgeneral
.
in
each
situation
the
criteria
existsfor eachparticularsituation
, but that
may well be
applied
ton (1989) a Kitcher-stylepluralist.He
" (MishlerandBrandon1987,403). Nor is Temple
different
- cohesiveness
andonecharaderonly is relevantto speciesstatus
thinksthat onecharacter
cancontributeto thiscohesiveness
eventhoughdifferentmechanisms
(seealsoDonoghue1985
andEreshefsky
1992).
'
. Because
3. Pluralismis at bottom incompatiblewith WheweUs conswenceof inductions
a surprise
it
comes
as
of
this
MichaelRuseis amongthe mostenthusiastic
,
prindple
supporters
to find him on Ereshefsky
, albeitas highly conservative
' S list of pluralists
pluralist. Although
into groups
, and
Ruse(1987, 238) arguesthat thereare"differentwaysof breakingorganisms
on thispoint.
" I happento thinkthathe is far too sanguine
theycoinddtf
mirror the decisions
4. I amawarethat the dedsionsI havemadeat this higherlevelof abstraction
to
considerations
.
historical
othershavemadeat thelevelof species
My
allowing
concepts
overridesimilaritywith respecito how I classifyspecies
conceptsis likely to imply something
level.
I find mostimportantat the species
aboutwhichconsiderations
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. Ada BiotheoretiC
R 45,
Shanahan
. T. (1997
, antirealism
, and the units of selection
) . Pluralism
117- 11
. 6.
. New York: Columbia University Press.
Simpson, G. G. (1961). Principlesof animal ta. ronomy
Sober, E. (19848). Discussion: Sets, speaes, and evolution. Comments on Philip Kitdter' s
"
." Philosophyof Science
Sl , 334- 341.
Species
Sober, E. (19Mb ). Thenatureof selection
. Cambridge, Mass.: MIT Press.
Sober, E. (1993). Philosophyof biology. Boulder, Colo.: Westview Press.
Sober, E., and D. S. Wilson. (1994). A aitical review of philosophical
work on the units of selec-
tion problem
. Philosophy
61, 534- 555.
of Science
Sober
: theevolution
andps.vchoiog
behaoiors
.
, E., andD. S. Wilson. (1~ 8). Untoothers
.vof unselfish
: HarvardUniversityPress
.
Cambridge
62,
Philosophyof Science
/
I. Monism, Pluralism
, Unity and Diversity
.. in
varies
Thereis nothing morecommonthan that the meaningof an expression
as a symptomand now as a criterion
is now considered
sucha way that a phenomenon
of a stateof affairs. And thenfor the mostpart in sucha casethe changeof
which allow exact
. In scienceit is usualto turn phenomena
meaningis not noticed
is
then inclinedto
one
and
an
criteria
;
into
measurements defining
of
expression
think that now the genuinemeaninghas beenfound. An enormousnumberof
arisein this way.
confusions
- Wittgenstein (1967)
Given the proliferation of speciesconcepts in recent years, it might seem
that the speciesproblem- the difficulty of reaching agreement about the
de6nition of the speciescategory- is as far from being solved as it has ever
been. On the contrary, the speciesproblem has, for the most part, already
been solved. Despite the considerablediversity among contemporary views
on species
, all are encompassedby a single, general concept that equates
with
segmentsof population level lineages. Becausethis population
species
modem ideasabout species
all
, it bearson
underlies
virtually
lineageconcept
careto ask
would
one
that
almost every historical and philosophicalquestion
about those ideas, including the major themesof this volume. In this essay, I
describethe general concept of speciesas segmentsof population lineages
es the diversity of modem views on species. I
and show how it encompass
then discusstwo assumptionsthat, despite widespreadagreementabout the
,"lead to incompatible speciesconcepts. I show how
generalnature of species
, which entails reconsideringthe defining
eliminating one of those assumptions
the
of
,
speciescategory effectively solves the speciesproblem.
properties
I then use this perspectiveto clarify severalphilosophical issuesconcerning
, including the role of the speciesconceptin biology , the individuality
species
of species
, whether the speciescategory is a relational concept, monistic
ve~sus pluralistic views of species, and speciesrealism. Finally, I briefly describe
the history of the lineageconceptof species.
.
THE GENERAL LINEAGE CONCEPT OF SPECIFS
In a previous paper (de Queiroz 1998), I argued that all modem species
I
conceptsare variants of a single general concept of species. In that paper,
Lineages
I have used the term lineage(de Queiroz 1998; seealso Simpson 1961, Hull
1980) for a seriesof entities forming a single line of direct ancestry and descent
. For example, a lineage can be traced Horn a given organism backward
though a parent, grandparent, great-grandparent, and so on, and forward
through a child, grandchild, great-grandchild, and so on. Biological entities at
several different organizational levels form lineages. Thus, biologists speak
of gene lineages, organelle lineages, cell lineages, organism lineages (as
describedin the above example), and population lineages. Becauseentities
that form lineages often make up, or are made up of, entities at different
organizational levels, the same is also true of the lineages themselves. An
organism lineage, for example, is (often) made up of multiple cell lineages,
and multiple organismlineagesmakeup a population lineage.
Lineagesin the sensedescribedabove are unbranched; that is, they follow
a single path or line anytime an entity in the serieshas more than one direct
descendant(figure 3.la ). Consequently, lineagesare not to be confusedwith
clades, clans, and clones- though the terms are often used interchangeably
in the literature} Clades, clans, and clones include all paths or lines of
descentfrom a given ancestorand thus are branched, which is to say that
they are composedo,f multiple lineages(figure 3.lb ). Moreover, clades, clans,
and clonesare monophyletic by definition; a clade, for example, is definedas
a monophyletic group of species..3Lineages, in contrast, can be paraphyletic
or even polyphyletic in terms of their lower-level components(see"Phyly " ).
They can even be paraphyletic in terms of their segmentsat the sameorganizationallevel. Thus, the later segmentsof a lineage commonly sharemore
. recent common ancestorswith separatebut recently diverged lineagesthan
they do with earlier segmentsof their own lineage(figure 3.2).
Species
(8) lneages
~
(b) c~
, clans
, orclones
)~
contrasted
with clades
, andclones(afterde Queiroz1998). All of
, clans
Figure 3.1 Lineages
th~ samephylogenywith differentlineageshighlightedin (a)
the branchingdiagramsrepresent
anddifferentclades
, clans
, or cloneshighlightedin (b). Noticethat the lineagesareunbranched
andeithernestedor
andpartiallyoverlapping
, clans
, or clonesarebranched
, whereastheclades
for pathsbeginningat various
canberecognized
. Additional(partial) lineages
mutuallyexclusive
internalnodes
.
/
de Queiroz: The GeneralUneageConcept of Species
(a) sexualreproduction
II
(b) asexualreproduction
Figure 3.3 Population lineages in sexually and asexually reprodudng organisms (adapted
from Brothers 1985). (a) Under sexual reproduction, organism lineagesare connected
through
the processof reproduction itself (representedby connections[ A] betweenvertical lines) to form
a population-level lineage. (b) Under asexualreproduction, no such reproductive connections
exist, but it is possiblethat the organism lineagesare bound into a population lineage by other
processes (representedby the spatial localization of the organism lineages). In both diagrams,
organismsare representedby vertical lines.
1974)- that is, to groups of organism lineages that are united to form
higher-level lineages. The formation of population-level lineages is most
evident in the case of biparental organisms, where the process of sexual
reproduction continually reconnectstemporarily separatedorganismlineages
to form a unified nexus (figure 3.3a). At least someauthors, however, believe
that uniparental organismsalso form species(figure 3.3b). Becausea general
speciesconcept (i.e., one that can encompassthe diversity of modem views
about species) must allow for this possibility, I use the term populationin the
general senseof an organizational level above that of the organism, rather
~than in the specificsenseof a reproductive community of sexual
organisms.
The population level is really a continuum of levels. Lineagesat lower
levels in this continuum (e.g., demesor demelineages) often separateand reunite
over relatively brief time intervals. Toward the other end of the con. tinuum,
lineage separation is more enduring and can even be permanent.
/
I. Monism, Pluralism, Unity and Diversity
Thus, when I say that a lineage is unbranched, I do not mean that it can
never exhibit internal branching; however, any such branching that it exhibits
would have to be judged as ephemeral. In any case, most authors
equatespecieswith lineagestoward the latter end of the continuum, though
they differ with regard to the precise point that they consider the line of
demarcationfor species.
Under the lineage concept of species, speciesare not equivalent to entire
population lineages, but rather to segmentsof such lineages. Just as a cell
lineage is made up of a seriesof cells and an organism lineage of a seriesof
organisms, a species(population) lineage is made up of a seriesof species.
Not just any lineage segment qualifies as a species, however. Instead, a
species corresponds with a lineage segment bounded by certain critical
events. Authors disagree, however, about which events are critical.
In short, speciesare segmentsof population-level lineages. This definition
describesa very general conceptualizationof the speciescategory in that
it explains the basic nature of specieswithout specifying either the causal
processes responsiblefor their existenceor the operational criteria used to
4
recognize them in practice. It is this deliberate agnosticismwith regard to
causalprocesses and operational criteria that allows the concept of species
just describedto encompassvirtually all modem views on species, and for
this reason, I have called it the generallineageconceptof species(de Queiroz
1998).
THE UNITY AND DIVERSITY OF SPECIFSCONCEPTS
By identifying the unity of contemporary speciesconcepts, the general lineage
concept of speciesprovides a context for understandingtheir diversity .
Statedin the most generalterms, that diversity results from different authors
emphasizingdifferent aspectsor properties of the entities conforming to the
general lineage concept. In the remainderof this section, I describesome of
the major differencesamong contemporaryideasabout speciesas well as the
relationship of those ideas to the general lineage concept. This exerciseis
not intended to describethe diversity of such ideasexhaustively, but rather
to illustrate that even what seem to be the most fundamental differences
among contemporary views on speciesare compatible with the generallin
eageconcept.
/
de Queiroz: The GeneralUneageConceptof Species
1985). These two classesof speciesdefinitions are not at odds with one another
, and both are entirely consistent with the general lineage concept of
species. As has been noted by several authors, a lineage (at the population
level) is a population extendedthrough time, whereasa population (in itself )
is a short segment- a more or lessinstantaneouscross section- of a lineage
(see Simpson 1951, 1961; Meglitsch 1954; George 1956; Newell 1956;
Rhodes1956; WestoIl1956 ).s Thus, definitions that equatespecieswith populations
consider the entities of interest over relatively short time intervals,
whereasthosedefinitions that equatespecieswith lineagesconsiderthem over
longer time intervals. In other words, the two categoriesof definitions do
not describedifferent conceptsof species
; they merely describetime-limited
and time-extendedversions of the samespeciesconcept.
Process es and Produds
/
I. Monism
, Pluralism
, Unity andDiversity
de QueiroZ:TheGeneralUneageConceptof Speaes
/
I. Monism. Pluralism, Unity and Diversity
(b) bifurcation
Anagenetic
(c) blastation
CIedOg8l8dC
genetic model equatesspeciation with lineage change, whereas the cladogenetic model equatesspeciationwith lineage splitting . The other main difference
between the models concernshow speciesare bounded relative to
speciationevents (however those events are defined). Under both the phyletic transformation and bifurcation models, species correspond precisely
with the segments of lineages between speciation events (though what
counts as a speciation event differs for the two models), whereasunder the
blastation model, speciescorrespondwith lineage segmentsthat originate in
speciationevents but do not necessarilyterminate in such events. The point
is that all three of thesemodels equatespecieswith lineage segments.8
Phyiy
Annther major difference among contemporary views on speciesconcerns
what might be termed phyly- that is, whether speciescan or must be monophyletic, p~ aphyletic, or polyphyletic . Different authors allow all three types
. de
Queiroz: f Ite GeneralUneageConcept of Species
monophyletic
-p.po
a-p.
"
"
Figure 3.5 Paraphyly andpolyphylyof speciesin termsof their components
, orgagenes
nelles
. (a) The species
, or organisms
on the right sideof the split is "paraphyletic
" in the sense
that someof its lower-levelcomponents
sharemorerecentcommonancestors
with the components
of anotherspeciesthanwith othercomponents
of their own species(butseenote9). (b)
The species
on the left sideof the split is polyphyleticbecause
someof its lower-levelcomponents
are only distantlyrelatedto one another
, coalesangin a remoteancestralspecies(not
shown). In both diagrams
, gene,organelle
, or organismlineagesthat havesurvivedto themost
recenttimearehighlightedso thattheirrelationships
canbeseenmoreeasily.
of species(e.g., Neigel and Avise 1986); or only paraphyletic and monophyletic species(e.g ., Brothers 1985, Crisp and Chandler 1996), or only
monophyletic species(e.g., Rosen1979, Mishler and Donoghue 1982). Other
authors argue that the conceptsof phyly do not apply to individual species
but only to groups of species(e.g., Wheeler and Nixon 1990; also seenote 3).
Some of the differencesregarding speciesphyly reflect differencesin the
level of organization under consideration. Thus, phyly in terms of component
genes or organisms, (as discussedby Neigel and Avise, 1986), should
not be confusedwith phyly in terms of componentpopulations (as discussed
by Bremer and Wanntorp, 1979). Paraphyly9 and polyphyly in the former
sense(figure 3.5) appear to be common initial stages in the divergence of
population-level lineages(Neigel and Avise 1986) and, in the caseof poly phyly , when species arise as the result of hybridization. Most authors
presumablywould not deny that speciescan be either paraphyletic or poly phyletic in this sense(but see Baum and Shaw 1995). In contrast, there are
probably few (if any) contemporary biologists whose concept of species
includes entities that are polyphyletic in terms of their component populations
- that is, who would
recognizeas parts of a single speciestwo or more
populations that are not particularly closely related to eachother (figure 3.6a;
Sosef 1997). Similarly, at least some authors (e.g., Rosen 1979, Bremer and
I. ~
sm. Pluralism
., Unity andDiversity
Speaesa
speaesb
(DOlvDhvletic
)
Speciesc
Specleax
(paraphyletlc
)
Spedesy
Pigure 3.6 Polyphyly and paraphyly of speciesin terms of their component populations. (a)
Polyphyly of speciesb, whose two component populations both share more recent common
ancestorswith heterospecificpopulations than with one another. It is assumedthat the two
populations of speciesb are consideredconspecific becauseof convergent rather than retained
ancestralcharacters
, so that their common ancestralpopulation would not be consideredpart of
b.
(b)
species
Paraphylyof speciesx, one componentpopulation of which is more closely related
to speciesy than to the other population of its own species
. In both cases
, phylogeniesof lowerlevel components(e.g., genes) are shown within the population lineages, with lineagesthat survived
to the most recent time highlighted.
I. ~ nism
, Unity andDiversity
, Pluralism
THECAUSFS
OFTHESPECIES
PROBLEM
ANDA SIMPLE
SOLUTION
Despite nearly universal acceptance of the general lineage concept of species,
at least two factors prevent a general consensus about the definition of the
species category . One of these factors compromises universal acceptance of
the general lineage concept itself ; the other creates incompatibilities among
'
the concept s numerous variants . Consequently , these factors are critical to
solving the species problem .
Onto logical and Taxonomic
Categories
The first factor concerns a basic assumption about how the species category
is interpreted , which bears on acceptance of the general lineage concept
itself . One interpretation is that the species category is an onto logical category
(see Ghiselin 1997 )- that is, one of the fundamental categories of biological
existence (other such categories are the cell and the organism ). The
other interpretation is that the species category is a taxonomic category that is, a level or rank in the Linnean hierarchy of taxonomic categories
(other such categories are the genus and the family ). These alternative interpretations
are not necessarily at odds with one another , but they often underlie
at least partially incompatible views on speciesd . Boyd , chapter 6 in this
volume ).
The interpretation of the species category as an onto logical category is
implicit in the general lineage concept of species, which equates the species
category with the onto logical category whose members are the biological
entities known as population lineages. On the other hand , the interpretation
of the species category as a taxonomic category is implicit in its use in biological
taxonomy , which equates the species category with one of the taxo nomic categories in the Linnaean hierarchy . These two interpretations have
several possible relationships with one another . ( 1) All of the taxonomic
categories are artificial ; none of them corresponds with an onto logical category
(cE. Ereshefsky, chapter 11 in this volume ). (2) Each taxonomic category
corresponds with a different onto logical category ; the species category corresponds
with one onto logical category , the genus with another , the family
with yet another , and so on . (3 ) All the taxonomic categories apply to the
same onto logical category , the members of which form nested hierarchies ;
t ~ 'e various taxonomic categories represent differe ~t ranks or levels in those
nested hierarchies. (4) The various taxonomic categories represent some
combinati .on of the first three alternatives .
I. ~ nism, Pluralism
, Unity and Diversity
/
de Queiroz:TheGeneralLineageConceptof Species
UENCFS
PHILOSOPHICAL
CONSEQ
In this section, I examine the implications of the perspective developed in
the previous sections for various philosophical issues concerning species,
including severalof the major themesand topics of this volume. My purpose
is to show how the general lineage concept, along with the reinterpretation
of the necessaryproperties of the speciescategory, either clarifiesor resolves
other issuesabout species.
Species and the Representation of Biological Diversity
I. ~
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. Pluralism
, Unity and Diversity
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de Queiroz: The GeneralUnease Concept of Species
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I. Monism, Pluralism
, Unity and Diversity
Species Individuality
/
: TheGeneral
deQueiroz
LineageConcept of Species
/
I. Monism, Pluralism
, Unity and Diversity
segments. This is not to say that specieshave regular and integrated ontogenleslike those of many organisms, but merely that they go through cycles
of genesisand termination, with other changesin between. Indeed, because
different processes are responsiblefor the unification of organisms(e.g ., cell
membranejunctions, cell to cell adhesion) and that of species(e.g ., interbreeding
, selection), care should be taken when drawing analogiesbetween
the two kinds of individuals. On the other hand, becausethe implications of
organismalindividuality are more familiar to us, suchanalogiesoften greatly
facilitate our ability to conceptualizethe implications of speciesindividuality .
Thus, both the similarities and differencesbetween organisms and species
provide insight into specieslife cycles.
With regard to origins, an obvious analogy can be made between reproduction
"
"
by fission and speciationby bifurcation (see Models of Speciation ),
where new species arise from large subdivisions of an ancestral species
(reviewed by Bush 1975). In both cases
, the descendantsoriginate from
often
more
or
less
of
(
major
equal) portions their ancestors. Similarly, reproduction
"
by budding correspondswith speciationby blastation (see Models
"
of Speciation ), where a speciesoriginates from a small founder population
(seeBush 1975). In both cases
, the descendantarisesfrom a small portion of
its ancestor. In all of these modes of genesis(fission, bifurcation; budding,
blastation), the production of new organisms or speciescoincides with lineage
splitting . If speciesare like organisms, then the model of speciation
"
"
by phyletic transformation (see Models of Speciation ) would seem to be
invalid (e.g ., Hennig 1966, Wiley 1978). A single organism changesconsiderably during the course of its life (e.g., zygote to adult human), so the fact
that a specieschangesduring its existencedoes not require that it changes
into a different species. Indeed, a speciesshould be able to change indefinitely
and still remain the samespecies,provided that the changeis more or
less gradual and continuous. Situations in which eachorganism or speciesin
a seriesproducesa single descendantvia budding or blastation should not be
confused with unbranchedlineages. Although we tend to think of a such
successionsas linear or unbranchedin the caseof organisms, they are really
branched if parent and offspring coexist temporally. On the other hand, if
a parent dies more or . less simultaneouslywith the propagation of a single
O
offspring, perhaps the lineage can be consideredunbranched} Similarly, if
an unbranchedpopulation lineagepassesthrough a severebottleneck, which
is similar in many respectsto a founder event, perhaps it is justifiable to
consider the lineage segmentson either side of the bottleneck as different
species.
. Despite the possibility of phyletic transformation, differencesbetween the
component organismsin earlier and later parts of an unbranchedpopulation
,lineage - even those that affect other biologically significant properties (e.g .,
the ability to interbreed)- are not particularly relevant to the question of
whether their respective lineage segmentsconstitute the same or different
species. Undifferentiated cells in an early embryo are not particularly similar
/
de Queiroz
: TheGeneralLineageConceptof Species
to the differentiated cells that makeup the sameorganismlater in its life, and
perhaps they would not be integrated into the later organism if given a
chance. Even changesin the emergentproperties of lineage segmentsdo not
necessarilyimply that they are different species. Some organisms change
from carnivoresto herbivores, or from femalesto males, during a single turn
of an organism life cycle, so a speciesshould be able to change from panmictic to subdivided, for example, during a single turn of a specieslife cycle.
Regarding termination, organisms sometimesend by ceasingto function
as integrated wholes- that is, by death. Speciescan end in an analogous
manner, normally termed extinction.And just as certain component cells can
continue to live after their organism dies, certain component organismscan
continue to live after their speciesbecomesextinct. The most obvious example
is a speciescomposed of organisms with obligate sexual reproduction
and separatesexes(and no sex-changing abilities) in which the only surviving
, organismsend
organismsare all membersof the samesex. In other cases
into
more
than
one whole, that is, by fission. The analogous
by separating
situation for speciesis ~ifurcation. Becausethe ancestorin both casesis no
longer identifiable after the lineage splits, it is consideredto terminate at the
!
splitting event} This is not to say that ancestors necessarily terminate
whenever lineagessplit. When the split is highly unequal, as in the casesof
organism budding and speciesblastation, the ancestorcan be consideredto
persist.
If there is a differencebetween organismsand specieswith regard to lineage
- that is,
splitting, perhaps it is the frequency of intermediate cases
casesin which the split is only moderately unequalso that it is ambiguousas
to whether the ancestorpersists. Organismal reproduction appearsstrongly
polarized into fission and budding modes, with few intermediate cases. In
contrast, bifurcation and blastation modes of speciationappear to be opposite
ends of a continuum in which the intermediatecasesare far more common
, particularly if extrinsic barriers are a common causeof speciation. In
any case, considering the life cycles of specieshelps us to formulate a fuller
description of the general lineage speciesconcept. Speciesare not just any
segmentsof population-level lineages; they are the segmentsof populationlevel lineagesthat correspondwith a single turn of the life cycle, from genesis
to termination.
An important difference between speciesand organisms concernsfusion.
Separateorganism lineagesrarely fuse as wholes (but see below). Even the
continual merging of sexual organism lineages usually involves only the
transfer of genetic material between cells or the union of specializedcells
(gametes); the organisms themselvesretain their separateidentities. In the
caseof population lineages, fusion appearsto be much more commonAl though certain speciesdefinitions are based on properties that would seem
to be correlated with irreversible separation(e.g., intrinsic reproductive isolation
), resulting in speciestaxa that are more like organismsregarding their
likelihood of fusion, there are no guarantees
. For example, premating barriers
/
I. Monism, Pluralism
, Unity andDiversity
71
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I. Monism, Pluralism, Unity and Diversity
Pluralism
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: TheGeneralLineageConceptof Species
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I. Monism, Pluralism
, Unity and Diversity
authors take this statementto mean that the existenceof speciestaxa is not
independent of the theoretical interests of biologists (e.g ., Stanford 1995).
Others take it to mean that there is no common and unique identifying
property of the speciescategory (e.g., Ereshefsky1998, chapter 11 in this
volume).
The secondform of antirealismis directly contradicted by the generallin eage concept of species, which is based on the identification o~ a common
and unique property of speciestaxa. All speciesare segmentsof population'
level evolutionary lineages. This position is consistent with Ereshefsky
s
(1998) view that speciesare genealogicalentities, but Ereshefskyarguesthat
being a genealogicalentity does not suffice as a unifying feature of species
becauseit also appliesto genera, families, and so on- that is, to higher taxa.
The apparentproblem is readily solved in the context of the general lineage
concept by recognizing a distinction between two different kinds of genea"
"
logical entities: lineages(as defined in this essay; see Lineages) and clades.
Speciesdiffer from higher taxa in that speciesare lineages(or more properly,
lineage segments), whereas higher taxa are clades (i.e., groups of species
sharing an exclusivecommon ancestry). The sameconclusionholds if (some)
higher taxa are allowed to be paraphyleticgrades.
Both forms of antirealism rest on a form of speciespluralism that views
alternative descriptions of the speciescategory as irreconcilable definitions
- a
position that in turn rests on the interpretation of certain contingent
properties of lineages as necessaryproperties of species. This position is
what allows antirealiststo concludethat a single organismcan belong simultaneously
to different types of speciesand thus to different speciestaxa (e.g .,
Ereshefsky1998, chapter 11 in this volume). If, for example, intrinsic reproductive
isolation is interpreted as a necessaryproperty of species
, it will lead
to the delimitation of one set of speciestaxa, and that set of speciestaxa will
likely differ (in terms of both the number of speciesand the assignmentof
organisms to speciestaxa) from the set of speciestaxa delimited under a
species definition that adopts a different property diagnosability, for
- as a necessary
property of species.
example
I have already shown how reinterpreting certain properties as contingent
rather than necessaryproperties of speciesresolves the conflict between
speciesmonism and speciespluralism. Becausethe antirealismargumentrests
on speciespluralism (or more accurately, antimonism), it is not surprising that
reinterpreting the significanceof those properties also nullifies the argument
against speciesrealism. If properties such as distinguishability, ecological
distinctiveness, and reproductive isolation (to mention only a few) are contirtgent rather than necessaryproperties of species, then they imply neither
alternative sets of speciestaxa nor the existenceof fundamentally different
kinds (ontological categories) of species. Instead, they merely imply that a
single speciescan belong simultaneously to several subcategoriesof the
general category species. For example, a speciescan simultaneouslybe phe, ecologically distinct, and extrinsically isolated from
netically -distinguishable
/
deQueiroz
: TheGeneralUnea~e Conceptof Species
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I. Monism
, Pluralism. Unity and Diversity
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de Queiroz: The GeneralUneageConceptof Species
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I. Monism, Pluralism
, Unity andDiversity
CONCLUSION
Fortunately, the situation is not as hopelessas it may appear. By losing sight
of the common thread running through virtually all modem views on
, both biologists and philosophershave overlooked a relatively simple
species
solution to the so-called speciesproblem. Virtually all modem biologists
have the samegeneral concept of species. Most of their disagreementsstem
from interpreting certain contingent properties of lineages as necessary
properties of species(i.e., speciescriteria), which leads to speciesdefinitions
that are incompatible both in theory (becausethey are based on different
necessaryproperties) and in practice (becausethey result in the recognition
of different speciestaxa). This situation fosters competition among alternative
speciescriteria and their associatedspeciesdefinitions, with each one
for
statusas thedefining property of the speciescategory. As a consequence
vying
, the common theme underlying all of the alternative views tends to
be obscured, and the perception of a major, unresolved problem concerning
the nature of speciespersists.
Recognizingthe common thread manifestedin what I have called the gene
ralline age concept of speciesrevealsa simple and straightforward solution
to the speciesproblem. All that is required is to drop the interpretation of
certain contingent properties of lineagesas necessaryproperties of species
,
and the speciesproblem will vanish. By reinterpreting what have been called
, or
speciescriteria as contingent rather than necessaryproperties of species
simply as different lines of evidence concerning the separationof lineages,
the conflicts among speciesdefinitions are removed. The definitions inquestion
are not alternative definitions of the speciescategory at all, but merely
descriptions of the diverse contingent properties of species. Consequently,
there is no longer any major unresolved problem regarding the nature of
speciesor the definition of the speciescategory.
The problem is that despite the existenceof a perfectly adequateconcept
and definition of species,most speciesare more like slime molds and sponges
than like highly organized and tightly integrated multicellular organismsat least in terms of their individuality . Not only can almost any part of a
speciesgive rise to a new lineage, but those new lineagesalso commonly reunite
after separating. Consequently, there will be many casesin which it
will be difficult to determinethe precisenumber and boundariesof species
just as it is difficult to determine the precise number and boundaries of
organismsin a fragmenting acrasialianpseudoplasmodiumor a multioscular
sponge. But such observationshave not led to the conclusionthat there is a
major unresolvedproblem concerningthe concept of the organism, and similarly
, they do not imply a major unresolvedproblem concerningthe concept
o~the speciescategory; .instead, they merely imply a practical problem about
establishingthe limits of speciestaxa in practice. Taxonomic traditions notwithstanding
, everything we know about speciestells us that they are inherently
difficult to circumscribe, particularly in the early stagesof divergence;
/
deQueiroz:TheGeneralUneageConceptof Species
that they are not always sharply distinct , easily recognized entities ; and that
unambiguous assignment of all organisms to species taxa will be difficult , if
not impossible . Attempting to solve this problem by treating operational
criteria as defining properties only aggravates the situation because it confuses
a purely practical problem with a theoretical one. The appropriate
solution to the practical problem is simply to accept the inherent ambiguities
'
of species boundaries (O Hara 1993 ). In any case, recognizing the conceptual
unity among modem views on species allows us to transcend their differences
. It helps us identify both the cause of and the solution to the species
problem , which clarifies a great deal concerning the concept of species itself
as well as its history and its significance for both biology and philosophy .
ACKNOWLEDGMENTS
I thank David Wake, Michael Donoghue, JacquesGauthier, Alan de Queiroz,
Bob O 'Hara, Allan Larson, Michael Ghiselin, David Hull , David Good,
Molly Morris , David Baum, Kerry Shaw, Jim Mallet , Marc Ereshefsky,Stewart
Berlocher, Darrel Frost, Ed Wiley , Ron Nussbaum, Linda Allison , and
Maureen Kearney for discussions(some many years ago) relevant to this
essay. I also thank Rob Wilson for the invitation to contribute .an essayto
this volume and for his commentson an earlier draft.
NOTFS
1. According to the proposed tenninology, a spedesconceptis an idea about the nature of the
entities that make up the speciescategory; a spedescrifm' on is a standardfor judging whether a
particular entity qualifiesas a member of the speciescategory, and a spedesdefinitionis a statement
and thus desaibing a Spedesconcept, usually in
spedfying the meaningof the term species
terms of necessaryand suffident properties.
2. Wilson (1995), for example, developed a view that equatesspecieswith what he called lineages
, but he used that term in a sensethat includesclades, clans, and clones.
3. The terminology for these entities has not been developed adequately. De Queiroz and
Donoghue (1988, 1990) used the term monophyleticto describethe generalclassof entities each
of whose membersconsistsof an ancestorand its descendants
, regardlessof organizationallevel.
They noted that the term .cladehad generally been used for monophyletic entities composedof
, and clonefor comparableentities at lower levels of organization. This terminology,
species
however, does not distinguish between monophyletic entities at severaldifferent organizational
levels below that of spedes, nor does it take into considerationthe distinction between diverging
and reticulating patterns of descentand the most common use of the term clonefor cases
involving asexual(nonreticulating) reproduction. O' Hara (1993) proposed using the term clan
for monophyletic entities at the organismallevel, regardlessof reproductive mode, but terms for
. other levels are currently lacking. Someauthors (e.g., Wheeler and Nixon 1990) object to using
'
monophyleticto describeentities below the spedes level, based on He Mig s (1966) distinction
- the former
between phylogenyand tokogeny
desaibing the descent of spedes, the latter the
descentof organisms. The term phylogeny
, however, is commonly used in a more general sense
to describedescentat various organizationallevels (e.g., " gene phylogeny" ).
/
I. Monism, Pluralism
, Unity andDiversity
The following tenninology makesmost of the distinctions that previous authors have considered
important, while minimizing discrepancieswith previous usage. Phylogeny(the genesisof
tribes) is usedfor (predominantly) branchingpatterns of descent, nuogeny(the genesisof bonds)
for (predominantly) reticulating patterns of descent. Both are general terms that can be used to
describedescent at various organizational levels, though each can be modified to specify the
level of organization (e.g ., gene phylogeny, organism nexogeny). Ramogeny(the genesis of
branches) is usedfor the descentof populations (from demesto species
), and tokogeny(the genesis
of offspring) for the descentof organisms. Correspondingterms for other levels are not proposed
here. The term phyly (-phyletic) can be usedin associationwith the prefixes mono, para, and
to
describe different patterns of descent (see Hennig 1966) regardlessof organizational
poly
level; the terms ramy (-rametic) and toky (-toketic) can be used forspeci Ac organizational levels
(e.g., monorametic, polytoketic). The generalterm entogeny(the genesisof things that exist), and
the related term enly (- mtetic
), can be used to encompassdifferent modes ( branchingand retiwould used for a single
clate) and levels (species
, organism
, etc.) of descent. Thus, tnonoentetic
ancestorand its descendants
, regardlessof whether that group is mutually exclusiveor partially
if the group is mutually exclusive(e.g., clades,
overlapping with other suchgroups, monophyletic
. retic if it is partially overlapping (e.g ., clans
clans/clones of uniparental organisms), and monone
within a biparental species
). Cladeis used for monophyletic (and monorametic) groups of populations
). Clan is usedfor monoenteticgroups of organisms, regardlessof
(from demesto species
reproductive mode - recognizing that clans of uniparental organisms will be monophyletic,
whereas clans of biparental organisms will be mononexetic. Clone is used for monophyletic
groups of asexually reproducing entities at or below the organismal level (e.g ., gene clones,
organeneclones, cen clones, although cen clonesin unicenularorganismsare also clans).
4. Two or more causalprocesses are implied: first, the processof descent, which is inherent in
the conceptof a lineage(at any level), and second, whatever processor processes unite organism
lineagesto form population lineages.
5. The concept of a population is not atemporal (truly instantaneous
) in that the processes
viewed as determining the limits of populations are temporal phenomena. For example, the pracessof interbreedingis commonly viewed as important in determining the limits of populations,
but as pointed out by O' Hara (1993), no population is composedof organismsthat are all interbreeding
at any given instant.
6. Severalauthors (e.g ., Brothers 1985, Templeton 1989) have emphasizedthat the exchangeof
genetic material among organism lineagesis not neatly dichotomized into asexualand sexual
reproductive modes, but insteadforms a continuum.
7. From the Greek blastos, meaningbud, sprout, shoot, or germ. The term is proposed to distinguish
-level process from the analogous organism-level process termed budding,
the species
which has also been used to designatethis model of speciation (e.g ., Foote 1996, de Queiroz
1998).
8. It should be noted that although thesegeneralmodels of speciationare logical consequences
of certain views on the properties of species(those properties used to define the models in
"
"Models of
Speciation ), the properties are most commonly stated without explicit referenceto
the modelsof speciationthat they imply . Other times, the models are implied by properties that
are a step further removed. For example, the view that every diagnosable lineage segment
representsa diHerent species(e.g., Nixon and Wheeler 1992) implies that speciation occurs in
model of speciation.
~ ranchedlineages, and this consequencein turn implies an anagenetic
9. Neigel and Avise (1986) used the term paraphylyfor casesin which certain gene or organism
lineages within a speciesshare more recent common ancestorswith heterospeciAc than with
.~onspeci6cgene or organism lineages(Agure 3.5). However, at least some of the speciesthat At
this desaiption are not paraphyleticin the senseof a group including an ancestorand some, but
not all of its descendants(e.g ., Hennig 1966, Wiley 1981), becausethe most recent common
/
de Queiroz:TheGeneralLin~ e Conceptof Species
ancestorof the lineagesin question is not part of the spedesidentified as paraphyletic, but of a
more distant ancestralSpedes. Spedesof this kind are polyphyletic rather than paraphyletic in
terms of their componentgenesor organisms; they differ from the spedesthat Neigel and Avise
consideredpolyphyletic only in the relative depth of coalescenceof their component gene or
organismlineages.
10. A similar situation involving polyphyly exists when hybridization between members of
separatebiparental spedes occurs multiple times to produce separateuniparental clones, the
componentorganismsof which are similar in most biologically significant respects. Cnemidopho
rus tesselatus
(reviewed by Wright 1993) is commonly dted as an example(e.g ., Kitcher 1984a,
Holsinger 1987, Wilson 1995). If interbreedingis the only processthat unites organismlineages
to form spedes, then neither the individual clonesnor the collection of them are spedes. However
, if processes other than interbreedingunite organism lineagesto form spedes, then it might
be argued either that the individual clonesare spedesor that the collection is a Spedes
, and the
collection is polyphyletic in terms of its component clones. If those clones represent separate
populations (e.g ., if they are allopatric), then the Spedesis polyphyletic in terms of its component
populations. This caseis similar to the caseof speciesparaphyly in that the issueis whether
to recognizea single spedesfor the entire set of populations (clones) as opposedto recognizing
eachindividual population (clone) as a spedes.
11. Even if the secondary properties always characterizethe same lineages, the alternative
spedesdefinitions basedon them might not be consideredrecondled in that the entities described
by those definitions are still conceptually, if not physically, distinct.
12. The situation is not quite as simple as stated in that not just any segmentof a population
"
lineagequali6esas a Spedes(see SpedesLife Cycles") .
13. It is often said that populations, not organisms, are the entities that evolve (e.g., Futuyma
1986), a view reflectedin the common definition of evolutionas changesin allele frequendesin
populations (e.g ., Wilson and Bossert1971, Hartl 1981). The evolution of populations, however,
is not the result of their organizationallevel, but rather of their temporal extent. Over short time
intervals (i.e., lessthan one generation), populations do not evolve any more than organismsdo.
Furthermore, organismlineages(as opposedto individual organisms) do evolve in the sensethat
they exhibit heritable change through descent. Thus, lineagesat all levels are the things that
evolve (Hull 1980), and a more accurategeneral definition of evolutionis heritable changesin
lineages.
14. This conclusion is analogousto the proposition that asexual(reproductively autonomous)
as analogouswith the term organism
organismsdo not form spedes. Considering the term species
that
the
situation
should
be
described
.
Becausewe talk about unicellular
differently
implies
organismsrather than saying that unicellular entities do not form organisms, it is more appropriate
to talk about uniorganismal species(provided that unisexual organisms do not form
population-level lineages) ~han to say that unisexual organismsdo not form spedes (see Hull
1980). An inddental benefit of this terminology is reconciliation (in theory, if not in practice)
of the proposition that asexualorganismsdo not form population-level lineageswith the taxonomic tradition that requiresall organismsto be members(parts) of spedes.
15. Spedespossessingdifferent contingent properties are useful for different kinds of studies.
Thus, just as one might examineonly sexually mature organismsin a study of mating behavior,
one might examine only reproductively isolated species(spedfically, those isolated by pre. mating barriers) in a study of reinforcement.
16. Severalauthors(e.g ., Chandlerand Gromko 1989, Mallett 1995) have arguedagainstspedes
definitions that treat putative speciationmechanismsor unifying processes as necessaryproperties
of spedes. They argue that suchdefinitions tend not only to restrict the generality of the
spedesconcept, but also to confusetheories about the origin and maintenanceof spedeswith
/
I. Monism, Pluralism
, Unity and Diversity
concept
for a generalspecies
are, in effect,arguments
itself. Thesearguments
theconceptof species
of the deAningpropertiesof
andthusarevery muchin keepingwith the reinterpretation
.
in thisessay
thespecies
categoryadvocated
of
do not differabsolutely
, asis revealedby theexistence
17. Eventheselevelsof organization
".
"
and
see
)
of
(
the
and possibility uniorganismal
unicellularorganisms
species Species Biology
18. Oneimportantreasonfor makingthis distinctionclearis that the wholemaybe morethan
thesumof its parts.
this resolutionof the individual
19. Ghiselin(1997; seealsoFrostandKluge1994) considered
" semantic
different
confuses
it supposedly
of species
andclass
trickery
" because
/ setinterpretations
in
resolution
the
the
.
contrary,
question
levelsin thehierarchyof biologicalorganizationOn
. If anyposition
requiresanexplicitdistinctionbetweendifferentlevelsof biologicalorganization
"
' s own positionthat the namesof taxa
assemantic
is to be characterized
trickery, it is Ghiselin
'
'
or 'Homosapiens
, not
... they aretermslike ' Mammalia
remainnamesof the taxathemselves
'"
' or ' human
'mammal
useof the
being (1997, 69). Thispositionbegsthe questionby assuming
their
of
the
sets
than
rather
wholes
level
the
organismal
to
termspecies designate population
is identical
, thereis nothingaboutthe idea
preference
parts. Althoughmy own terminological
to designate
areindividualsthat requiresusingthetermspecies
thatpopulationlineagesegments
.
their
of
the
sets
than
rather
themselves
organisms
thelineagesegments
component
and
not
do
, . producesingleoffspring
offspringthatdie youngor
,
20. Most organisms
, of course
.
still countasbranchingdespitetheearlyterminationof theirlineages
fail to reproduce
, then terminationby bifurcation
21. If speciesextinctionis analogousto organismaldeath
. At
. The followingterminologymakesthe relevantdistinctions
shouldnot be calledextinction
termed
is
which
birth
is
called
),
(
reproduction
the organismallevel
, the processof origination
. At thespecies
level
if it is highlyunequal
fissionif thedivisionis moreor lessequalandbudding
if thedivisionis moreor
, whichis termedbifurcation
of originationis calledspeciation
theprocess
is calleddefunction
. The terminationof organisms
if it is highly unequal
lessequalandblastation
with fission.
associated
when
called
it
is
;
disjunction
(death) whenthe lineageitself terminates
it
is calleddistinction
terminates
itself
the
;
when
ertinction
is
called
of
The termination species
lineage
with fission.
whenassociated
to includespecies
considered
conceptsthat do not conform
22. Thepluralistpositionis sometimes
- for example
in thisessay
, viewsin whichspecies
to thegenerallineageconceptdiscussed
of
basis
the
on
as setsdefinedsolely
organismalsimilarity(e.g., Kitcher
are conceptualized
such
to designate
the termspecies
use
of
.
volume
in
this
1
)
1993
, chapter
Although
1984a
; Dupre
one), it is doubtful
is
a
semantic
issue
the
on
(because
be
dismissed
cannot
grounds
logical
groups
"
in thisway (seeTheory and
thatanycontemporary
species
biologistsadual1yconceptualize
"
Operations).
- for example
, thoseregarding
differences
23. This positiondoesnot denycertainconceptual
do not, however
,
. Thosedifferences
esthat uniteorganismlineagesto form species
the process
es
abouttheprocess
. Instead
of species
, theyreflectdifferenthypotheses
reflectdifferentconcepts
) that form entitiesfitting the general
and thusthe kindsof organisms(e.g., sexualvs. asexual
.
lineageconcept
, nevermentioningthe possibilitythat
24. Darwin emphasized
divergencein this discussion
eventhe mostrecentlydivergedlinesof descentmight reunite.Thus, it is not clearwhetherhe
.
viewedthoselinesasbeingunifiedby somethingotherthantheirrecentcommonancestry
which
in
of
Anition
de
a
minimalist
species
species
Beatty(1985) arguedthat Darwinadopted
asspecies
by his fellownaturalistsandusedit to arguethat
weresimplythosetaxarecognized
'
, a theoryto explainthe
aslineageswas, therefore
evolved.Darwins conceptof species
. ' species
as
of the entitiesthat his fellow naturalistsrecognized speciesratherthana prescriptive
existence
.
definition
. 83
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Smith, A B. (1994). Systematics
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. London
J. Huxley, ed., Thenewsystematics
II
andLife' s Complications
Species
THEaLl ATEPOINTOFVIEW
Areas of Dissent
Beginning biologists are sometimestold they have to get inside the skins of
the organismsthey study, to think like a bird or a beetle. More experienced
biologists clearly have their perceptions shaped by the organisms they
study. Some biologists indeed come to look like fruit flies or mice. In this
essay, as a ciliatologist, I look at biological speciesfrom the perspedive of
the ciliated protozoa. A careful look at any particular organism might be
illuminating to philosophers of science, a useful corrective to airy speculations
and premature generalizations, but the ciliated protozoa have a considerable
"
"
"
"
history of providing not just a real perspective, but a really
different point of view ( Nanney1983).
In a thoughtful paper, Jan Sapp (1991) develops this thesis of the eccentricity
of the ciliates and ciliatologists by considering episodesof historical
turbulence generatedby work on ciliates. His paper includes the subject of
his earlier book (Sapp 1987), Beyondthe Gene: CytoplasmicInheritanceand the
, which describesthe studies on cytoplasmic
Strugglefor Authority in Genetics
inheritance in the ciliated protozoa that challengedthe information hegemony
of the nucleus.
The impact of the ciliate studies in heredity is illustrated by a comment
madeby John Maynard Smith (1983):
There are a few well-establishedexceptions[to nuclearcontrol] of which the
'
'
phenomenonof cortical inheritance in ciliates is perhaps the most important
. Neo-Darwinists should not be allowed to forget these cases
, because
the overwhelming
our
views.
However
threat
to
the
constitute
,
only significant
they
majority of inherited differencesare causedby differencesbetween
chromosomalgenes. (p. 39)
Sapp also discusses some whispers of heresy generatedby studies of mor
phogenesisin ciliated protozoa. This material has been superbly summarized
: Ciliate Studiesand
by JosephFrankel (1989) in his book PaHernFormation
Models. Ciliate developmentalstudiesraisevexing doubts about the adequacy
/
ll . Spedesand Ufe' s Complications
"
jar organisms) that appearedsuddenly in the geological record about halfway
through the history of life. The eukaryotessharean astonishing set of
complex adaptations. Unlike any prOkaryotes, they all possesssimilar electrically
excitable membranesthat facilitate communication and interaction
between cells and with their environment. All eUkaryotesexhibit homologous fiber systemsthat shapetheir bodies and move efficient motor organ
elles. All eukaryotes package their DNA on nucleosomic histone spools,
arrangedin conventional chromosomesthat undergo meiosis and mitosis at
appropriate times.
Ciliates differ &om higher eukaryotes, (as well as &om algae and fungi),
however, in the organization of their germplasm and soma. Like higher
organisms, they have compoundedtheir basic equipment to provide a larger
organismic entity than can be achieved within the domain of a single
genome, but they have done this without aggregating and differentiating
populations of discrete cells. They have pooled many identical genomes
&om dozens to hundreds within the bounds of a single amitotic macronucleus
, and they have pooled their cytoplasmic componentsinto a communal
cytoplasm with an elaborately differentiated cortex. They reserveone or
a few diploid micronuclei as germinal reservoirs that dance the mitotic and
meiotic pavannes, but that are incapableof ordinary transcriptional activities
until the next round of sexualactivity . The germinal information is periodically
summonedinto action in sexual episodes, after which a new somatic
genetic system is developedto replacethe old one.
How , or even if , these organizational featuresaccount for the discomfort
'
is uncertain. That
generatedby ciliatologists observations and conclusions
"
"
of
in
find
themselves
often
exceptional observations
possession
ciliatologists
"
"
and exceptionable interpretations is, however, undeniable.
THETETRAHYMENINFS
TheSimpleLife
My principal focus is a particular kind of ciliated protozoan, an organism (or
group, set, assembly, in.dividual, category see Ghiselin 1997) first dubbed
by Waldo Furgasonin 1940. They are small, deceptively simple
Tetrahymena
ciliated protozoa that have been seen and studied seriously only since the
perfection of the optical microscopein the last quarter of the nineteenth century
. Certain features of their organization were unknown, however, until
other technologiesemergedmore recently for the description of submicrosc.opic anatomy. These technologies particularly include the silver staining
cell surf
of
the
proceduresdevelopedfor visualizing the detailed architecture
.ace(Klein 1926, Chatton and Lwoff 1936) and the transmission electron
.microscopethat revealedthe complex patterns of underlying granules, fibers,
. Silver staining,
and membranesconstituting the architecturalsuperstructures
/
Nanney: When Is a Rose?
.
.U....".
M
18
-LwoEE
ciliaterevealedby O1atton
Figure4.1 Corticalfeaturesof a tetrahymenid
silverstains
.
Eachdot is a kinetosome
- - thebaseof a typical9 + 2 ciliumor flagellum
. Eachlongitudinalrow
of kinetosomes
is referredto asa ciliaryrowor kinety.Thetetrahymena
! oral apparatus
consists
of an undullatingmembrane
( UM
) andthreeothermembranelles
composingan adoralzoneof
membranelles
(AZM) ; theystainby virtueof theirpackedranksof kinetosomes
. Theciliaryrow
is designated
as kinety 1 (K1), or the stomatogenic
extendingbackfrom the oral apparatus
kinety, andthe new oral mlagen(OA) beginsto the left of this row asa proliferating6eldof
kinetosomes
. Theciliaryrowsarenumberedto the cell's right aroundthe circumference
of the
- paM) is designated
cell so that the row to the cell's left of row 1 (or the postoralmedian
as
kinetyn ( Kn), usuallya numberfrom 16to 20. Also associated
with kinety1 at theposteriorend
of thecellis the irregularlystainingoutletof thecytoproct(CYP
) . Approximatelyone-fourthof
the distancearoundthe cell to its right arethe openingsof the water-balanceorganelles
- the
contractilevacuolepores(CVP). (FromNanney1966)
for example, first allowed Furgasonto describethe four membranellescharacterizing the remarkably complex buccal apparatus(the tetrahymena! feeding
device).
. The tetrahymenines(Agure 4.1) are smaller than the more familiar
large
ciliates, such as Parameciumor Stentor. They are shapedlike a u .s. football,
for the samereason. They reach about 60 microns in length, but their size
varies during the vegetative cell cycle when they double in volume and
during starvation when they may be drastically reduced in size. The cell
/
D. Species
andUfe's Complications
/
Nanney: When Is a Rose?
SexComplicatesThings
'
At about the sametime, interest was rising in tetrahymena
s genetic system,
and crosseswere attempted. Theseattempts failed with the availablelaboratory
strains, which- it was belatedly discovered- were all amicronucleate
and asexual.Renewedisolationsfrom nature, however, quickly provided large
numbers of strains with micronuclei, and they were fully capableof undergoing
conjugation under appropriate circumstances(Elliott and Nanney
1952, Elliott and Hayes 1953). The major condition required for controlled
matings was the presenceof organismsof complementarymating types.
Earlier, Tracy Sonneborn(1937) had discovered mating types in another
ciliate, Paramecium
aurelia. The first mating strainsof tetrahymenahad a more
complex mating system: whereasonly two mating types constitute a mating
, the first tetrahymenasystem studied had seven
system in most paramecia
mating types ( Nanneyand Caughey 1953). Conjugation occurred when any
two mating types were brought together. Mating systemsof great diversity
abound in the ciliated protozoa, and the genetic economiesthey delineate
confound easy summary: Somegenetic specieshave only two mating types;
others have multiple mating-type systems that may facilitate mating with
"
"
strangers in an outbreeding economy. Other genetic specieshave characteristics
of inbreeders; they have short life cycles- terminated by autogamy
(self-fertilization) or death in a few weeks if a suitable mate isn' t found.
Others have no autogamy, but life cycles that last for years. The differences
among the speciesmay be interpreted as methods of coping with different
kinds of environmental challengesby meansof the differential utilization of
physiological plasticity and mutational variety. None of these differences
challengesthe Modem Synthesisor violates the esseritialcharacteristicsof
the biological speciesconcept, though they raisequestionsabout the units of
natural selection. The reader is directed to informed discussion of these
breeding systemsby Sonneborn (1957), Nanney (1980, chap. 10), Nyberg
(1988), and Oini and Nyberg (1993). The major studies in Tetrahymena
population geneticsare those of Ooerder et al. (1995).
Tetrahymenastudies soon revealedanother feature previously reported in
Paramecium(Sonneborn 1939, 1947); the morphologically indistinguishable
strains are actually divided into severalgenetically isolated systems(Gruchy
1955, Elliott 1973). Tetrahymena
pyriformis, like Parameciumaurelia, consists
of a set of noninterbreeding cryptic biological species. Becauseat first the
biological speciescould only be identified by the use of living tester strains,
'
they were referred to by numbers instead of by names, with Sonneborns
biological speciesidentifiable
. neologism syngenused to designate cryptic
'
only with live referencestrains. Sonneborns refusalto assignLatin binomials
to sibling speciesearnedhim the scorn of Ernst Mayr .
T. pyriformiswas clearly not a single biological speciesin the sensedefined
in the lexicon of the Modem Synthesis, but rather a collection of such
species. Yet T. pyriformiscontinued to be a useful collective term for awhile,
/
andLife's Complications
Species
100
/
ll . Species
andLife's Complications
Troplcalis
~
Group
I Pyriformis
I
G
I roup 81
7: /imacls
I Borealis I
) I
I Group@
Thermo~Hila
Group e
Lambomel
/a
Caudata
Group
Egg
I American 's I
I GrouP .
I
Glaucoma
102
ll . S~
103
Gene/PheneComplementarityin Tetrllhymenll
104
105
106
II. S~
clum, P. bursaria- harbor symbiotic algae, which are not essentialfor life,
but are clearly beneficialto the hosts.
With some other kinds of eukaryotic protists, the mutual dependenceof
diverse lineagesis more obvious- as, for example, in the caseof the fungus/
alga unions in lichens(Purvis 1997). Not so obvious are the multiple lineages
that commonly populate the cytoplasmic communities of sarcodinids. One
reasonthat we have so few of theseprotists under full domesticationis that
many of them are dependent on associateswe feel we have to get rid of.
Careful study has been given to some of the ciliate symbionts (Gortz
1988, Quackenbush 1988), mainly becauseof theoretical interest in the
"killer " character
, which was associatedwith a classicalcaseof cytoplasmic
inheritance.
Thanks to studies in molecular phylogenetics and to the achievementsof
"
of an
Lynn Margulis (1981) as an advocate, we now recognize the reality
'
ancient genetic conjunction at the base of the eUkaryotes that converted
bacteria into mitochondria and that may have converted a group of prokaryotes into the first eukaryote. Similarly, we acceptthe algal origin of plastids in green plants. We also have come to realize that the associationsare
guaranteed in part by the transfer of essential genetic elements from the
peripheral to the central genetic depot.
,
Perhapsbecausewe are fixated on closed gene pools, we have assumed
however, that the horizonal transfer of genetic elementsfrom separatephylogenetic systemsoccurred only rarely and in an earlier geological age. Jan
Sapp (1994) has discussedthese issuesat some length. The tight little gene
pools assumedin the Modem Synthesisprobably allowed much more leakage
in the protists via commensaltransfersand even in higher organismsand
in modem times more than is often assumed
.
The " cataclysmic" origin of new plant speciesby allotetraploid fusion of
separatedgenealogieswas acceptedearly in the Modem Synthesis as an
"
"
exceptional departure from the isolation of evolutionary lineages, but we
now know from DNA measurementsthat severalrounds of polyploidization
occurred within the ancestry of metazoanlineages. Did these chromosomal
doublings always involve closely related lineages, or did some of these significant
saltatory episodesinvolve hybridization of truly distinctive evolutionary
stems?
DISCUSSION
Historical Perspectives
This essaywas introduced with the observation that the ciliated protozoa
often fail to conform to the generalizationsthat apply to more familiar labora~ory denizens, and it has thus far summarizedsome of the major conclusions
concerning the evolutionary history and genetic economiesof the
tetrahymeQid ciliates. These organisms have been examined not because
107
Nanney: ~
Is a Rose?
108
109
:~
Nanney
Is a Rose
?
Prokaryotic Species
The situation with respectto prokaryotic speciesis far more serious. Microbiologists
have essentially bypassed the Modem Synthesis, considering it
irrelevant within their territories. Modem microbiology textbooks abound
"
"
with Latin binomials, but the species associatedwith the Linnaeanterms
makeno claimsof associationwith closedgenepools. The bacterialspeciesis
essentiallya set of organismswith an arbitrarily defined degreeof molecular
"
"
similarity and hence a group of presumed recent common origin (Goodfellow, Manno, and Chim 1997; d . Mishler, chapter 12 in this volume). The
mechanismsof genetic exchange and recombination in bacteria are not
Mendelian and no Mendelian population may exist ( MaynardSmith 1995) in
them. Conjugation- the mechanismmost similar to synkaryon formation in
eukaryotes- only rarely involves whole genomes. A large but uncertain
portion of genetic exchangesin natural populations is mediated by bacteriophages or plasmids, which insert genetic elementsand sometimesthemselves
(as in lysogenic phages) into the genomesof the organismsinvaded.
When released, the progeny may be inserted (via transduction) into another
host with another load of genetic baggage. The early studies of host range
(h) mutants demonstrated genetic control of the limits of transmission of
viral genetic elements, but little evidence has been reported that the gene
pools have any particular associationwith named species. Indeed, the rapid
spreadof antibiotic resistanceamong pathogens in clinical settings appears
to be a consequenceof low host specincity of resistance(R) plasmidsamong
even the most diverse microbes. What goes on in nature is of coursemuch
more obscure than what happens in the researchlab ( Embley and Stackebrandt 1997).
"
"
The concept of the selfishgene (Dawkins 1976) is right at home among
the prokaryotes, where individual genes appear capable of moving easily
acrosshighly diverse genomesunder appropriate selective conditions. The
modem genetic fluidity among the bacteria may be a relic of primordially
"
"
open genetic systems. The coadaptedgene complexes of the Modem Synthesis
and the isolating devicesrequired to maintain their integrity may have
evolved gradually though gradients of accessibility. In such an evolutionary
"
"
scenario, the species- i.e., the closed gene pool- might not have its
'i
"
origin until halfway through the history of life. As Doolittle (1998)
recently observed, the evidencecontinuesto accumulatethat eukaryotesare
110
constructed
committee
and
by
organisms
memories
of
several
within
their
carry
diverse
adventures
genomes
significant
genetic
Rediscovering
Time
Perhaps the theory of the Modem Synthesis, despite its acknowledged success
, has been relegated to a marginal position in modem biology because its
custodians failed to learn well the first Darwinian lesson. In their book Discovery
of Time ( 1965 ), Toulmin and Goodfield propose that the recognition
of the indefinite extensibility of time was the most fundamental feature of
the so-called Darwinian revolution . Like all intellectual discontinuities , the
scaling of temporal reality continues to have aftershocks. Neither the age of
the earth nor the time course of life has permeated fully the thinking of biologists
. We still focus on the last chapter of the book and ignore the long
stretch es back to a beginning nearly four billions of years ago .
The classification scheme we employ for living creatures, with its " higher
"
categories , was invented before the Darwinian realization of the depth of
time and was modeled on higher plants and animals, which are a fairly recent
phenomenon on the geological playing field (d . Ereshefsky, chapter 11 in
this volume ). Moreover , the playing field considered by the synthesizers was
refereed by umpires of Lyellian persuasion, and Cuvier was dismissed by
whiggish Darwinians . We now accept that life is long and well -punctuated ,
though we dispute the agencies of discontinuity . In any case, the number of
'
punctuation marks in an organism s evolutionary history probably reflects
the length of time lapsed since the origin of its " kind " ; the " natural " hierarchies
may be explained as consequences of evolutionary pulses. We can
hardly impose on all forms of life the same set of nested boxes by which we
rationalize the evolutionary history of organisms only a few million years
old - the species, genera, families , classes, phyla , and kingdoms of the metazoa. The number of nested boxes in the metazoan domain is insufficient to
order the ciliates , so protozoologists have rushed to elevate the ciliates to a
phylum . This move is interpreted , probably correctly , as a political move
instead of a scientific judgment , but clearly the ciliates need some new boxes
for the products of old genetic ferments .
A goal of molecular cbronometricians is to formulate appropriate standards
for dating the evolutionary branch es generated by cladistic analysis .
Though skirmish es over choices of molecular chronometers and modes of
analysis are fierce, a faithful cladist does not doubt the eventual resolution of
the details of evolutionary history . We shall soon date the evolutionary
junctions , and those junctions will define the problems to be resolved in a
new synthesis .
APPENDIX
A few words of explanation
are necessary
- about the methodolo Y
-. Lesused in
the molecularanalysisof tetrahymenastrainsbecause they are not very
111
: wt;;;;. Is a Rose
?
Nanney
/
II. Species
andLife's Complications
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118
II. Species
andLife's Complications
asEcologicalMosaics
Species
KimSterelny
SPEaFSAS A GRADE
The mechanismsof evolution have produced on Earth an astounding variety
of life forms. Together with adaptive design, the evolution of that diversity
is the central explanatory target of evolutionary biology . Though great,
however, the diversity of life on Earth is limited in important ways. Diversity
is bunchedor clumped. A walk in Australia' s eucalypt woodlands reveals
many parrots: eastern and crimson rosellas; red-rumped parrots; rainbow,
scaly, and musk lorikeets; galahs, gang-gangs, and an assortment of other
cockatoos. These birds are all readily recognizableas having some overall
similarity of form and behavior; their parrotness is apparent. But the divisions
within the group are apparent, too. There is no spectrumof birds from,
say, typical galahs to typical sulphur-crested white cockatoos. Equally, parrots
do not " fade into " the equally recognizableand distinctive pigeons. We
do not find a spectrum of birds from typical parrots through to typical
pigeons.
Life's mechanismshave produced phenomenological
: recognizable,
species
reidentifiable clusters of organisms. This fact makespossible the production
of bird and butterfly field guides, identification keys for invertebrates and
regional floras, and the like. However, the significanceof this clustering is far
from uncontroversial. The speciesconceptis one of the key conceptsof " folk
" - our set
of commonsense concepts for dealing with the living
biology
world (Atran 1990, and chapter9 in this volume; Berlin 1992). The extent to
which theoretical biology vindicates folk biology ' s categories is an open
question. We are large mammals, with a lifespan that is long by the standards
of animalecology, but not by the standardsof evolutionary change, and
we have a distinctive set of perceptualmechanisms
. We have no guarantee
that the distinctions that strike us as salient coincide with the natural kinds
of evolutionary biology . For example, one natural way to read Dawkins' s
book TheExtendedPhen~type(1982) is to seeit as arguing that folk biology ' s
C.<>
DCeptof an organism does not correspond to any natural kind of evolutionary
'
biology . Whether we accept Dawkins s caseor not, we cannot just
assiuneth.atour ordinary conceptof organismidentifiesa kind of evolutionary
, that phenomenological
biology . Equally, it has to be shown, not assumed
constitute
a
kind.
Minimalists
about
species
biological
speciesdo not expect
theoretical biology to vindicate the speciescategory: they think it is merely
a phenomenologicalkind. We can clump organisms into species
, but the
biology explaining the clumping pattern is so diverse that speciesdoes not
namea biological kind.
In this essay, I mount a limited defenceof the idea that speciesare objective
featuresof the world , existing independentlyof our perception and categorizatio
, and that they are not just epiphenomenalproducts of ecological
and evolutionary processes. I argue that some speciesare evolutionarily
linked metapopulations. Suchspeciesare ecological mosaics, for the separate
populations that composethem are scatteredthrough quite different ecological communities. When this occurs, the relationship between evolutionary
unit and ecological forces is complex and complex in a way that limits the
possibility of evolutionary change, so speciesthat are ecological mosaicsin
this senseare distinctive evolutionary units. Not all organismsare organized
into evolutionary species, so not all phenomenological speciesare evolutionary
species. But I argue that when this mosaicorganization does evolve,
it profoundly influencesthe future evolutionary possibilities of the species.
One traditional reason for skepticism about the signi6canceof phenomenological speciesis the idea that our capacity to identify them is an artifact
of our limited temporal and spatial perspective. Adelaide and crimson rosellas are now readily distinguishable, but if we trace thesepopulations back in
time, they begin to merge. Equally, if we could trace them forward in time
"
"
future Adelaide rosellas may look increasingly unlike their contemporaries
. If a smooth continuum of change links these birds to the earlier
parrots from which they evolved, then we might supposethat there can be
no fundamentaldifferencebetween the Adelaide rosella and its ancestor. Our
recognition of those species depends on our temporal standpoint. If we
were able to track the slowly changing lineage backward in time, we would
'
come to organisms different enough from today s rosellas to distinguish
them. But from a future vantage point , as the lineage continues to change,
'
today s rosellas would be the intermediate gradations between two other
parrot species Aqelaide rosellas as they have become and Adelaide
rosellas as they have just been. Hence, our speciesidenti6cations are not
the recognition of objective units in nature, for if lineageschangesmoothly
and seamlessly
, they canbe segmentedinto speciesin many ways. The choice
on
the
baselineand the amount of change that counts as enough
depends
for us to recognizea new species.Furthermore, evolutionary changeis often
seamless
. Though plant speciesare often formed rapidly through hybridization, jumps will be rare in animal lineages.
There is a secondreasonfor skepticismabout the signi6canceof phenomenological species. These clusters of morphologically similar organisms are
, and the clustersthemproduced by very different evolutionary mechanisms
120
/
ll. Species
andLife's ComplicatioN
121
/
asEcologica
J Mosaics
: Species
Sterelny
122
/
n. SpedesandUfe's Complications
SPECIES
ARE MOSAICS
In a famousmetaphor
, Dobzhansky(1937) picturedcloselyrelatedspeciesas
,
li J1ked
togetherthroughbeing on adjacentpeaksin an adaptivelandscape
with eachpeakrepresenting
a speciesniche. He wrote:
/
: Species
asEcological
Mosaics
Sterelny
124
/
ll. Species
andLife's Complications
125
126
127
Stere
Jny: ~
Mosaics
esasEcological
128
/
II. Species
andUfe's Complications
-129
Mosaics
Stere
Iny: s; eaesasEcological
If Eldredge, V rba, and those of similar views are right , two populations
consist of organismsof the samespeciesif they evolve (or more likely fail to
evolve) together. The extent to which populations are evolutionarily coupled
is not, of course, independentof the phenotypesof individual organisms, nor
is it any simple function of those phenotypes. Populations can becomepermanently
decoupledfrom one another through geological, environmental, or
ecological changesthat leave no obvious initial trace in individual phenotypes. An invasion can force one population to change its daily cycle; the
extinction of a long distancepollinator can disjoin two plant populations. Of
course, the idea that speciesidentity and speciationis just a matter of population
-level properties is controversial. Some, perhaps most, evolutionary
biologists take speciation to occur only when there have been intrinsic
changesin the two populations that make renewedgene flow impossible, so
their test for speciationis stronger than mere de facto permanentseparation
of the two populations. Yet on the faceof it , this criterion is puzzling, for the
view that speciesare historically defined entities is close to the consensus
view in evolutionary biology . As a consequenceof such a view, the facts
that make an organism a "member of a particular speciesare now considered
to be the relations of the organism to others, not its intrinsic physical
characteristics
.
Of course, even if we accepta thoroughly historical and relational account
of what makesan organism a memberof a given species, we still have good
epistemic reasonfor an interest in intrinsic charactersand changesin those
characters
. Often, we cannot tell of contemporary populations whether
extrinsic isolating mechanismsare permanent. The extinction of a critical
pollinator may be local, or a new pollinator may evolve and rejoin populations
that seemedpermanently separated. When the factors that have segmented
a previously coupled metapopulationinto disjoint parts are extrinsic,
we may be unable to tell whether we are dealing with a single speciesthat is
now a geographicaland ecologicalmosaicor a cluster of incipient species.In
this sense, as O 'Hara (1993) has noted, identifying specieshas a forward
looking elementto it.
However, I suspectthe insistencethat speciationrequiresintrinsic change
also stems from the idea that the speciesevolutionary biologists identify
should map closely onto phenomenologicalspecies. If Australian and New
Zealand boobook owls count as separatespeciessimply becausethe two
populations are permanently segmentedby 2,000 kIn of the Tasman Sea,
there would be no way a taxonomist identifying a specimenin a museum
could place that organism in its correct evolutionaryspecies
from the intrinsic
featuresof the specimenalone. I think this motivation is inapprophysical
'
priate becauseit stemsfrom a lingering adherenceto the idea that there is a
9
single and universal.speciescategory. I think, therefore, we should instead
see phenomenologicalspecies identiAable clusters of organisms- as fallible
cluesto the existenceof evolutionarily linked metapopulations. We have
already been forced to accept that the concordancebetween phenomeno-
.131
SterelnyrSpedesas EcologicalMosaics
132
ll . Spe~
133
UPSHOT
It ' s time to draw some general morals from this discussion. Dawkins
(1982) has argued that the developmental cycle is of great significancein
adaptive evolution. He argues that the evolution of adaptive complexity in
multicelled organisms depends on a developmental cycle from
single-cell
bottleneck to single-cell bottleneck. A genetic change can make
important
differencesto the whole organism when development is funneled
through
.
. this bottleneck. The change, by acting early in this process, can have global
. The developmentalcycle matters, for only at some points in
consequences
an organism's life history can a change make a global difference to the
134
ll. S~
andLife's ComplicatioN
organism (Dawkins 1982, chap. 14). The idea that specieshave ecological
structure, together with the turnover pulse hypothesis, suggeststhat species
too have developmentalcyclesthat are important in the sameway. Adaptive
~
phenotypic change13is possible only becauseof the metapopulation
to
take
the
with
place
change tending
population ~ metapopulation cycle,
at the most simplified point in that cycle, when the incipient speciesis a
both
single population. The change is entrenched only if that population
stock.
the
from
isolated
is
and
survives
parental
permanently
More generally, the importance in evolution of the geographic and ecohave
logical structure of speciesshows that speciesare real. We do not
to defend speciesselection to show that specieshave an ineliminable role
in evolution. In their 1993 review of punctuated equilibrium, Gould and
Eldredge identify standardmodem Darwinism as having three central commitments
: (1) the organism is the main unit of selection; (2) natural selection
"
"
novelty and hence is the creative
explains the existenceof evolutionary
"
"
force in evolution; and (3) genealogicalchange at all taxonomic levels is
nothing more than the accumulationof small change in local populations
"
'
his opponents as ultra(Gould 1995, 4). Eldredge, in particular, sees
"
"
14
"
Darwinians who acceptan extrapolationist view of evolution, and it is
this final clause that Eldredge takes himself to have refuted. Macroevolutionary
change is not just the accumulationof change in local populations,
. We
for such changes do not accumulateexcept in special circumstances
characteristics
cannot explain the evolution and entrenchmentof adaptive
without both explaining the role of Mayr' s Brake in buffering speciesfrom
'
s Brake
changein normal times and the brakes occasionalrelease. But Mayr'
and
existence
its
.
Hence
not
,
is a feature of metapopulations,
populations
a
within
evolution
from
operation cannot be understood by extrapolating
.
scales
single population to evolution within larger temporal and geographic
Natural selection within a population is the only mechanismthat generates
of standardDarwinism
adaptive phenotypic change: the first two commitments
in
however,
generating permanent
survive. That selection is only effective,
featuresof the
to
distinctive
for
least
at
(
adaptations
phenotypic change
local community) under relatively rare background conditions. The attempt,
then, is to defend a modest version of the view that evolutionary speciesare
kinds whose structure is of profound significancein evolutionary change.
; not all speciesare
Not all organismsare organizedinto evolutionary species
do evolve, they are
complex mosaics. But where specieswith this structure
not just epiphenomenona of phenotypic change. They profoundly influence
that changebecausethey are a central feature of the environment of change.
ACKNOWLEDGMENTS
Thanksto David Hull and Rob Wilson for their feedbackon an earlier
version of this paper .
.135
NOTFS
1. Bacteriado not fall easilyeveninto phenomenological
- at leastnot on the basisof
species
the featuresof theirmorphologyaccessible
. Theycomein a number
throughlight microscopes
of differentshapes
, but otherwisehavefew identifyinv
.
. characteristics
2. Thereis geneflow betweenspecies
because
hybridsarefertilewith bothparentalspecies
: the
in plantsof a mulebeingableto breedwith bothdonkeyandhorse.Buttherearealso
equivalent
formedby hybridizationin whichthe hybrid is fertilebut
species
reproductivelyisolatedfrom
both parentalspecies
. The sunflowerHelianthus
anomalus
seemsto be an example
; its parent
seemto beHelianthus
annuus
andHelianthus
species
-fertile
. Helianthus
anomalus
is cross
petiolaris
with neitherof theotherHelianthus
, but hasbeenexperimentally
species
recreated
fromthemby
hybridization(Niklas1997,64- 65)
3. Alternatively
, the generalityproblemis avoidedat thecostof problemsequallygreat. Some
versionsof the phylogenetic
species
concepttreatanypopulationwith a distinctiveandinvariant
featureasa species
. This treatmentdoesavoidrevivingall the problemsof the
biological
, but only at thecostof countinginnumerable
species
concept
.
ephemeral
populationsasspecies
Thespecies
sodefinedis not anevolutionarykind, in termsof eitherthemechanisms
category
that
producetheseclumpsor theirevolutionaryupshot.
4. As Hull (1978) pointsout, theseideasareinterdefinable
. Theissueis not whetherspecies
are
kindsor individuals
, but wheth~r theirdefiningfeaturesarehistoricalandrelationalor intrinsic.
5. I think this view is true accordingto both Elton's conceptionof a nicheasa functionalrole
within a communityand its Hutchinsonian
successor
in whichnichesareessentiallylinkedto
their occupants
(Elton 1927, Hutchinson1965). Damuth(1985) introducedthe termat1atarto
thefractionof a species
in a particularcommunity
designate
. Unlesswe definenichesvery thinly
- not spedes
- haveniches
indeed
, avatars
. Van Valen(1976), in his blendof an evolutionary
andecologicalconceptionof a species
, mayhavehada very thin definitionin mindindeveloping
hisnotionthat species
haveuniqueadaptivezones
. Still, themorework onewantstheniche
to do in explainingthe phenotypiccoherence
of a species
, the moredifficultit is to downplay
the ecologicaldifferences
betweenthedifferentpopulationsthatmakeup a species
. Theneedto
defineanadaptivezonethinly enoughso all organisms
in a species
sharean adaptivezonecuts
acrossboth the ideathat adaptivezonesexplainspecies
andthe ideathat they are
phenotypes
.
species
sped6c
6. I think we shouldbe very cautiousaboutacceptingthis idea
. Notice first that this idea
'
that species
are typicallyat equilibriumand that thoserangesareset by the
supposes
ranges
's
organism
powersof dispersal
. Moreover
interactingwith its limitsof tolerance
, peripheralisolates
cansometimes
outruntheir usualparasitesandpredators(Thompson1994, 159). It may
well bebest, therefore
, to thinkthatthemix of selectionin peripheral
rather
populations
changes
than to supposethat thesepopulationsare in suboptimalconditions
to organisms
compared
' distribution
closeto thegeographic
centerof the species
.
7. Eventhis beliefmaybe no morethandifferentideasabouthow to count
stability. Vrbaand
Eldredgearepalaeontologists
, andThompsonis not.
8. Fora very accessible
introductionto thisliterature
, seeWilson1992, 131- 141.
9. Biologicaltaxonomyhasnow evolvedinto systematics
, but the concernthat species
differences
be markedby features
identiAablein eachspecimen
alsobea lingeringvestigeof traditional
may
"
" in that
taxonomyandtheroleof the type specimen
.
taxonomy
"
. 10. Thus, Vrbacomplains
~ To saythat a parentspedesmustceaseto existonceit givesoff a
. branch
, andbe recognised
asa new spedesif it persistswithout changeafterbranchingis like
"
sayingI ceasedto existat the birth of my daughterandmustbe namedasa new individual
(Vrba 1995, 27).
136
ll . Sp~
'
"
"
11. In contrast to the isolation concept, Patersons synonym for the biological speciesconcept
. The point is that identifying isolating mechanismsimplicitly refers to another population,
the population from which one is isolated.
12. This point is bad news for someversionsof the phylogenetic speciesconcept.
13. Lessboldly , adaptive phenotypic changeof a certain kind- the kind driven by communityspecificecologicalfactors.
14. I am very skeptical of Eldredges identification of these ultra-Darwinians. He seems to
- the
believe that accepting gene selectionismcommits one to the minimalist view of species
view that they are a merely phenomenologicalkind. But theseare independentdebates:Williams
invented gene selectionism,but he defendsspeciesselection(1992).
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Brandon
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Cracraft
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"
Damuth
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Dawkins
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Eldredge
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Eldredge
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's
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Eldredge
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, S., ed. (1993). Evolution
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, S. (1985). Theontogeny
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.
Wilson
, E. O. (1992). Thediversityof life. New York: W. W. Norton.
138
III
Homeostasis
, andHigherTaxa
, Species
RichardBoyd
INTRODUCTION
Overview
idea that speciesare homeostatic property cluster kinds together with this
distinction to clarify other issuesabout the metaphysicsof species. In the
first place, I conclude that individual specieshave (homeostatic property
"
"
cluster) essences
, so that a form of essentialism is true for species
, albeit a
form of essentialismquite different from that anticipatedby Mayr and others
who have discussedessentialismin biology . Furthermore, I indicate how recognizing
speciesas homeostatic property cluster phenomenaand drawing
the distinction betweentypes of definitions allows us to makebetter senseof
issuesregarding " realism" and " pluralism" about species-level taxa.
I extend the application of the accommodationthesis to consideration of
the question of the reality of higher taxa. I argue that some higher taxa are
probably real natural kinds in the senseof the term required by the accommodation
thesis- indeed, probably homeostatic property cluster natural
kinds. I deploy that thesisto identify a crucial relation betweenjudgments of
arbitrariness or conventionality of representationalschemes
, and to show
how a referenceto that relation can help to clarify and to evaluate claims
about the conventionality of higher taxa.
Homeostatic Property Cluster Kinds
In the empiricist tradition since Locke, the standardconcepHon of scientific
"
"
(and everyday) kinds has been that they are defined by nominal essences
or by other purely convenHonal specificationsof membership conditions.
Part of that concepHon has been a concepHon of linguisHc precision, according
to which a properly defined kind will be defined by necessaryand sufficient
membershipcondiHons. Becausethe boundaries of kinds are, on the
nominalist concepHon characteristicof empiricism, purely matters of convenHon, any failure of scientific conceptsto correspond to this standard of
precision could, in principle, be remedled by the adoption of more precise
nominal definiHons.
The realist critique of Lockean nominalism that arose with naturalistic
conceptions of natural kinds and of the semantics of natural kind terms
(Kripke 1971, 1972; Putnam 1972, 1975a, 1975b) was articulated around
examplesof a posteriori definitions of natural kinds that likewise specified
necessaryand sufficient membershipconditions- such as natural definiHons
of chemical kinds by molecular formulas (e.g., " water = H2O" ). These critiques
thus gave support to what many authors call the " traditional" essentialist
concepHon of natural kinds, according to which, among other things,
suchkinds possessreal (as opposed to nominal) essencesthat define them in
1
. terms of necessaryand sufficientmembershipconditions.
At the time I began thinking about theseissues, philosophical conceptions
of kinds and categoriesthat did not treat definition by necessaryand sufficient
condiHons as the relevant standardof precision were pretty much limited
to Wittgensteinian and other " ordinary language" conceptions whose
extrapolation to scienHfic casesdid not seemto me very plausible.
142
/
RethinkingNaturalKinds
/
, and Higher Taxa
, Species
Boyd: Homeostasis
7. In suchcases
, the relative importanceof the various properties in F and of
the various mechanismsin determining whether the thing falls under I- if it
can be determinedat all- is an a posteriori theoretical issuerather than an a
priori conceptualissue.
8. Moreover, there will be many casesof extensionalindeterminacy, which
are not resolvableeven given all the relevant facts and all the true theories.
There will be things that display somebut not all of the properties in F (and/
or in which some but not all of the relevant homeostaticmechanismsoperate
) such that no rational considerationsdictate whether or not they are to
be classedunder t, assumingthat a dichotomouschoiceis to be made.
9. The causal importance of the homeostatic property cluster F, together
with the relevant underlying homeostaticmechanisms
, is such that the kind
or property denoted by t is a natural kind.
10. No refinementof usagethat replacest by a significantly less extensionally
vague term will preserve the naturalnessof the kind referred to. Any
such refinementwould require either that we treat as important distinctions
which are irrelevant to causalexplanation or to induction, or that we ignore
similarities which are important in just theseways.
11. The homeostaticproperty cluster that servesto define t is not individuated
extensionally. Instead, the property cluster is individuated like a (type
or token) historical object or process: certain changesover time (or in space)
in the property cluster or in the underlying homeostatic mechanismspreserve
the identity of the defining cluster. In consequence
, the properties that
determine the conditions for falling under t may vary over time (or space),
while t continues to have the samedefinition. The historicity of the individuation
conditions for the definitional property cluster reflects the explanatory
or inductive significance (for the relevant branches of theoretical or
practical inquiry ) of the historical development of the property cluster and
of the causalfactors that produce it , and considerationsof explanatory and
inductive significancedetermine the appropriate standardsof individuation
for the property cluster itself. The historicity of the individuation conditions
for the property cluster is thus essentialfor the naturalnessof the kind to
which trefers.
Examples
In almost any philosophical discussionabout the nature of natural kinds, the
author will illustrate her claims with especiallypersuasiveillustrative exam. pIes. It will , no doubt, seemodd to readerswho are biologists or philosophers
of biology that in my own paperson the subject, I deployed biological
speciesas such examplesof HPC natural kinds. It is a peculiarity of the literature
that in mainstreamanalytic philosophy, biological speciesare - along
with chemical elements and compounds- the paradigmatic natural kinds,
whereasamong philosophically inclined biologists and philosophersof biol -
144
/
HI. RethinkingNaturalKinds
. They differ &om natural kinds in that what unites their membersis their
historical relationshipsto one another ratherthan their sharedproperties.
I maintain that the first three of these considerationsdraw their current
plausibility &om a profoundly outdated positivist conception of kinds and
that the fourth participates in both this sameerror and in a misestimateof
the explanatory role of speciesconceptsin biology . I offer an alternative to
,
the positivistically motivated conception of natural kinds and their essences
and explain why , in the light of this alternative, biological speciesproperly
count as natural kinds, defined by real essences
, even if in some sensethey
are also like paradigm casesof individuals.
I then indicate how the insights of the alternative accountcan be extended
to provide resourcesfor the treatment of other aspectsof the speciesproblem
, and even to certain issuesabout higher taxa.
The Essenceof Essentialism: Toward a New Understanding
One implication of the HPC conception of (some) natural kinds is that the
reflectedin the four considerationsand
positivist conceptionof natural kinds
"
"
suggestedby examplessuch as water = H2O misleadsus about what is
145
/
, Species
, and Higher Taxa
Boyd: Homeostasis
146
/
DI. RethinkingNaturalKinds
(1969) that the theory of natural kinds is about how schemesof classiAcation
contribute to the formulation and identmcation of projectible hypotheses(in
the senseof Goodman 1973). The naturalnessof natural kinds consists in
'
their aptnessfor induction and explanation; that s why (on one scienti Acally
central notion of deAnition) deAnitions of natural kinds are reflections of the
properties of their membersthat contribute to that aptness.
thesis
The thesisI defend here (the accommodation
) makesthe further claim
that what is at issue in establishingthe reliability of inductive and explanatory
practices, and what the representationof phenomenain terms of natural
kinds makespossible, is the accommodationof inferential practicesto relevant
causalstructures.
Here is the basic idea. Consider a simpliAed casein which reliable inductive
practices depend on our having a suitable vocabulary of natural kind
terms. Supposethat you have been conducting experiments in which you
, the flame
exposedvarious salts of sodium to flames. In ea~ ~of many cases
turned yellow . You conclude that always (or almost always) if a salt of
sodium is heated in a flame, then a yellow flame results. You are right , and
'
.
your inferenceis scienti Acally respectable
is
a reflection of the fact that the categories
in
this
matter
success
Your inductive
salt of sodium, flame, and yellow are natural categoriesin chemistry,
and of the fact that the hypothesis you formulated with the aid of reference
to thesecategoriesis a projectableone.
Now , anyone who has read Goodman (1973) can come up with indefinitely
many unprojectablegeneralizationsabout suchmatters that At all past
data equally well, but that are profoundly false. You were able to discernthe
true one becauseyour inductive practicesallowed you to identify ageneralization appropriately related to the causal structures of the phenomenain
question. In this particular case, what distinguished the generalization you
acceptedfrom the unprojectable generalizations(which also At the extant
data) was that for any instantiation of it that makesthe antecedenttrue, the
state of affairs describedby the antecedentwill (in the relevant environment)
causethe effect describedby the consequent. Your deployment of projectable
categoriesand generalizationsallowed you to identify a causallysustained
generalization.
What is true in this simpliAed example is true in general of our ability in
scienti Ac (and everyday) practice to identify true (or approximately true)
generalizations: we can identify such generalizationsjust to the extent that
we can identify generalizationsthat are (and will be) sustainedby relevant
causalstructures. Things may be hairier than they are in our example; perhaes the truth makers for the antecedentsof true instantiations are symptomatic
.
effectsof causesof the statesof affairs describedby the consequents
Perhapsthe generalizationsspeak of causalpowers and propensities rather
th' anof determinateeffects so that it is the causalsustenanceof propensities
rather than the causation of effects that is relevant. Perhapsthe generalizations have a more complex logical form. And so forth.
/
, Species
, and Higher Taxa
Boyd: Homeostasis
De6nition
148
/
III. RethinkingNaturalKinds
kind is provided
categories. In one senseof the term, a definitionof a natural
use of a natural
the
that
role
by specifying a certain inductive or explanatory
demandsof a
kind term referring to it plays in satisfying the accommodation
kind a programmaticdefinition
disciplinary matrix. Call this sort of definition of a
inductive
the
an
element
/ explanatory role indicated by
. Defining
by
an
its location in the periodic table would be exampleof offering aprogram
matic definition for it .
There is another perfectly legitimate senseof definitionaccording to which
a definition of a natural kind is provided by an account of the properties
sharedby its members- in virtue of which, referenceto the kind plays the
role required by its true programmaticdefinitions. Call this sort of definition
an explanatorydefinition. Defining a chemicalelement in terms of its atomic
number and the associatedvalence structures is an example of offering an
explanatory definition.
'
To a good first approximation (I m ignoring here the issues of partial
, subtle questions about the individuation
denotation, nonreferring expressions
of disciplinary matrices, translation of natural kind terms between different
etc.) one can
languagesemployed within the samedisciplinary matrix,
of
satisfaction
the
of
terms
in
characterize true explanatory definitions
accommodationdemandsas follows:
Let M be a disciplinary matrix and let t1, . . . , tn be the natural kind terms
es
success
deployed within the discoursecentral to the inductive/ explanatory
definitions
of M . Then the families Fl' . . . ' Fn of properties provide explanatory
of the kinds referredto by t1, . . . , tn just in case:
. Epistemicaccesscondition. There is a systematic, causally sustained tendency
- established by the causal relations between practices in M and
causalstructuresin the world - for what is predicatedof ti within the practice
of M to be approximately true of things that satisfy Fi, i = 1, . . . , n.
. Accommodationcondition. This fact, together with the causal powers of
use of 4 , . . . , tn in M
things satisfying Fl ' . . . , Fn, causallyexplains how the
inferential
the
of
contributes to accommodation
practicesof M to relevant
for
causalstructures: that is to the tendency
participants in M to identify
.
causallysustainedgeneralizationsand to obtain correct explanations
To put the matter slightly differently, one can say that the explanatory
definition of a natural kind is provided by an account of the family of properties
shared by its members which underwrite the inductive/ explanatory
roles indicatedby its true programmaticdefinitions.
A (Sort of ) Continuum of Definitions The best-known treatments of
in the philosophical literature
programmatic and explanatory definitions
the definition of psychological
probably lie in functionalist discussionsof
of such states proposed
definitions
abstract
and
states. The very general
at
efforts
are
programmatic definitions:
by so-called analytic functionalists
characterized
they define psychological states in terms of very broadly
149
functionalistaccounts represent
explanatory roles. By contrast, so-called psycho
eHorts at explanatory de6nitions of the same states. (Excellent discussions
of theseconceptionsare to be found in Block [1980].)
There are, however, many ways in which the literature on functionalism
raisesissues- about the analytic-synthetic distinction and about the properties
of mental states in physically impossible organisms, for example - that
are irrelevant for our presentpurposes(for a discussionof some of them see
Boyd forthcoming a). For that reason, it is probably better to take as paradigm
casesof programmatic de6nitions the de6nitions of chemicalelements
in terms of the inductive/ explanatory roles indicatedby their positions in the
periodic table and to take their de6nitions in terms of atomic number as paradigm
casesof explanatory de6nitions.
What theseexamplesillustrate - and what is true in general- is that both
programmatic and explanatory de6nitions of a natural kind embody claims
about the causalpowers of its members. In fact, although there is an important
diHerencebetween the aims of the two sorts of de6nitions, there is
something like a continuum between the most abstractly formulated programmati
de6nitions of a natural kind and its explanatory de6nitions. Thus,
for example, a chemicalelementmight be programmatically defined in terms
of the causal/explanatory role corresponding to a particular place in the periodic
table, but the causal/explanatory role it occupiesmight equally well be
spelledout in term of valenceor in terms of the structure of orbitals, and so
on, with ever-increasingspecificationof the details of its causal/explanatory
role in chemistry until the characterizationsin terms of causal/explanatory
role converge to an account of an explanatory de6nition of the element in
question.
Thus, the relationship between proposalsfor programmaticde6nitions, on
the one hand, and proposals for explanatory de6nitions, on the other, is
quite complex. As the literature on analytic functionalism and psychofunctiona
suggests, even when proposed programmatic and explanatory
definitions for a natural kind are quite diHerent, there needbe no incompatibility
"
"
between them. Once the continuum just discussedis recognized, we
can seethat the samecan be true of two quite diHerent programmatic definitions
of the same kind, provided that they are cast at diHerent levels of
abstraction. At the sametime, becauseprogrammaticde6nitions are a posteriori claims about the relation between the causalpotentials of things and
the accommodationdemands of disciplinary matrices, unobvious conflicts
between programmatic and explanatory de6nitions of the same kind, or
between programmatic de6nitions of a kind involving diHerent levels of
abstraction, are possible.
What will prove important for our purposesin considering de6nitions of
individual speciesis the simple point that programmatic formulations of
speciesde6nitions in terms of explanatory roles are not, in general, rivals to
explanatory de6nitions in terms of common factors, relations of descent,
gene-exchange, and so on.
ISO
/
III. RethinkingNaturalKinds
IS1
Boyd: Ho~
stasiS
, andHigherTaxa
, Species
1lI. ~ hinkingNaturalKinds
153
, Species
, andHigherTaxa
Boyd: Homrostasis
Relationally defined categories, such as social roles, are natural kinds just
in casedeployment of referencesto them contributes to the satisfaction of
the accommodationdemandsof the disciplinary matrices in question. Their
explanatory definitions include relational properties just in casethe shared
causal powers and dispositions among their members- upon which that
contribution to accommodationdepends- are causallysustainedby (among
other things) shared relational properties. That an imaginary and unpracticable
disciplinary matrix might embody the project of, for example, predicting
and explaining the behaviors of social animals by deriving them &om
independently formulated intrinsic physical characterizationsof the animals
and of their environments is irrelevant to the question of whether (partly )
extrinsically defined socialkinds are natural kinds in the disciplinary matrices
in which we actually work.
It may be somewhat more difficult to see why the definitions
Historicity
of natural kinds need not be ahistorical and unchanging. Consider first the
question of whether the explanatory definition of a natural kind can be such
that membersof the kind are necessarilyrestricted to some spatial or temporal
region, or such that it involves referenceto a particular space-time region
or individual.
The obvious casesof natural kinds with just these properties are the historical
periods recognizedby an explanatorily relevant periodization of the
history of some phenomenaor other. Supposefor the sakeof argument that
important causalfactors in Europeanhistory are revealed if we distinguish,
for any given political and economicregion, between a feudal period, on the
one hand, and the period of transition to recognizably modem organization
of trade, production, and governance. If this is so, then the distinction in
question will correspondfor each region to two different natural categories
of historical events and processes, such that the consequencesof a historical
event will tend to be significantly determinedby its situation with respectto
this periodization. Of course, the natural historical periods in question would
"
"
have vague boundaries- they would possesshomeostaticproperty cluster
explanatory definitions but as we have seen, this vaguenesswould not
undermine their status as natural kinds in the sense appropriate to the
accommodationthesis.
If an example in which the members of the kinds are historical events
seemstoo atypical to be fully convincing, consider the (homeostatic property
cluster) distinction between feudal and capitalist economicsystems. It is
almost certainly true that recognizing this distinction contributes fundamentally
to accommodationin the disciplinary matrix that includeseconomicand
social history .
Now , according.to some economic theories (Marxist ones, for example),
"
"
this distinction correspondsto quite general (inexact) laws of economic
development such that in any suitably situated human society there would
154
, andHigherTaxa
Boyd: Horn~ tasis, Species
156
1lI. ~ hinkingNaturalKinds
area perfectlyordinaryphenomenon
in disciplinarymatricesconcerned
with
thehistoryof complexphenomena
.
Homeostasis , Compositional Semantics, and Disciplinary Matrices
The accommodationthesis has one more consequenceof that we need to
examine before we turn to issues about biological species. Disciplinary
matricesare themselvesHPC phenomena.What establishes the coherenceof
an intellectual discipline is a certain commonality of methods, explanatory
strategies, relevant Andings, and the like. We may seehow this sort of commonality
results in disciplinary coherenceby recognizing that, within any
disciplinary matrix, very, very many accommodationdemandsarisefrom the
enormous range of quite particular phenomenafor which explanationsand/
or predictions are sought. What we recognize as an intellectual discipline is
the phenomenon manifested when a cohesive set of laws, generalizations,
, technical and inductive methods, and explanatory
conceptual resources
strategiescontributes to the satisfactionof a very wide spectrumof accommodation
demands.
The conditions of satisfactionof these accommodationdemandsare thus
themselves homeostatically related: the satisfaction of various demands
tends systematicallyto contribute to the satisfactionof many other demands.
In typical disciplines, this homeostasisis in large measurea matter of widely
applicable causalknowledge: the commonalitiesamong or systematicity in
the significant causalinteractions between the factors that produce the phenomena
under study are such that the knowledge of such factors necessary
to solve one disciplinary problem will conduce to the solution of a great
many other problems.
'.
This homeostatictendency is reflectedin the very phenomenonof natural
kinds. What we recognizeas a natural kind is a multipurpose category, reference
to which facilitates the satisfaction of a great many accommodation
demandswithin a disciplinary matrix. Here, then, is a particular aspectof the
homeostasisjust mentioned: typically , the kind distinctions central to meeting
one of the accommodationdemandsof a disciplinary matrix will facilitate
the satisfactionof many of its other accommodationdemands.
What is important for our purpose is the way in which this particular
aspectof disciplinary homeostasisis related to the compositional semantics
of natural kind terms. We are usedto the idea that natural kinds are the kinds
that are the subjectsof natural laws- not perhapseternal, ahistorical, exceptiorness laws, but at least explanatorily significant causalgeneralizationsof
some sort. It is important to note that even this concessionto the positivist
tradition overstatesthe connectionbetweennatural kinds and laws. The naturalness
of many natural kinds is indicated not by their being the subjectsof
natUrallaws, but by the fact that referenceto them is crucial for the formulation
of laws with more specificsubject matters. Goodman's (1973) contrast
between greenand grue illustrates this point . There are no interesting laws
/
, Spedes, and Higher Taxa
Boyd: Homeostasis
about green things generally, but referencesto colors like green are important
in formulating explanatorily important psychologicalgeneralizations.
More scientifically important examplesof the samephenomenonare provided
, ion, and compoundin
by, for example, the categories acid, element
.
Few
chemistry
explanatorily important generalizationsapply to all of the
membersof any of these categories, but referenceto them is central to the
formulation of important laws. The contribution that recognition of these
categoriesmakesto the satisfactionof accommodationdemandsin chemistry
, ion, and compound
dependson the compositional roles of the terms acid, element
in specifying the subjectmatters of important generalizations.
Even when a natural kind exhibits its naturalnessby being the subject
matter of explanatorily important causal generalizations, the homeostatic
contribution that its recognition makesto the satisfactionof accommodation
demandsin the relevant disciplinary matrix will typically depend to a great
extent on the compositional role of natural kind terms referring to it . The
paradigmaticnatural kinds (speciesexcepted) chemicalelements provide
a spectacularillustration of this point . There are, to be sure, laws regarding
each of the elements. Nevertheless, the overwhelming majority of chemical
natural kinds are compounds rather than elements, so the overwhelming
majority of chemicallaws do not have elementsas their subjectmatter. Thus,
the main contribution that the use of terms referring to elementsmakesto
the satisfactionof accommodationdemandsin chemistry arisesfrom the use
of suchterms in formulas for chemicalcompounds.
Two related points follow that are important for the later discussionof the
metaphysicsand epistemology of the speciescategory. In the first place, the
naturalnessof a natural kind is not a matter of its being somehowfundamental
, with less fundamentalkinds being somehow less natural than more fundamenta
ones. Thus, for example, with the discovery of the phenomenon
of chemicalisotopes, there was no methodologically or philosophically significant
"
II
problem about the true or real elementallevel in chemistry, with
conflicting positions regarding the question of whether the true or morefundamenta
elementallevel consistedof categoriesdefinedjust by atomic number
or of categories defined by atomic number and atomic weight. The
decision to adopt the practice of using the term elementfor categoriesof the
. first sort was a matter of convenience, not a matter of fundamental metaphysicso
fundamental chemistry. What was important- and not just a
matter of convenienceor convention- was that either choicewould result in
the establishmentof a vocabulary for chemistry in which the sameclassof
causallyand explanatorily relevant distinctions could be drawn. The naturalness
of a natural kind is a matter of the contribution that referenceto it
makes to the satisfaction of the accommodationdemandsof a disciplinary
matrix, in the context of a system of a compositional linguistic resourcesfor
the representationof phenomena.
158
/
1lI. RethinkingNaturalKinds
159
/
, Species
, andHigherTaxa
Boyd: Homeostasis
"
"
kind may be natural from the point of view of some discipline or disciplinary
"
matrix, but not from the point of view of " another. Perhapsjade is a
natural kind in gemology or the history of art, but not in geology ( because
somejade is jadite, and some is nephrite, and these two minerals are chemically
quite different). This relativity to a discipline or disciplinary matrix
does not compromisethe naturalnessor the " reality" of a natural kind. Natural
kinds simply are kinds defined by the ways of satisfying the accommodation
demandsof particular disciplinary matrices.
1993
(
) makesa similar point about the relativity of the naturalness
Dupre
of kinds to particular projects. He arguesfor a " promiscuousrealism" about
natural kinds accordingto which, among other things:
There is no God-given, unique way to classify the innumerableand diverse
products of the evolutionary process. There are many plausible and defensible
ways of doing so, and the best way of doing so will depend on both the
purposesof the classificationand the peculiaritiesof the organismsinquestion
, whether those purposesbelong to what is traditionally consideredpart
of scienceor part of ordinary life. (p. 57)
The accommodationthesis- according to which the naturalnessand the
"
"
reality of a natural kind consist in the contribution that referenceto it
makesto the satisfactionof the accommodationdemandsof a particulardisciplinary
matrix- supports and provides a metaphysical rationale for this
'
aspect of Dupre s conception ( but probably not to his other critiques of
unificatonist conceptionsof science- critiques I confessto not fully understanding
). Different disciplinary matrices and different accommodation
demandswithin a disciplinary matrix will - given the complexity of the biological
world - require referenceto different and cross-classifying kinds in
order to achieveaccommodation, and this fact in no way demeansthe naturalness
"
"
or the reality of those kinds.
'
One of the criticisms of Dupre s conception offered by Wilson (1996) is
that the classificatorycategoriesof ordinary life and languageare not natural
"
kinds at all; he deniesthat common senseand common languageare in the
"
businessof individuating natural kinds at all (p. 307). According to Wilson,
ordinary language lacks the systematic purpose of uncovering order in
nature, which governs scientific practice and language, and which makes it
necessaryfor scientific terms (as opposedto ordinary languageones) to refer
. Dupre (1993) himself indicatesthat
to natural kinds definedby real essences
the plurality of natural kind classificationsin ordinary languageis unsurprising
becausecommon sense aims to gather information about the world ,
rather than primarily to achieve a unified picture of it . Wilson agrees, but
identifies the latter aim with the sciencesand seesreferenceto natural kinds,
definedby real essences
, as appropriateonly to the latter task.
"
I
allows one to " split the difference between these
advocate
The position
two conceptionsof everyday kinds. Although my choosing the term disciplinary
matrix undoubtedly betrays my special concern with the issue of
kinds in the theoretical sciences
, everyday life provides disciplinesor at any
160
/
III. RethinkingNaturalKinds
161
/
, Species
, and Higher Taxa
Boyd: Homeostasis
I have in mind, of course, the casesin which referenceto what are plainly
scientific kinds (of diseasesand medicines; of semiconductorsand other electronic
parts; of reagents for photographic development, etc.) plays a role
in everyday practical or recreational endeavors. But I also have in mind
a general feature of ordinary linguistic usage that seemsto point toward a
general recognition of the everyday relevanceof theory-driven standardsof
classification.
Dupre (1981) launched the casefor (what becameknown as) promiscuous
realismby insisting that in the ordinary everyday sense of the term lily ,
'
'
onions (among other plants) arent lilies. Although it is true that we don t
' decorative our
,
judgments
ordinarily count onions as lilies becausethey arent
are
lilies
are
remarkabout
whether
onions
ones
even
our
)
(
ordinary
"
ably sensitive to the ways the question is put . Someonewho says, Onions
"
are lilies, may seem to have spoken falsely or misleadingly, but someone
"
"
who says, Onions are a kind of lily , says something that many would
"
intuitively accept. There are lots of similar cases( Birds are a kind of dinosaur
" "
"
" ; The glass snake is a kind of lizard ; Tomatoes are a kind of
"
" '
fruit ; Mushrooms are not really a kind of plant ) in which the expression
"kind of "
signals referenceto (or, if you prefer, deferenceto ) scientific and
theoretical standards. The fact that ordinary language has such a semantic
device for marking out and thus making available reference to scientific
standardsprovides, I believe, further reason for recognizing that ordinary
kinds and scientificnatural kinds lie along a continuum. They do so precisely
becausethey are all kinds of natural kinds- that is, resourcesfor achieving
accommodation.
Natural Individuals
When we presently turn our attention to the famous (or infamous ) question
of whether biological species are kinds or individuals (see also de Queiroz ,
chapter 3 in this volume ), we need to recognize that it is a consequence of
the accommodation thesis that the question may not have as deep a metaphysical
import as the literature would suggest . Once we begin to think of
natural kinds as fea~ es of human inferential architectures - as artifacts rather
than as Platonistic entities - as the accommodation thesis requires, the distinction
between natural kinds and natural individuals becomes less important .
A number of philosophers have suggested something like this conclusion in
discussing the species-as-individuals issue. ~ pre ( 1993 ) concludes that the real
"
question about whether species are individuals or kinds is whether the same
set of individuals can provide both the extension of a kind and the constituent
"
parts of a larger individual . And the answer to" this is clearly yes ( p . 58 ).
"
Ereshefsky ( 1991 ) understands the traditional notion of a natural kind
"
approximately along the lines indicated in the earlier section " entitled strategy
"
; he therefore concludes that species are not kinds , but historical entities
'
. ~ Still , he does maintain that some of them are individuals as well ,
162
III. RethinkingNaturalKinds
whereas others are not, so he does not take the category individual to be
incompatible with the much more kindlike category historicalentity, which
includesthe higher taxa.
'
Finally, Wilson (1996) seemsto hold that Dupre s conception, if developed
'
in Dupre s promiscuousor pluralist style, would commit one to " the
absurdity of saying that one and the same thing is a natural kind and an
individual" (p. 310). But even he then goes on to say that the choice
between the two conceptionsof speciesis " merely pragmatic," suggesting, I
believe, that neither has an advantage in satisfying the accommodation
demands of biology . What I propose is that by seeing the similarities
between the inductive and explanatory roles played by referenceto natural
kinds, on the one hand, and by referenceto individuals, on the other, we can
seewhy the distinction between natural kinds and (natural) individuals is, in
an important way, merely pragmatic.
After all, successfulinduction and explanationdependjust as much on the
accommodation of our individuative practices for individuals to relevant
causalstructures as on the accommodationof those practices for kinds. A
failure to be able to recognize the various stages in the maturation of an
organism as stagesof the sameorganismwould undermine induction and
explanation in biology just as much as a failure to deploy accommodated
schemesof classificationfor the organismsthemselves.The fact that it is, for
certain familiar cases
, easierto get this sort of thing right should not prevent
our recognition that the classificationof temporal stagesas temporal stages
of the sameindividual must meet just the sameconstraints of accommodation
as the classificationof individuals into natural kinds. Nor should this
fact lead us to miss the point that sometimesaccommodationof inferential
practices for individuals is a real scientific achievement, as in the case of
organismswhose larval and adult stagesare so dissimilar as to appearcontraspecific
. If the truth be known, the spatial or temporal stagesof a natural
individual form something like a natural kind.
It may seemodd to think of the stagesof someordinary object- that rock
over there, for example - as forming a natural kind; .after all, particular rocks
aren't typically explanatorily important enough to make the honorific title
natural kind seemappropriate. This is lessclearly so for somebigger rocksthe rock of Gibraltar, for example- or for other sorts of individuals- Oliver
Cromwell, let' s say. In thesecasesand many others, the accommodationthat
underwrites cogent explanations(I assumethat historical explanationscount
ascausaland require accommodation) dependon our capacitiesto .individuate
explanatorily important individual entities. Of course, if biological species
are i!\dividuals, then they are individuals with the explanatory importance
characteristicof natural kinds.
Eyen with respect to the casesof inconsequential( but still natural) individuals
, our capacitiesto individuate are central to successfulaccommodation
of inferential practicesto causalstructures. Thus, for example, experimental
trials on ordinary (and individually explanatorily unimportant) mice, trees,
163
/
, Species
, andHigherTaxa
Boyd: Homeostasis
mineral specimens, DNA ~amples, fossils , rivers , and so on depend for their
'
inductive cogency on experimenters abilities to properly individuate these
things . Experimental studies on gruified mineral samples would represent
failures of accommodation in just the same way and to just the same extent
that such studies of ( properly individuated ) grue samples would .
Just think about a Quinean hydrologist studying river -kindred water
stages. The distinction between natural kinds and natural individuals is
almost just one of syntax . In particular , the metaphysics of accommodation is
the same for natural kinds and for natural individuals .
A Humean Note
I have just argued against a conception of natural kinds according to which
they must be defined by unchanging necessary and sufficient membership
conditions and must figure in eternal , ahistorical , exceptionless laws. I suggested
that the current plausibility of this conception arises not from any
important features of actual scientific practice , but from the' demands
(together with a bit of physics envy ) of the logical empiricists project of
providing Humean rational reconstructions of causal notions .
Now , I have argued elsewhere ( Boyd 1985b ) that such Humean reconstructions
'
must always fail . ( Here s the argument in brief . Scientific realism is
true , so we have [unreconstructed ] knowledge of factors such as the charge
of electrons . But charge just is a causal power , so knowledge of unreconstructed
causal powers is actual.) What is important for our purposes is that a
rejection of the Humean project of rational reconstruction is not necessary in
order to accept the conclusions of the preceding sections of this essay.
'
"
'
Perhaps there is some metaphysically innocent notion of iaw or of iaw
"
likeness in terms of which an antimetaphysical reconstruction of causal
notions can be provided . Whether this notion exists or not , scientific (and
historical and everyday ) knowledge often depends on our being able to
identify causally sustained generalizations that are neither eternal nor ahistorical
nor exceptionless , and our ability to do so depends on our coordination
of language and classificatory categories with causal phenomena
involving and defined by imperfect property homeostasis. Any adequate
Humean rational reconstruction , whether of science or of other areas of empirical
knowledge , will need to be compatible with the recognition of these
facts and will thus be compatible with (a suitably reconstructed version of )
the homeostatic property cluster conception of natural kinds advanced here.
. SPECIES AS HOMEOSTATIC
Species as Homeostatic
KINDS
Phenomena
164
/
1lI. Rethinking Natural Kinds
/
, Species
, andHigherTaxa
Boyd: Homeostasis
166
/
III. RethinkingNaturalKinds
areatleastverymuchlikenatural
: theyreflect
kinds
solutions
to the
Species
accommodation
demands
. Moreover
in
ofbiology
the
which
reference
, ways
to themcontributes
to satisfying
these
demands
makes
themresemble
paradigmatic
natural
kindsasopposed
to theleastkindlikenaturalindividuals
kinds
(whicharethemselves
).
verymuchlikenatural
I propose
thatbiological
are
natural
kinds
. Whatis
HPC
species
simply
is thatthebestarguments
in favorof thealternative
view- that
interesting
ratherthankinds actually
I amproposing
thethesis
theyareindividuals
support
. Whentheresidual
of
kinds
is
,
away
positivist
conception
stripped
whatthebestarguments
thatspecies
areindividuals
ratherthankindscome
in thesame
downto, atleastto agoodfirstapproximation
, isthatorganisms
must(a) bemembers
of someinitialpopulation
of that
biological
species
its
that
a
temor
descendants
of
members
so
cannot
become
(
species
species
, bemembers
) and(b) if contemporaneous
porarilyextinctandthenreevolve
eitherof thesame
or
of
that
are
relevantly
population populations
reproductively
of species
haveimportant
(so thatthe constituents
integrated
internal
witheach
other
ofparadigm
individuals
do).
relations
, asconstituents
Themorecogentreasons
for insisting
thatspecies
musthavethetwo
characteristics
onoutdated
of science
.oneddonotdepend
justmenti
philosophy
. Whenafamilyof populations
of organisms
satisfies
, butonbiology
(a)
is a
and(b), thefactof theircommon
descent
andreproductive
integration
serious
source
of a tendency
towardevolutionary
unity. Thebiologically
claimthatwithouttheoperation
for (a) and(b) restonthescientific
arguments
the
a
of
will notpossess
of thefactors
, family populations
theyrequire
. (Considerations
of this
ev.olutionary
of species
leveltaxa
unitycharacteristic
sortareexplicitin, forexample
, Hull[1978
] andin Ghiselin
[1974
].)
'
in favorof
of argument
, forthesake
, thattheconsiderations
. Letssuppose
a
and
b
are
correct
.
Then
common
descent
and
()
()
integration
reproductive
evolutionary
of thesorttheyrequire
areessential
to establish
thehomeostatic
inferences
and
:
the
by
unityof biological
unity
anticipated
species
167
/
, Species
, andHigherTaxa
Boyd: Homeostasis
168
/
III. Rethinking Natural Kinds
more than indicate why this is a better or more natural way of formulating
taxonomic claims.
In the first place, I have offered a general theory of the nature of natural
kinds (the accommodationthesis) that affords a rebuttal to the more philosophical (and positivist ) argumentsagainst the thesis that speciesare natural
kinds. It does more than that however. The category natural kind is itself a
natural kind in metaphysicsand epistemology, and the accommodationthesis
is a thesisabout its essentialor explanatory definition. It follows from this
definition that biological speciesare natural kinds and not marginal examples
either. Their homeostaticproperty cluster structure is perfectly ordinary for
natural kinds; they are deeply important to the satisfactionof the accommodation
demands of a very, very successfuldisciplinary matrix; and their
'
departuresfrom the positivists conception of natural kinds are all essential
to the accommodationthat referenceto them helps to achieve.
In fact, just as philosophers have usually thought, biological speciesare
paradigmatic natural kinds. The natural kinds that have unchanging definitions
in terms of intrinsic ~ecessaryand sufficient conditions and that are
the subjectsof eternal, ahistorical, exceptionlesslaws are an unrepresentative
minority of natural kinds (perhaps even a minority of zero). Every sort of
practical or theoretical endeavor that engageswith the world makesaccommodation
demandson the conceptualand classificatoryresourcesit deploys.
Recognition of the sorts of kinds beloved by positivists can meet the
demandsfor very few (perhapsnone) of these endeavors. Instead, the sort
of kinds (many of them homeostaticproperty cluster kinds) required for the
inexact, messy, and parochial sciencesare the norm. Of thesekinds, biological
speciesare entirely typical, indeed paradigmatic, examples.
SPECIFS
AMONGTHETAXA
Pluralistic
Realism
169
/
, Species
, andHigherTaxa
Boyd: Homeostasis
170
/
flI. RethinkingNaturalKinds
i71
/
, Species
, and Higher Taxa
Boyd: Homeostasis
considering propose that there is no such nature - that, instead, there are
many different, (approximately) equally methodologically important ways of
, each corresponding to a different legitimate way of
demarcating species
understandingspecies-level taxa. If the solution is so easy, why does it represent
a fairly recent proposal?
One reason, no doubt, has been the admirably motivated but (in the light
of the complexity of homeostatic mechanisms
) ultimately fruitless effort to
establish something like a universally applicable" operational deAnition" of
conspecificity (or at least a unitary formula that determinesthe relevant deAnition
for any group of organisms) and thereby to establishconsistencyand
uniformity of classiAcatory and nomenclaturalpractice. Arguably, the articulation
of the species-as-individuals conception contributed to the plausibility
of this project. If speciesare thought of as unique among the taxa in being
evolutionary individuals in nature rather than human constructs (as many
believe), then perhapsit is more plausible that a single unitary conception of
conspeciAcity deAned in terms of the relevant notion of individuality
will be forthcoming.
What I suspect, however, is that the main sourceof the speciesproblem is
practical. Many disciplinesare like biology in that there are schemesof classiAcationthat- by themselves- are almost adequatefor the satisfactionof a
wide variety of different accommodationdemands- for example, the classiAcation of the elementsin chemistry and the standardclassiAcation of (what
are called) mineral speciesin geology. In eachof these disciplines, the compositional
"
"
characterof natural kind terms is exploited to Ane-tune these
almost adequatecategoriesto At more particular accommodationdemands.
Thus, we speak
, for example, of the isotopes of chemicalelements, the different
physical forms of elementalsulfur, and the different varieties of quartz
in order to achievemore nearly complete accommodation. There is no persisting
"
"
" elements
"
problem in chemistry and there is no speciesproblem in
geology precisely becauseby using suitablenatural adjectival terms to modify
'
other natural terms, we can achieve accommodation, and it s merely a
matter of conveniencejust how we do this. This is just the point I made
earlier- that the compositional semanticsof natural kind terms is important
to the ways in whicb the accommodationdemandsof disciplinary matrices
get satisAed.
'
Why cant we do this in biological taxonomy as well? The answer, I suggest
, is that the compositional semanticstructure of the standard Linnaean
system of taxonomic nomenclatureis inadequatelyflexible. Thus, for example
, one might hope to take advantage of the tight homeostasisbetween
. the factors sustaining homeostasiswithin each particular speciesby settling
(it might not matter exactly how) on some one reasonableway of deAning
the species-level taxa and then satisfying the accommodationdemandsof
explanatory programs not perfectly served by this classificationby deploying
additional natural adjectival terms to differentiate further between
groups of organisms or populations. (Wilson [1996 and chapter 7 in this
172
/
III. Rethinking Natural Kinds
/
, Species
, and Higher Taxa
Boyd: Homeostasis
/
III. RethinkingNaturalKinds
over and above a natural kind term together with its use in satisfying accom'
moda Hon demands. ( What elser ' you ask. Well , there ' s whatever is necessary
to accommodate transla Hons that preserve saHsfacHon of accommoda Hon
demands and to accommodate phenomena such as reference failure and par Hal denota Hon.) Or , better yet , the establishment of a natural kind (remember
that natural kinds are legisla Hve achievements - that is, arHfacts) consists
solely in the deployment of a natural kind term (or of a family of such terms
connected by prac Hces of transla Hon) in saHSfymg the accommoda Hon
demands of a disciplinary matrix . Given that the task of the philosophical
theory of natural kinds is to explain how classificatory prac Hces contribute
'
to reliable inferences, that s all the establishment of a natural kind could consist
in : natural kinds are the workmanship of women and men.
The causal structures in the world to which accommoda Hon is required
are, of course, independent of our practices (except when our practices are
[part of ] the subject matter ; see Boyd 1989 , 1990, 1991 , and 1992 for better
formula Hons). SHll, natural kinds are social artifacts . That ' s why asking
whether a kind exists independently of our prac Hce is the wrong way to
inquire about its reality . No natural kinds exist independently of prac Hce.
The kind natural kind is itself a natural kind in the theory of our inferen Hal
'
practice . That s why the reality of kinds needs to be understood in terms of
the saHsfacHon of the accommoda Hon demands of the relevant disciplinary
matrix .
Natural Individuals , Again The very same points can be made about
natural individuals, such as organisms. The relations of causal continuity ,
similarity, or whatever that unite the temporal stagesof an organism exist
independently of our practices, and they have the causaleffects that make
referenceto that organism important to the satisfaction of accommodation
demandsindependently of our practice. But the grouping of those temporal
stagesunder a common linguistic or conceptualheading- treating them as
constituting an organism- is just as much a matter of social practice in service
of accommodationas the establishmentof a natural kind.
'
It s tempting to argue that this view can't be right becauseeven if we
become extinct, dogs might continue to exist, so they must be organisms
that exist independently of us. Of course, dogs might continue to exist: the
persistenceconditions (properly) associatedwith the notion of an individual
dog might continue to be satisfied, but the fact that these persistenceconditions
are natural ones- the fact that persisting dogs are individuals " in
"
nature, as one might say- is a fact not about nature alone, but about how
bilogical practicesare accommodatedto nature. After all, some organisms
would be in Mammalia even if we becameextinct, and they would continue
t~ occupy placesin the -relevant continuing historical lineages: in that sense
Mammalia too exists independentlyof us.
Nature makes temporal stages similar and different, continuous and discontinuous
, but things are the workmanship of women and men.
/
, Species
, and Higher Taxa
Boyd: Homeostasis
Realism
Higher Taxa and Accommodation Neither for kinds nor even for individuals
is the question of their reality best understood as a question about
'
"
"
"
independencefrom our practices. That s why questionsof reality or realism
"
about them are best understood as questionsabout the accommodation
of disciplinary matrices to causal structures. Thus, no simple contrast between
speciesand higher taxa with respectto their independenceof practice
can establishthe unreality (or diminished reality) of higher taxa. They may
yet be unreal (or less real), but this unreality is not a matter of their being
the results of human conception and practice. If they are unreal, it will be a
matter of their failure to contribute effectively to accommodation.
Assessing Accommodation : Methodological Spectra and the Equifertility
Principle I want to make a proposal about how we might fruit fully approach issuesconcerning the contribution that referenceto higher
'
taxa (or to any other kinds) makes to accommodation. Let s say that the
choice between two alternative classificatory schemeswithin the context
of a disciplinary matrix is arbitrary, just in case neither scheme reflects
-relevant causalstructuresbetter than the other. When sucha
aceommodation
f
choice is arbitrary, the disciplinary matrix would (from the point of view of
accommodation) be equally well servedby either scheme.
Now , one measureof the extent to which a classificatory schemecontributes
"
"
to accommodation- one measureof its reality - is given by the
range of alternative schemeswith respect fo which a choice would be arbitrary
. Philosophersor biologists who differ about the reality of higher taxa
will differ about which choicesbetween higher taxonomic schemesare arbitrary
ones. How are we to assesscompeting claimsabout sucharbitrariness?
It will help to answer this question if we consider the methodological
reflectedin a disciplinary
import of such claims. By the substantiveconception
matrix at a particular time, let us understandthe theories, doctrines, putative
insights, and so on regarding the relevant subject matters acceptedat that
time. Of course, in any actual case, there will be issuesand controversiesof
varying degreesof importance within a disciplinary matrix, so referring to
the theories and so on that are acceptedat a particular time involves some
degreeof idealization, but nothing in what I argue here dependson any subtleties
about how the idealization is understood. I do intend that substantive
conceptionsbe thought of as conceptualentities: as representationsof phenomena
deploying the conceptualresourcesof the matrix rather than, for
. example, as sets of propositions understood as nonconceptua1entities. The
substantiveconception CM of a disciplinary matrix M is thus the representation
within M of the causalknowledge putatively achievedin M .
The inferential practiceswithin a disciplinary matrix M will be (except in
caseswhere practitioners reasonbadly) justified by the substantive conception
'
CM. That s how the accommodationof inferential practices to causal
176
/
1lI. RethinkingNaturalKinds
177
/
, Spedes
, andHigherTaxa
Boyd: Homeostasis
These points are obvious, but important. They allow us to identify ways
of specifying and assessingconventionality estimatesregarding disciplinary
matrices. One way of specifying an estimate of conventionality EM for a
matrix M with substantivecontent CM is to specify a range of alternatives
to CM such that the choice between CM and any of these alternatives is
to be understood as arbitrary or conventional in the sensethat disciplinary
matricesjust like M , except that they deployed anyone of these other representatio
, would equally well reflect facts about the relevant subject
matters ).
The exampleswe have just consideredillustrate a quite general and fundamental
methodological principle concerning conventionality and its relation
to methodology- a principle that indicates another (related) way in
which conventionality estimatescan be specified(and sometimesassessed
).
to
the
when
the
choice
between
two substantive
According
equifertility principle,
contents is arbitrary or conventional, the two substantive contents
are methodologically
equifertilein the sensethat no methodological principle
or practice is justified by one unless it is also justified by the other. The
equifertility principle is about as obvious a methodological principle as there
can be. It follows via a pretty straightforward application of the accommodation
thesis- provided that one rejects the neo-Kantian view, apparently
advocatedby Kuhn (1970), that the adoption of a paradigm or conceptual
framework can noncausallydetermine the causal structures of the relevant
phenomena(seeBoyd 1990, 1992).
What is especially important for the present discussionare the implications
of the metaphysicalinnocencethesis in casesin which it is proposed
that the prevailing conventionality estimateEM for a matrix M is too modest
and that there are alternativesto CM with respectto which the choice of
CM is unexpectedlyconventional. Sucha proposal entails that any inference
or inferential practice that would be justified ( by the standardspreviously
prevailing in the matrix) given CM, but not given anyone of the alternative
, is thereby shown to be itself unjustified. No inferencesthat
representations
on
conventional
or arbitrary choicesof representationalschemesare
depend
Justified.
By the methodological
spectrumof a disciplinary matrix M at a given time,
let us understandthe inferential strategiesand methodological practicesjustified by CM. What we have just seen is that any proposal of unexpected
conventionality within a disciplinary matrix entails that the methodological
spectrumof the matrix is narrower, in a systematicallyspecifiableway, than
. Thus, we have two ways of specifying
practice within the matrix assumes
the
of
a
claim
of
import
unexpectedconventionality. One way characterizes
'
the conventionality in terms of the representationswith respectto which the
.
. choice of prevailing substantive content is said to be arbitrary or conven
. tional; the other way indicatesthe dimensionsof the narrowing of the methodological spectrumof the disciplinary matrix thereby required in the light
of the equifertility thesis.
178
DI. Re~
NaturalKinds
179
180
III. Ret~
ng NaturalKinds
-181
Boyd: H ~
eostasis,
Taxa
, andHigher
Species
important way. There are very good reasonsto believe that at least some
genera are real in this way. I have already indicated why pluralist realism
about speciessuggeststhat somegenus-level categoriesare real. If, as
many
authors have suggested, there are casesin which homeostasisat
approximately
the specieslevel obtains in families of populations between which
gene exchangeis minimal or nonexistent (in the caseof asexuallyreproducing
"
," for example), we have reasonsto believe
reptilian or amphibian species
that the samesort of homeostasismight obtain in at least some
recognized
genera, perhapsin most.
Moreover, if some higher taxa are real kinds that are important in evolutionary
theorizing, it is difficult (although, no doubt, not impossible) to
see what their importance could be except as (representationsof ) stasis'
'
producing factors. If that s what real higher taxa are, then it s equally difficult
, given the complexity of evolutionarily relevant causalfactors, to see
how the contribution to stasis in any particular case could fail to involve
homeostasisof severaldifferent factors. I propose, therefore, that insofar as
some higher taxa are real and important categoriesin evolutionary
theory
(above and beyond their "important role in representingpatterns of ancestry
and descent), they are probably, like species, homeostatic
property cluster
kinds.
If there are higher taxa that are real in this way, it is important to note that
there is no particular reasonto believe that their homeostatic
property cluster
definitions will honor strict monophyly, which is not to deny that the
homeostasislinking the members of such a taxon might always crucially
involve facts about the effects of their common ancestry. Thus, even if a
requirementof strict monophyly is appropriatefor someother higher taxa, it
need not be so for taxa in question.
Modest Cladism Suppose, for the sake of argument, that some higher
taxa- some genera for example- are real homeostatic property cluster
kinds in the way indicated. What are we to make of the concern that efforts
to discern evolutionary patterns in the fossil record- the causesof which
define higher taxa- will identify patterns for which no explanation in terms
of evolutionary tendenciesexists?
The obvious answer is that this problem may arise for some higher taxa
and not others. Perhapstaxa at the genus level- as taxa at that level are
generally erected- are usually real in the special sensediscussedhere, but
order-level taxa are usually not. Perhapssome such pattern obtains, but it is
different acrossphyla, given extant practices. Perhapstaxa of shorter historical
duration are more likely to reflect genuine stasis-sustaining properties.
Perhapstaxa erectedto accountfor the earlier stagesin the history of life are
more or less likely to be real than those taxa erected to account for later
.
. stages. Perhaps, in this regard, things are really a messfor which there is no
simple characterization.
182
1lI. Ret~
ng NaturalKinds
In any event , barring the extremely unlikely possibility that the standard
criticisms of evolutionary systematics are somehow without force in light of
the slight modification to this position we are considering, there will be
some domain of higher taxa about which the cladistically inclined system atist can reasonably maintain that the only important fads about the evolution
of life----which we can reflect in ereding such taxa - are historical fads
about relations of ancestry and descent. About erection of taxa of this sort ,
the only nonconventional or nonpragmatic constraint would then be one of
monophyly . This modest version of cladism is the one in which I am inclined
to believe .
ACKNOWLEDGMENTS
In formulating my approachto natural kinds, I have bene6tedgreatly from
conversationswith Eric Hiddleston, BarbaraKoslowski, Ruth Millikan , Satya
, SusannaSiegel, Jason Stanley, Zoltan Szabo,
Mohanty , Satya Shoemaker
and JessicaWilson. My thinking about biological taxonomy bene6ted
greatly from conversations with Christopher Boyd, Kristin Guyot , and
Quentin Wheeler.
NOTES
of naturalkindspart of whathe calls
1. Wilson(1996) goesso far asto makesuchconception
"
"traditionalscient
of scienti Acrealismwascentered
tradition
that
the
to
me
. It seems
i Acrealism
"
-veriAcationist
on the issueof refutingempiricist
argumentsagainstknowledgeof unobserv
"
ables ratherthanon the issueof whetheror not scientifickindsareindividuatedby essences
. Earlyon, the traditionalrealistturn
conditions
andsufficientmembership
that specifynecessary
andto realismaboutmental
of
behaviorism
a
rise
to
in the philosophyof science
critique
gave
in this critique
who participated
realists
scientific
hold
that
to
.
It
is
and
states
.
properties implausible
believedor werecommittedto believingthat thenaturalkindsof psychologyalwayshave
.
conditions
andsufficientmembership
detennined
by necessary
sharpboundaries
. 183
Boyd: ~
eostasis,Species
, and Higher Taxa
It is likewiseimplausible
that traditionalessentialist
viewsalwaysincorporated
sucha conception
of kind definitions
. Thosebiologistswho haveheldthathumanraces
, astheyareordinarily
shouldnot be understoodto haveheldthe additional
recognized
, havediHerentbiologicalessences
absurdpositionthatsuchracesalwayshavesuchsharpboundaries
.
1.. I thankProfessor
DavidHull for suggesting
thisclariAcation
.
3. One metaphysical
commitment
madeby Linnaeus
himselfthat Ereshefsky
criticizesis that
taxaat the levelsof genusandspecies
aredefinedby mind-independent
essences
whereastaxa
abovetheselevelsaresubjectto only pragmaticconstraints
. Ereshefsky
deniesthedistinctionon
the groundsthat thereareno taxon-speci6cessences
at any level. If the conception
of essences
defended
hereis correct
andprobablymanytaxaabovethe species
, thenspecies
leveldo have
essences
(albeitnot of the sort Ereshefsky
hasin mind), but all biologicaltaxaare, in a certain
senseof theterm
, minddependent
, or at leastpracticedependent
.
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CFS
BlockN. (1980). Readings
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, N. (1973). Factfictionandforecast
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'
/
, andHigherTaxa
, Speaes
Boyd: Homeostasis
REALISM
NATURALKINDSAND SCIENTIFIC
The idea that there are natural kinds hasa history in and an aptnessfor articulating
realist views of science. Realistshave traditionally held something like
the following view of natural kinds: natural kinds are what the sciencesstrive
to identify ; they feature in laws of nature and so scientific explanation; they
are individuated by essences
, which may be constituted by unobservable(or
" theoretical"
) properties; and they are conceiver-independent classifications
"
"
of what there is in the world - they carve nature at its joints.
The traditional realist view of natural kinds extends the following naive,
commonsense view. There are objects and properties that exist independently
of human observers. For example, supposethat we have before us a
of
piece rock. It has properties, such as a certain massand constitution, and
the rock and its properties exist independently of human observers. Scientists
investigate such objects, uncover certain relationships between their
properties, and develop taxonomies natural kinds that make these relationships
more apparent. Supposeour rock has the property of being made
of molten lava (composed, say, of 50% silica) and so has a certain melting
point and various other chemicalproperties. By taxonomizing it as an igneous
rock, scientistscan both recognizeits relationship to other kinds of rock and
explore the relationshipsbetween the properties that igneous rocks have.
The traditional realist view of natural kinds goes beyond suchacommonsenseview, chiefly in the depth of its metaphysicalcommitments. Distinctive
'
is the realist s view of why certain relationshipsbetween properties hold and
why scientific taxonomies that identify natural kinds reveal further relationships
between properties. Some properties are coinstantiated or correlate
with one another becausethey feature in laws of nature, and these laws hold
becauseof how nature is structured. In addition, the properties that featurein
laws of nature are intrinsic properties of the entities that have them: they are
properties that would .be instantiated in those entities even if those entities
.were the only things that existed in the world . Natural kinds, then, categorize
objects in terms of the intrinsic properties they have: same intrinsic
that
properties, samekind of thing . This in turn explains why taxonomies
identify natural kinds lead to further revelations about how properties are
related to one another, assumingthat the most fundamental
properties in the
world are intrinsic properties. In moving from traditional realism in
general
to critiques of it within the philosophy of biology - within the literature on
"
the speciesproblem" in particular- I want to focus on two further
aspects
of this overall metaphysical conception of natural kinds, essentialismand
.
unificationism
Essentialismis the view that natural kinds are individuated by essences
,
where the essenceof a given natural kind is a set of intrinsic (
perhapsunobservable
) properties, each necessaryand together sufficient for an entity ' s
a
being member of that kind. Realiststhus say that scientific taxonomy proceeds
by discovering the essencesof the kinds of things that exist in the
world and that this explains, in part, the theoreticaland practical success
es of
science. The endorsementof essentialismprovides a way of
distinguishing
natural kinds from arbitrary and conventional groupings of objects. Natural
kinds are kinds(rather than mere arbitrary collections) becausethe entities so
grouped sharea set of intrinsic properties- an essence- and natural (rather
than conventional or nominal) becausethat essenceexists
independent of
human cognition and purpose.
The rejection of essentialismabout speciesand, along with it , of the idea
that speciesare natural kinds at all has been central to the claim that
species
are individuals (Ghiselin 1974, 1997; Hull 1976, 1978).
According to this
view, the traditional realist misconstruesthe ontological nature of
species:
speciesare individuals rather than kinds individuated by essences
. Essentialism
about specieshas also been attackedindependently in the
philosophy of
1970
and
Sober
1980
.
(
)
by
biology
Mayr
(
)
As a general thesis, unificationismis the view that scientific
knowledge
is unifiedin some way; for the traditional realist, it is the view that because
natural kinds reflect preexisting order in the world , they are unified or integrated
. But realists are not alone in holding some version of uni Acationism
about scientific knowledge. The strongest versions of unificationism were
held by the logical positivists as the " unity of science" thesis (e.g.,
OppenHelm and Putnam 1958) and came with a reductive view of the nature of
"
"
higher -level scientific categories. More recent unificationist views have
been nonreductive- cast in terms of the notions of constitution or realization
, rather than in terms of identity . Traditional realism, whether in its
reductionist or nonreductionist guise, implies views about the basisof membership
in a given natural kind, the relationship between the various natural
kinds and the complexities in nature, and the way in which natural kinds
. themselvesare ordered. We might expresstheseviews as follows:
. the commonality
: there is a common, single set of sharedproperties
assumption
that form the basisfor membershipin any natural kind
. the priority assumption
: the various natural kinds reflect the complexities
one ~ ds in nature rather than our epistemicproclivities
188
/
Rethinking NaturalKinds
. the orderingassumption
: natural kinds are ordered so as to constitute a
unity
For a traditional realist about species
, the commonality assumptionamounts
to essentialismabout natural kinds; the priority assumption points to the
world rather than to ourselvesas the source of the variety of natural kinds
one finds; and the ordering assumption, typically expressedin the view that
natural kinds are hierarchicallyorganized, saysthat there is oneway in which
different natural kinds are related to one another.
Pluralists about speciesreject either the priority assumptionor the ordering
assumptionor both. For example, Mishler and Donoghue (1982) reject
the ordering assumption, but maintain the priority assumption when they
"
to adequatelycapture
say that a variety of speciesconceptsare necessary
"
the complexity of variation patterns in nature (p. 131). Dupre (1981, 1993),
by contrast, would seem to reject both the priority and ordering assumptions
"
in suggestingthat the best way of [classifying species] will dependon
both the purpose of the classificationand the peculiarities of the organisms
"
in question (1993, 57; d . D.upre, chapter 1 in this volume). Kitcher seemsto
"
sharethis view when he saysthat there is no unique relation which is privileged
in that the speciestaxa it generateswill answer to the needs of all
"
1
biologists and will be applicableto all groups of organisms (1984, 317).
Traditional realism about species is indefensible, and in the next two
sectionsI indicate just how this view has motivated the individuality thesis
(the second section) and pluralism (the third section). But reflection on the
similarities between the case of speciesand the case of neural taxo~omy
leavesme skepticalabout the plausibility of the inferencesto thesetwo views
about species (the fourth section). Moreover, I argue that the resources
'
afforded by Richard Boyd s (1988, 1991, chapter 6 in this volume) homeostatic property cluster view of natural kinds provide a view of speciesthat
lies between traditional realism, on the one hand, and the individuality thesis
and pluralism, on the other (the fifth section). I suggest that rather than
,
rejecting the connection within traditional realism between realism, essence
us
leaves
a
that
in
and kind, we need to complicate those relationships
way
closer to traditional realism than we might have expected.
TAXA
INDMDUALITYAND SPECIFS
A natural way to apply traditional realism to specieswould be to hold that
membersof particular speciesshare a set of morphological properties or a
set of genetic properties, eachnecessaryand together sufficient formembership
in that species. Let me take the morphological and genetic versions of
this view separately. For example, according to the former of these views,
domestic dogs, members of Canisfamiliaris, share some set of observable
'
properties presumably determinate forms of phenotypes such as having
four legs, hair, a tail, two eyes, upper and lower teeth- eachnecessaryand
189
/
Wilson: Realism
, andKind
, Essence
190
treats species as kinds rather than individuals , but they are nominal kinds
rather than natural kinds because the measure of overall morphological similarity
is a function of the conventional weightings we assign to particular
morphological traits or DNA segments.
By contrast , proponents of the individuality thesis respond to the failure
of essentialism with respect to species taxa by claiming that species are not
natural kinds at all , but individuals or particulars- with individual organisms
being not members of the species kind , but parts of species because a species
itself is an individual . Species have internal coherence, discrete boundaries ,
- all
properties that particulars
spatiotemporal unity , and historical continuity
have. Viewing species
kinds
nor
nominal
natural
neither
which
have , but
as individuals rather than as kinds allows us to understand how species
can have a beginning (through speciation ) and an end (through extinction );
how organisms can change their properties individually or collectively and
still belong to the same species; and why essentialism goes fundamentally
wrong in its conception of the relationship between individual organism and
species.
CATEGORY
AND THESPECIFS
PLURALISM
The individuality thesis is a view of the nature of particular speciestaxafor example, of Canisfamiliaris. BecauseI suggestedthat the individuality
thesis was a competitor to both traditional realism and pheneticism, I also
think of the latter two views as making claims about particular speciestaxa.
But pheneticismis also often taken as a view about the speciescategory
that is, as a view about what definesor demarcatesspeciesas a concept that
applies to a unit of biological organization. So construed, pheneticismis the
view that speciesare individuated by a measureof overall phenetic similarity
, with organisms having a certain level of overall phenetic similarity
counting as species, and higher-level and lower level taxa having, respec
tively , lower and higher levels of similarity .
Apart &om pheneticism, the various proposalsthat have been made about
what characterizesthe speciescategory are often divided into two groups:
(1) reproductiveviews, which emphasizereproductive isolation or interbreeding
as criteria- including Mayr' s (e.g., 1982) so-called biological species
'
of it , such as Patersons (1985) recognition concept
concept and relaxations
'
views,
and Templeton s (1989) cohesionspeciesconcepti and (2) genealogical
which give phylogenetic criteria the central role in individuating speciesand
are typified by Cracraft (1983) and Wiley (1978). Unlike pheneticism, both of
thesefamilies of views fit naturally with the individuality thesisas a view of
speciestaxa.
. The focus of both reproductive and genealogicalviews, as views of the
speciescategory, is on two questions: (a) what distinguishes species&om
other groupings of organisms, including varietiesbelow and generaabove, as
well as more clearly arbitrary groupings? and (b) how are particular species
Wilson: Realism
, andKind
. Essence
distinguished from one another? The question that preoccupies pheneticists- namely, what properties of individual organisms determine species
membership receivesonly a derivative answer from proponents of reproductive
and genealogicalviews. If one answerseither (a) or (b) or both, one
determineswhich speciesindividual organismsbelong to not by identifying
a speciesessence
, but by seeing which group, individuated in accord with
the relevant answer to (a) or (b), those organismsbelong to. Thus, "belonging
"
to can be understood in terms of part-whole relations, as it should
according to the individuality thesis. Moreover, proponents of reproductive
views conceiveof speciesaspopulations
, whereasproponents of genealogical
views conceive of speciesas lineages
and
both populations and lineagesare
,
understood
as spatiotemporal, bounded, coherent individuals, rather
easily
than as kinds, be they natural or nominal.
It is widely acceptedthat there are strong objections to the claim that any
of theseproposals- pheneticism, reproductive views, or genealogicalviews
- are
adequate.Theseobjectionshave, in turn, motivated pluralismabout the
speciescategory, the idea
. being that eachof the three views, or eachof the
more specmcforms that they may take, provides a criterion for specieshood
that is good for some, but not all purposes. The commonality assumptionis
false because
, broadly speaking, phenetic, reproductive, and genealogical
criteria focus on different types of properties for speciesmembership, so
there is no one type of property that determineskind membership. The priority
assumptionis also false becausethe different speciesconceptsreflect
the diverse biological interests of (for example) paleontologists, botanists,
ornithologists, bacteriologists, and ecologists, so these concepts depend as
much on our epistemic interests and proclivities as on how the biological
world is structured. And the ordering assumption fails becausewhere we
locate the speciescategory amongst other scientific categoriesdepends on
which researchquestionsone choosesto pursueabout the biological world .
Like pheneticism, reproductive and genealogicalviews of the speciescategory
recognize the phenotypic and genotypic variation inherent in biological
populations, so they concede that there is no traditionally conceived
essencein terms of which speciesmembershipcan be defined. But even aside
from viewing heterogeneity amongst conspecifics as intrinsic to species
,
these two views sharea further feature that makesthem incompatible with
the sort of essentialismthat forms a part of traditional realism. In contrast
with the traditional view that essencesare sets of intrinsic properties, reproductive
and genealogicalviews of the speciescategory imply that the properties
determining speciesmembershipfor a given organism are not intrinsic
. properties of that organism at all, but depend on the relations the organism
bearsto other organisms. Let me explain.
Although we are considering reproductive and genealogicalviews of the
speciescategory, I mentioned earlier that theseviews have a derivative view
of what determinesspeciesmembershipfor individual organisms. Reproductive
views imply that a given individual organism is conspecificwith organ-
192
/
HI. RethinkingNaturalKinds
isms with which it can interbreed (Mayr ), with which it sharesa mate recognition
system (paterson), or with which it has genetic or demographic
exchangeability(Templeton). Genealogicalviews imply that conspecificity is
determined by a shared pattern of ancestry and descent (Cracraft) or by a
"
sharedlineage that has its own distinctive evolutionary tendenciesand historical
"
fate ( Wiley 1978, 80). According to theseviews, conspecificity is not
'
determinedby sharedintrinsic properties, but by organisms standing in certain
relations to one another. We can see this most clearly if we consider
both views in conjunction with the individuality thesis, since conspecificity
'
is then determined by an organisms being a part of a given reproductive
population or evolutionary lineage, where neither of these is an intrinsic
property of that organism. Here, we seema long way from the traditional
.
realist' s conceptionof essentialism
a
for
more
serious
Any
integrative conception of speciesmust
proposal
reflect the inherent heterogeneity of the biological populations that are species
, and it is difficult to seehow the traditional realist view of natural kinds
can do so. Also, given the implicit commitment of both reproductive and
'
genealogicalviews of the speciescategory to an organisms relational rather
than its intrinsic properties in determining conspecificity, the prospects for
resuscitatingessentialismlook bleak.
193
/
Wilson: Realism
, and Kind
, Essence
"\'"
Figure 7.1 The neural crest. A representationof the localization of the neural crest and neural
crest cells (black) between neural ectoderm (stippled) and epidermal ectoderm (white) at neural
plate (a), neural fold (b, c) and subsequentstages(d- f ) of neural crest cell migration to i Ilustate
patternsof migration in relation to neural tube closurein various vertebrates. The time of initial
migration varies betweendifferent vertebratesand can also vary along the neural axis in a single
embryo. In the rat, cranial neural crest cells migrate while the neural tube is still at the open
neural fold stage (c). In birds, neural crest cells remain in the neural folds until they close (d),
only then beginning to migrate (f ), whereasin amphibians, neural crest cells accumulateabove
the closedneural tube (e) before beginning their migration (f ). ( Reprintedwith pennission from
Hall and Horstadius 1988.)
194
/
1lI. RethinkingNaturalKinds
195
/
Wilson: Realism
, andKind
, Essence
"
(1995) says that we now know more about the anatomicaland functional
properties of retinal ganglion cells than we do about any other neurons of
the mammalianbrain" (p. 37), suggestingthat the neural categorieshere are
the product of relatively well-developed neuroscience
. Over the last thirty
years, a number of taxonomieshave beenproposedfor retinal ganglion cells;
some of these taxonomies(e.g., alphajbeta/gammatrichotomy ) are basedon
morphological criteria, suchas dendritic morphology and axon size, whereas
others (e.g ., the Y / X / W trichotomy) are basedon physiological properties,
such as the size of the receptive field (table 7.1). The functional distinctness
of eachof thesekinds of retinal ganglion cell suggeststhat they form distinct
visual channelsthat operatein parallel in visual processing.
As with neural crest cells and their determinatekinds, such as adrenergic
and cholinergic cells, the taxonomy of retinal ganglion cells proceeds by
identifying clustersof properties that each type of cell has. No one of these
properties is deemednecessaryor any set of them deemedsufficientfor classification as a Y, X, or W cell; thus, there is no essencefor any of theseneural
categories. Again, however, I want to suggestthat it is implausibleto see,
for example, the taxa of Y cells as individuals rather than as a natural kind or
to claim that this way of categorizing retinal ganglion cells has a pluralistic
rather than a unificationist basis. The clustering of the various morphological
and physiological properties in these cells again points us to a middle
ground here. Large numbersof retinal ganglion cells tend to sharemany of a
cluster of properties in their normal environments. This fact, together with
the distinctnessof theseclustersof properties, provides the basisfor individuating
retinal ganglion cells into various kinds.
The biological facts in these areas of neurosciencedefy philosophical
views that posit traditionally conceived essences
. Equally clearly they suggest
an alternative to the corresponding individuality thesis and pluralism
about taxonomy in the philosophy of biology more generally.
HOMEOSTADCPROPERTYCLUSTERS
AND THE REVIVALOF
FSSENTIALISM
The middle-ground position I have in mind is basedon a view introduced by
Richard Boyd (1988, 1991, chapter 6 in this volume; see also Komblith
1993), which he calls the homeostatic
propertycluster(hereafter, HPC) view of
natural kinds. I shall adapt this view, noting explicitly where I depart from
Boyd. Boyd initially introduced this view as part of his defenseof anaturalistic version of realismin ethics, but from the outset he clearly intended for it
to speciesin particular. Precursorsto
, to apply to natural kinds in scienceand
the HPC view include Wittgenstein' s discussionof cluster conceptsvia the
'
; Putnams (1962) introduction of a law cluster
metaphor of family .resemblance
view of scientific concepts; and Hull ' s (1965) argument that biologists
who recognize higher taxa as cluster concepts should extend this view to
'
speciesthemselves. Boyd s previously published discussionshave been rela-
196
/
III. RethinkingNaturalKinds
Table7.1
Y Cells
XCells
Wcells
smalL10'- 10
linear
usually absent
slow, 15- 2,3m/ sec
absent
Somasize,peripheralretina
present
fast, 30- 40m/ sec
large, > 22JIm
medium
. 14- 22Jun
Proportion.of population
/0
< 100
approximately 40%
approximately
50- 55%
Retinaldistributiol1
concentratenear ~ a
centralis, more numerous
relatively in peripheral
retina
concentrateat area
centralis
concentrateat area
centralisand in streak
Central projections
to laminaeA. AI , and
CI2.of LGN; thenceto
area17; to midbrain(a
minority), but probably
not to SC
to SC, to C-laminaeof
LGN and thence
visual cortex area17
and/ or 18, and 19
Nasotemporaldivision
nasalcensproject
nasalcellsproject contralaterally
nasalcellsprojectcontralaterally
contralaterallYi most
, temporalcells
; mosttemporal
cellsipsilaterally
; stripof
temporal censalso
project ipsilaterally;
narrow strip of intermingling project contralaterallYi
interminglingcentered
about 40% of
centeredon
slightlytemporalto area
areacentralis
centralis
temporal censproject
ipsilaterally
tively programmatic, and his current view of the implications of the HPC
view for issuesconcerning species(see Boyd, chapter 6 in this volume) is
somewhatdifferent from the view I advocatehere.
The basic claim of the HPC view is that natural kind terms are often
definedby a cluster of properties, no one or particular n-tuple of which must
be possessedby any individual to which the term applies, but some such
n-tuple of which must be possessedby all such individuals. The properties
mentioned in HPC definitions are homeostaticin that there are mechanisms
'
that causetheir systematiccoinstantiation or clustering. Thus, an individual s
possessionof anyone of these properties significantly increasesthe probability
that this individual will also possessother properties that featurein the
definition. This is a fact about the causalstructure of the world : the instantiation
of certain properties increasesthe chancethat other particular properties
will be coinstantiated becauseof underlying causalmechanismsand
processes.
'
197
/
Wilson: Realism
, and Kind
, Essence
The view is a " cluster" view twice over: only a cluster of the defining
properties of the kind needbe presentfor an individual to fall under the kind,
and such defining properties themselvestend to cluster together- that is,
tend to be coinstantiatedin the world. The first of thesefeaturesof the HPC
view of natural kinds allows for inherent variation among entities that
belong to a given natural kind.
The secondof thesefeaturesdistinguishes the HPC view as a realisticview
of kinds from the Wittgensteinian view of conceptsmore generally to which
it is indebted. On the HPC view , our natural kind conceptsare regulated by
information about how the world is structured, not simply by conventions
we have establishedor languagegameswe play. Before moving to the case
of species
, considerhow the HPC view appliesto our pair of neural kinds.
First, take the caseof the individuation of neural crest cells. For a cell to be
adrenergicis for it to have a certain cluster of properties that scientistshave
discovered; amongst other things, it is to originate in the posterior of the
neural tube, to follow one of a given number of migratory paths, to function
in the sympathetic nervous system, and to produce the neurotransmitter
norepinephrine. Factsabout the structure of the biological world - facts still
being uncovered- explain why these properties tend to be (imperfectly)
coinstantiatedby certain kinds of cells. This clustering is the result of incompletely
understoodmechanismsthat govern an embryo' s developmentand is
absent, either partially or wholly , just when the normal function of those
mechanismsis disrupted. No single one of these properties is, however,
strictly necessaryfor a cell to be adrenergic. The presenceof all of them,
however, is sufficientfor a cell to be adrenergic, at least in the environments
in which developmentnormally occurS.3 This feature of the HPC view marks
one of the affinities between it and traditional realism, about which I say
more later. On this view, adrenergicneural crest cells are a natural kind of
cell, and individual cells are membersof that natural kind in virtue of satisfying
the homeostaticproperty cluster definition of that natural kind.
Second, take the caseof the individuation of retinal ganglion cells. Consider
in particular the physiological taxonomy of Y, X, and W cells. The tendency
of the various physiological properties- such as the axonal velocity ,
soma size, and retinialdistribution- to be coinstantiated by particular types
of cells is no accident, but the result of underlying mechanismsgoverning
neural development and neural functioning. Again, a determinate form of
anyone of these properties could be absent in a particular cell, yet the cell
will still be a certain kind of cell- say, a Y cell- so no oneof theseproperties
is an essentialproperty for being a member of that kind of retinal ganglion
cell. Nevertheless, there is a general definition of what it is to be a Y
cell, one basedon the homeostaticcluster of properties that one finds instantiated
in somecells and not inothersY cells are a natural kind of cell with a
sort of essence
, albeit one different from the sort of essencecharacterizedby
traditional realism. Moreover, there is a kind of integrity to being a Y cell
that .invites a unificationist rather than a pluralistic view of it .
198
/
III. RethinkingNaturalKinds
A more ambitious way to apply the HPC view to this example is worth
noting . Although it is a substantivehypothesis that the morphological and
physiological taxonomiesof retinal ganglion cells are roughly coextensive, it
is a hypothesis that is reasonably well confirmed (seeChalupa 1995, 40- 42).
The HPC view provides a natural way of integrating the two taxonomiesin
effect by adding together the two lists of properties in each cluster. This
, of course, that certain common mechanismsexplain the
integration assumes
new
cluster of properties qua cluster, without which we
this
of
presence
would simply have a disjunction of two homeostatic clusters, not a new
homeostaticcluster of properties.
I suggest that the HPC view applies to speciestaxa as follows. Particular
speciestaxa are natural kinds defined by a homeostatic cluster or morpho
.
This
features
and
reproductive
logical, genetic, ecological, genealogical,
cluster of featurestend to be possessedby any organism that is a memberof
a given species
, though no one of these properties is a traditionally defined
essentialproperty of that species
, and no proper subsetof them is a species
is
caused
This
.
essence
by only partially understood mechanisms
clustering
that regulate biological processes (such as inheritance, speciation, and morphological development) and the complex relations between them. More
generally, the homeostatic clustering of these properties in individuals
belonging to a single speciesis explainedby facts about the structure of the
biological world. For example, organismsin a given speciessharemorphol
ogy in part becausethey share genetic structures, and they share these
structuresbecauseof their common genealogy. This is not to suggest, however
, that anyone of theseproperties is more basic than all of the others or
that there is some strict ontological hierarchy on which they can all be
placed, for the dependencyrelations between these properties are complex
and almost certainly multifarious.
Having severedthe connectionbetween the HPC view and traditional realism
, let me now indicate an important affinity that the two views share.
Although possessionof individual properties or n tupies of the relevant
homeostatically clusteredproperties are not necessaryfor membershipin the
corresponding specieskind, possessionof all of them is sufficient for membership
in that kind. If the homeostatic property cluster definition is sufficiently
detailed, this circumstancewill likely remain merely an idealization,
uninstantiatedin fact and approximated to a greater or lesserextent in particular
cases.This in turn points to one way in which the sort of essentialism
that forms a part of traditional realism is a limiting caseof the sort of essentialism
implicit in the HPC view of natural kinds.
The HPC view can also be applied to the speciescategory, allowing a
definition of what sorts of thing a speciesis that marks it off from other biological
categories. First, the general nature of the cluster of properties
JDorphology, genetics, genealogy, and so on- will distinguish speciesfrom
nonevolutionary natural kinds, suchascells(in physiology), predators(in eco
logy ), and diseases(in epidemiology). Second, specieswill be distinguished
-199
Wilson: ~ iSm
. Essence
, andKind
200
1lI. R( hinkingNaturalKinds
Note how the HPC view of natural kinds preservesanother idea that is a
part of traditional realism: all and only membersof a natural kind satisfy the
corresponding definition of that kind. Anything that is a speciesand only
things that are specieswill satisfy the HPC definition for species; any individual
that is a memberof a particular speciesand only such individuals will
- likewise for neural crest
satisfy the HPC definition for a particular species
cells and retinal ganglion cells (as well as their determinateforms).
But what does it mean " to satisfy" such a definition1 Thus far, I have
"
"
"
"
implied that to satisfy is to possess enough of the properties specified
in the HPC definition. Here, we might suspect the vaguenessthis implies
regarding (say) the delineation of the speciescategory and membershipin
'
particular speciestaxa is the Achilles heel of the view. I want to offer two
responsesto this concern.
"
"
First, what counts as having enough of the relevant properties- as with
what are the relevant properties in the first place - is an a posteriori matter
determined in particular casesby the practitioners of the relevant science
rather than by philosophers with a penchant for crisp universality. There
need be no one answer to the question of what is " enough," but whatever
answersare given in particular caseswill be responsiveto the clusters that
one finds in the world .
Second, even once there is general agreement about what counts as
"
"
enough, there clearly will be casesof genuine indeterminacy with respect
to both the speciescategory and membershipin particular speciestaxa. Yet
this indeterminacy seemsto me to reflect the continuities one finds in the
complex biological world , whether one is investigating species, neurons, or
other parts of the biological hierarchy. There will be genuine indeterminacy
about the rank of given populations of organisms, and particular organisms
may in some casessatisfy more than one HPC definition for particular species
taxa. The former of these indeterminacies
, however, is a function of the
fact that under certain conditions and over time varieties becomespecies
, and
the descendantsof a given speciesbecome membersof a particular genus;
the latter reflects the process of speciation (and its indeterminacies
) more
directly.
2ql
their view negative (speciesare not natural kinds), another part positive
(speciesare individuals). For example, it has been argued that the heterogeneity
within biological populations implies that speciesare not natural kinds
and that their status as historical entities within evolutionary theory supports
a view of them as individuals. Insofar as the former types of argument
presumea two -way conceptualconnection between traditionally conceived
essencesand natural kinds, they carry no force against the view that species
are HPC natural kinds. Thus, the view I have defendedunderminesnegative
arguments for the individuality thesis. But the HPC view of natural kinds
also shows both types of argumentsfor the Ghiselin-Hull view in a new light
becauseparity of reasoningshould lead one to abandonthinking of neuronal
populations as natural kinds and embracea view of them as individuals. Of
course, such parity considerationscan always be undermined by the differences
betweenhow the term species
is used within evolutionary biology and
how (say) the term retinalganglioncellis usedwithin visual neuroscience
. The
HPC view, however, places the burden on those who think that there is
sometlung specialabout speciestalk that warrants a unique ontological view
of speciesas individuals to show this uniqueness
.
Alternatively , perhaps reflection on the neuroscientmccasesshould lead
one to extend the individuality thesis beyond the caseof speciesto other
biological categories. Interestingly, at least some researchersin the relevant
neurosciencemay be amenableto this idea. For example, following Tyner
(1975), Rowe and Stone (1977, 1980a, 1980b) advocate what they call a
parametricor polythetic approach to the individuation of retinal ganglion
cells, viewing these cells not as kinds with some type of essence
, but as
of
cells
that
have
their
own
internal
intrinsically heterogenouspopulations
coherenceand duration. (Indeed, Rowe and Stone explicitly take their cue
&om the modem speciesconcept.) The problem with sucha view, it seemsto
me, is that central to neural taxonomy is the idea of identifying categoriesof
cells that at least different organisms in the same speciesinstantiate, and
these instancesconsideredtogether do not form an individual. For example,
your adrenergic cells and my adrenergic cells considered together are not
spatially bounded, occupy different temporal segments, and do not form an
integrated whole. Perhapsthis points the way to how the positive arguments
for the individuality thesis can be sharpenedin light of the parity
considerationsintroduced with respectto the negative part of the argument
for the thesis.
PLURALISM
To remind you of what the pluralist holds about species, consider what
Dupre (1993 ) says in articulating his version of pluralism :
There is no God - given , unique way to classify the innumerable and diverse
products of the evolutionary process. There are many plausible and defensi-
202
1lI. RtthinkingNaturalKinds
ble ways of doing so, and the best way of doing so will dependon both the
purposesof the classificationand the peculiarities of the organismsinquestion
. . .. Just as a particular tree might be an instanceof a certain genus (say
Thuja) and also a kind of timber (cedar) despite the fact that thesekinds are
only partially overlapping, so an organism might belong to both one kind
defined by a genealogicaltaxonomy and another defined by an ecologically
driven taxonomy. (p. 57)
In introducing pluralism as the denial of either or both of two assumptions
central to traditional realism- the priority and ordering assumptions- 1
meant to suggest that there is some tension between pluralism and realism
punkt. The metaphysicalangst that many realists experiencewith pluralism
concernsthe extent to which one can make senseof the idea that there are
"
"
incompatible but equally natural (i.e., real) ways in which a sciencecan
taxonomize the entities in its domain. There is at least the suspicion that, to
"
'
use Dupre s terms, pluralism is driven more by the purposesof the classifi'
"
"
cation than by the peculiarities of the organismsin question," as Dupre s
own analogy suggests. In rejecting the priority assumption, such pluralism
would move one from a realist view toward a nominalist view of species(see
"
Wilson 1996; d . Hull , chapter2 in this volume).
Yet the most prominent forms of pluralism about specieshave all labeled
' "
"
' "
"
themselves realist," from Dupre s promiscuousrealism to Kitcher s pluralistic realism." Moreover, Boyd (chapter 6 in this volume) views at least
'
Kitcher s brand of pluralism as compatible with his own articulation of the
HPC view of natural kinds- suggesting a form of realism that acceptsthe
priority assumption, but rejectsthe ordering assumption. The idea that Boyd
and Kitcher share is one Mishler and Donoghue express(cited earlier): the
various speciesconcepts that one can derive and thus the various orders
within which one can locate speciesare merely a reflection of complexities
within the biological world . This view has two problems- one with pluralistic realism itself, the other with viewing the HPC view of natural kinds as
compatiblewith suchpluralistic realism.
As pluralists say, one can arrive at different speciesconceptsby emphasizing
either morphological, reproductive, or genealogicalcriteria for the species
category. Yet it is difficult to seehow the choicesbetween thesesorts of
alternativescould be made independently of particular researchinterestsand
epistemicproclivities, which calls into question the commitment to the priority
assumptionthat, I claim, needsto be preservedfrom traditional realism
in any successorversion of realism. Perhapspluralistic realists would themselves
'
reject the priority assumption, although Boyd s own emphasison what
"
"
he calls the accommodationdemands imposed by the causalstructure of
the world on inductive and explanatory projects in the sciencessuggestthat
he himself acceptssomeversion of the assumption.
'
~oyd s own view of the compatibility of the two views seemsto me to fail
to capitalize on the integrationistpotential of the HPC view, one of its chief
appeals. One of the striking featuresof the various definitions of the species
203
/'
Wilson: Realism
, and Kind
, Essence
category is that the properties that play central roles in eachof them are not
independent types of properties, but are causally related to one another in
various ways. These causalrelationships and the mechanismsthat generate
and sustain them form the core of the HPC view of natural kinds. Because
the properties specified in the HPC definition of a natural kind term are
homeostatically related, there is a clear sensein which the HPC view is integrationist or unificationist regarding natural kinds. By contrast, consider the
view of pluralists. Kitcher (1984) says that we can think of the speciesconcept
as being a union of overlapping speciesconcepts(pp. 336- 337; d . Hull
1965), so it is unified in some sense, but without a further emphasison
something to play the metaphysicalrole that underlying homeostaticmechanisms
play in the HPC view, the unity to the speciesconcept remainsallusive within Kitcher' s view.
Consider how the differencesin views manifest themselvesin a concrete
case- whether asexualclonelinesform species. For the pluralist, the answer
to this question dependson which speciesconcept one invokes- in particular
, whether one appealsto interbreeding criteria to define the speciescategory
. By contrast, on the HPC view, asexualclonelinesare speciesbecause
they share in the homeostatic cluster of properties that defines the species
'
category, even though they don t have at least one of those (relational)
properties, interbreeding.
Likewise, consider the issue of whether there is a qualitative difference
between speciesand other (especially higher) taxa (see Ereshefsky
, chapter
11 in this volume). Again, a natural view for a pluralist to adopt is that how
one construesthe relationship between speciesand other taxa dependson
which speciesconcept one invokes. For example, on Mayr' s biological species
concept, specieshave a reality to them provided by their gene flow and
its boundaries, which higher taxa lack; alternatively, pheneticistsview both
speciesand higher taxa asnominal kinds becausetaxa rank is determinedby a
conventional level of overall phenetic similarity. By contrast, on the version
of the HPC view of speciesI have defended, although the general difference
between various taxa ranks will be apparent in their different HPC definitions
, there will be caseswhere questions of the rank of particular taxa
remain unresolvedby the HPC view.
CONCLUSION
My chief aims here have been to clarify the commitmentsof a realist view of
natural kinds and to suggest a way of modifying rather than abandoning
traditional realism in light of the challengeof biological heterogeneity. Both
the individuality thesisand speciespluralism seemto me to be extreme reactions
to the failure .of traditional realism in the biological realm, but I have
stopped short here of trying to make a full casefor the middle-ground position
I have advocatedas an alternative to both of theseviews. That remains
for ~ other day.
204
III. R~ inkingNaturalKinds
ACKNOWLEDGMENTS
Precursors to this paper were presented at colloquia in the philosophy
'
departments at the University of Western Australia, Queen s University ,
Canada, and the University of Colorado, Boulder in 1995- 96; early thoughts
about the examplesdiscussedin the third section were first presentedat the
Cognitive Studies Workshop at Cornell University in May 1991. I would
like to thank audienceson all four occasionsfor useful feedback
, as well as
Allison Dawe, David Hull , Ed Stein, and Kim Sterelny for helpful comments.
NOTES
1. Later, I return to this appearanceand the putative contrast between traditional realism and
pluralism when I considerso-called pluralistic or promiscuousrealism.
2. J. D. Trout ' s (1988) brief ms Msion of neural crest cells, including the distinction between
adrenergic and cholinergic cells, stimulated my interest in this example; I also owe Barbara
Finlay a note of thanks for helpful, early discussionsof both this and the following example.
Neither should be landedwith the interpretation I give to the example.
3. Alternatively , one could make the normal developmentalsequencepart of the homeostatic
property cluster definition itself. In either case, the sufficiencyis not one that allows for a consid~ tion of all logically or even nomologically possiblecases.
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of nerve
Rowe, M. H., andJ. Stone(1980b
). The interpretationof variationin the classmcation
17, 123- 151.
cells. Brain,Behaoior
, andEoolution
47, 350. Philosophy
of Science
Sober
, populationthinking
, E. (1980). Evolution
, andessentialism
.
1992
in
383. Reprinted Ereshefsky
. In Otte and
: A geneticperspective
andspeciation
ton, A (1989). Themeaningof species
Temple
. Reprintedin Ereshefsky
land, Mass.: Sinauer
. Sunder
andits consequences
Endier
, eds., Speciation
1992.
sensepsychology
of common
: A scienti Bcdefense
, andmethod
Trout, J. D. (1988). Attribution, content
.
York
New
,
. PhiD. thesis
, CornellUniversity,Ithaca
: Applicationsof taxonomictheoryto the studyof
Tyner, C. F. (1975). The namingof neurons
12, 75- 96.
. Brain,Behaoior
, andEoolution
cellularpopulations
. Systematic
Zoology27,
Wiley, E. O. (1978). The evolutionaryspeciesconceptreconsidered
1992.
17- 26. Reprintedin Ereshefsky
. New York: Cambridge
andphysicalminds
University
Wilson
, R. A (1995). Cartesian
psychology
.
Press
47,
. BritishJournalfor the Philosophy
realism
of Science
Wilson
, R. A. (1996). Promiscuous
303 316.
207
/
Wilson
: Realism
, and Kind
, Essence
TO INDMDUALISM
FROM~ ENTIALISM
Chemicalelementsand biological specieshave been the two stock examples
of natural kinds &om Aristotle to Putnam. Samplesof lead or individuals of
"
the speciesPantroglodytesare not only similar in various respects, but are of
"
the samekind in somemuch.deepersense. One way to expressthis deeper
commonality is to say that the membersof a kind sharean essence- a property
common to all the membersof a kind and responsiblefor eachmember
being the kind of thing that it is. My gold watch resemblesyour gold navel
ring in many respects, someknown to us and somenot, becausethe atoms of
: their atomic number. Essentialism
which both are composedshare an essence
ErasmusresemblesSocks the
kitten
that
in biology would suggest
my
cat becausethey too share an essence(albeit a less well understood one).
Essentialismtook on a new life in the 1970s, largely becauseof the work of
the philosophers Saul Kripke (1980) and Hilary Putnam (1975). Biological
specieswere one of the stock examplesin this essentialistliterature, even
though by this time essentialismwas regardedby many biologists as inconsistent
with the basictenets of Darwinism!
The perceived antithesis between evolution and essentialismwas largely
due to the work of Ernst Mayr (1959). Mayr argued that biology before
Darwin was characterizedby typologicalthinking in which types or kinds of
organismshad ontological. and explanatory priority over concrete individuals
. Darwinian populationthinking gives populations of concrete individuals
ontological and explanatory priority instead. Elliott Sober has argued convincingly that the core of population thinking is the Darwinian approachto
variation (Sober 1980). The typo logical approachexplains the resemblances
"
"
betweenthe individuals in a speciesin terms of the underlying natural state
betweenatoms
of eachindividual, just aschemistry explainsthe resemblances
The
.
microstructure
shared
of
their
in
terms
element
of the same
typo logical
approachexplains variat~ons between the individuals in a speciesas perturb
~tions of the natural state of that species.The Darwinian approachexplains
both resemblanceand variation at the population level. Organisms resemble
one another not becauseof something inside each of them, but becauseof
something outside eachof them: the genealogicaland ecological factors that
make these organismsa population or a group of related populations. The
properties that differ between individuals are ontologically on a par with
those properties they share. Variation is not noise obscuring the essential
samenessof the membersof a species
, but an important, heritable property
of populations consisting of the aggregatereal differencesbetween its members
. Sober concludesthat becausethese explanations of samenessand difference
are central to the Darwinian tradition, Mayr is correct in concluding
that Darwinism precludesidentifying any phenotypic or genotypic features
as a speciesessence
. However, Sober notes that it would be quite consistent
to be a Darwinian essentialist, given the right choice of essentialproperties
(1980, 209). Population thinking excludesessentialintrinsic properties, but it
does not excludeessentialrelational properties. This paper defendsjust such
a relational essentialism
.
Mayr famously tried to characterizethe relational properties that unite the
membersof a species. His biologicalspeciesconcept(BSC) defined speciesas
I'
groups of actually or potentially interbreeding natural populations which
are reproductively isolated from other such groups" (Mayr 1940, cited in
Mayr 1963, 19). Underlying this formal definition of speciesin terms of interbreeding
is the idea of a genealogicalnexus. A nonreproductive worker in
one beehiveneither actually nor potentially interbreedswith nonreproductive
workers in other hives, but that individual is united in a genealogicalnexus
with reproductiveswho actually or potentially interbreedwith reproductives
in other hives. Thesereproductivesin turn are united in a genealogicalnexus
with nonreproductives in their hives, so the several nonreproductives are
membersof the samespecies.Attempts to extend the BSC to asexualspecies
also rely on this underlying genealogicalelementin Mayr' s speciesconcept,
. According to such proposals,
using it as one of two criteria for specieshood
an asexualspeciesis a well-defined segmentof a genealogicaltree of asexual
individuals that meets some other criteria, such as containing individuals
roughly as morphologically similar to one another as membersof a sexual
species.This secondcriteria is designedto distinguish the species-level genealogical tree segmentfrom the larger segmentsin which it is embeddedand
from the smallersegmentsthat it embeds. Robert Brandonand Brent Mishler
(1987) have generalizedthese two criteria for specieshoodinto groupingcriteria
and rankingcriteria, and they have argued that any speciesconceptmust
have both a grouping and a ranking criteria. In most modem speciesconcepts,
including modem versions of the BSC, the grouping criteria is genealogical.
. Speciesmust be characterizedby some version of monophyly descent
or
similar
from a single population, a single speciation event,
any
unique
point of origin . The ranking criteria serves to distinguish species from
equally monophyletic genera, families, and so forth. Although there is some
disagreementover the best definition of species-level monophyly, the main
disagreementsbetweenthe twenty or so current speciesconceptsare in their
210
/
III. RethinkingNaturalKinds
different ranking criteria. (For a very clear look at species- level monophyly,
seeKomet 1993.)
Individualismabout speciesis an idea with close links to antiessentialism
,
both conceptually and historically. Individualists argue that speciesare not
kinds or types at all, but big individual objects. Organisms are not members
of a species, but parts of a species. The individualist argumentsof Michael
Ghiselin (1974a, 1974b) and David L. Hull (1976, 1978) strongly resembled
someof the earlier argumentsagainst essentialism
. They argued that species
must be able to evolve and that kinds or types do not evolve. Speciesmust
be able to undergo unlimited changein any of their genetic or phenotypic
characters
. If specieswere kinds or types of
, not only in peripheralcharacters
organism, then it would not be speciesthat evolved, but organismsor populations
that changed from one speciesinto another. The kinds or types
would form a sort of biological absolute spaceagainst which evolutionary
change occurred. Hull and Ghiselin also pointed to the practical failure of
attempts to define speciesby lists of charactersor statistical clustersof characters
. This argument took on new force in light of the cladistic revolution in
'
, in which attempts to discerncommon descentreplacedattempts
systematics
to identify taxa by statistical clusters of characters. These and other arguments
convinced the individualists that speciescould be defined only in
terms of the pattern of ancestryand descentamong organisms, but the next
step in their reasoningis the most relevant to the issuesof this paper. Hull
and Ghiselin concludedthat becausespeciesand other taxa must be defined
in terms of genealogy, they must be moved from the ontological category of
types or kinds to the category of individual objects. If taxa are genealogically or historically defined, then they cannot be natural kinds.
This last step in Ghiselin and Hull ' s argument depends on a traditional
conception of natural kinds in which they are the subjectsof spatiotemporally unrestricted laws of nature. If natural kinds are to figure as the subject
of universal laws, they themselvesmust have universal applicability. Laws
that make ineliminable mention of things that can exist only at a particular
location in time and spaceare not, in the relevant sense, universal laws. If
Ghiselin and Hull are correct, then biological taxa have just such a unique
origin in spaceand time. No part of a taxon can exist outside the cone of
causalinfluence extending from its origin event, so taxa are restricted to a
.particular portion of space-time and cannot be mentioned in genuine laws of
nature.
The conclusion that there are no laws of nature concerning taxa has been
welcomed by many theorists as part and parcel of antiessentialism
. Hull
that
is
such
has
welcomed
the
conclusion
there
no
198
( .6)
liberatory
thing as
"
"
human nature. Attempts to distinguish normal from abnormal humansare
simply misguided. John Morss (1992) has argued that there are no laws of
ontogeny and particularly of child development. We should be suspiciousof
theories that describea seriesof stagesthrough which every child passesto
reachmaturity . The downside of the anomalousnessof biological taxa is that
211
/
Griffiths: Squaringthe Circle
212
/
HI. Rethinking Natural Kinds
213
Griffiths
: SquaringtheCircle
214
III. RethinkingNaturalKinds
215
/
Griffiths:Squaringthe Circle
216
/
III. RethinkingNaturalKinds
toothless. It does not licensethe conclusion that any way of classifying nature
is as good as any other. Natural kinds are ways of classifying the world
that correspondto some structure inherent in the subject matter being classified. They contrast to arbitrary schemesof classificationabout which the
nominalist claim that the membersof a kind share only a name is actually
true. Furthermore, the minimal account of naturalnesslends itself to successive
restrictions that allow us to distinguish between kinds of greater or
lessernaturalnessand henceof greater or lessertheoreticalvalue.
Although it is not possiblein this essayto give an adequatetreatment of
the principles for choosingbetween alternative taxonomiesof nature, a brief,
generaloutline may be helpful. The value of a lawlike generalizationcan vary
along two independentdimensions, which we might call scopeandforce. Force
is a measureof the reliability of predictions made using that generalization.
Scopeis a measureof the size of the domain over which the generalizationis
applicable. A theoretical category about which there are generalizationsof
considerablescopeand force is more natural than one about which generalizations tend to have more restricted scopeand lesserforce. For example, the
"
"
claim that cladistic taxonomy is maximally predictive of the unobserved
properties of taxa is intended to show that cladisticsis superior to other systems
in terms of force. There will not always be a clear winner when we
compare two sets of theoretical categorieson the basis of scope and force.
Scope and force may trade off against one another. The scope of general
the
include
than
rather
izations madewith one set of categoriesmay overlap
scopeof generalizationsmade with the other taxonomy so that neither tax
onomy can be discardedwithout loss of understanding.
Theoretical categoriescan also differ in the number of generalizationsinto
which they enter so that one category can seemthe focus of a richer scientific
project than another, irrespective of comparisonsof the strength of the
generalizationsthey yield . Finally, theoretical categoriesare tied up in wider
researchprograms whose relative prospectsmay causeus to prefer that set
of categoriesto another despite a paucity of currently establishedgeneralizations about the preferred set of categories. None of these considerations,
however, refutes the basic idea that some theoretical categoriesare superior
to others and that some are of no foreseeablevalue whatever. Even if different
are
categoriesare valuablefor different purposes, it is still true that some
no
foreseeable
have
some
that
better for a particular purposethan others and
use at all. Embodying these ideasin the languageof natural kinds links it to
a broadly realist perspectivein which the predictive and explanatory value
of categoriesis taken to be prima facie evidence that they capture part of
"
"
th~ structure of the world. The enthusiasmfor natural kinds embodiesthe
realization that there is more structure in the world than can be capturedby
a.single taxonomy of nature.
. Richard Boyd has outlined a similar conception of natural kinds using his
idea of causalhomeostasis
(Boyd 1991). According to Boyd, we judge a kind
217
/
Griffiths:SquaringtheCircle
218
/
DI. Rethinking Natural Kinds
'
Boyd s proposal is a substantialrevision of the traditional ideasof essence
"
"
and natural kindhood (see also Boyd, chapter 6 in this volume). Natural
kinds that have never been seen before can be created by social processes
unique to a particular society. The fact that people think certain things form
a kind can function as the essenceof that kind! The justification for these
conceptualrevisions is that they allows insights about the formation and use
of theoretical categoriesto be extended to the special sciencesrather than
. The
restricted to a (dwindling ) core of kinds with microstructural essences
'
for
a
continuity
Frank
Keil
has
used
s
ideas
to
C.
1989
)
(
Boyd
argue
psychologist
between category formation by developing children and category formation
in science(seealso Keil and Richardson, chapter 10 in this volume). I
have argued that the formation of theoretical categories in psychology,
including categories unique to particular cultures, is best understood as a
searchfor causalhomeostasis(Griffiths 1997).
In this section, I have tried to motivate a very general conception of
natural kinds, one that discardsmany of the traditional associationsof the
natural kind concept. Natural kinds are neededfor induction and explanation.
They representtheoretical categoriesthat we judge to be projectable, which
requires them to enter into lawlike, counterfactual supporting generaliza
tions. It does not require that these generalizationsbe universal, deterministic
laws: lawlike generalizations of more limited scope and force are
enough. Finally, kinds are defined by the processes that generate their
instances, and for many domains of objects, these processes are extrinsic
rather than intrinsic to the instancesof the kind. The causal homeostatic
mechanismthat guaranteesthe projectability of a kind plays the traditional
.
role of an essence
, but it neednot be a traditional, microstructuralessence
HISTORICAL
ESSENCES
Cladistic taxa and parts and processes deAned by evolutionary homology
. Nothing that does not sharethe historical origin of
have historical essences
the kind can be a memberof the kind. Although Lilith might not have been a
domestic cat,1 as a domestic cat she is necessarilya member of the genealogical nexus between the speciation event in which that taxon originated
and the speciationor extinction event at which it will ceaseto exist. It is not
possible to be a domestic cat without being in that genealogicalnexus. Furthermore
, cladistic taxa and parts and processes deAned by evolutionary
homology have no other essentialproperties, which is why processstructur
alists such as Goodwin and Webster do not think that these categoriescan
b~ adequatefor developmentaland structural biology . They do not seewhy
kinds whose only essentialproperties are historical should be the subjectsof
lawlike, counterfactual-supporting generalizationsabout morphological and
.physiological properties. Yet there is a well-known Darwinian ground for
expecting groups deAned by common descent to share morphological and
:
physiological characters
"219
/
Griffiths:SquaringtheCircle
~I,;~,.
Re~.....IAU
~ Natural Kinds
221
Griffiths:~
Jarin
~ theCircle
WHY GHISELIN
AND HULLWEREWRONG
Antiessentialistsand individualists about biological taxa were wrong to suppose
that there are no lawlike generalizationsabout thesetaxa. A hierachical
taxonomy based on strict phylogenetic prindples will collect more of the
correlations between characters
, from molecular to behavioral, than any
other taxonomy we know how to construct. Such a taxonomy will group
organismsinto natural kinds becauseit will predict with considerableforce
many properties of individuals. Although such a taxonomy will predict the
properties of unobservedgeneraor spedes, it will function most powerfully
in predicting the properties of new membersof taxa at or below the spedes
level. A number of competing (though not necessarilyexclusive) explanations
of the specialstatus of spedesare embodied in some of the twenty or
so currently proposed spedes concepts. These explanations draw attention
to causalprocesses such as gene exchange( biological speciesconcept) or
selection for the requirementsof a niche (ecological speciesconcept). These
mechanismsreinforce phylogenetic inertia in keeping the membersof a spedes clusteredtogether in the spaceof biological possibilityd . de Queiroz,
chapter3 in this volume).
Generalizationsabout taxa are exception-ridden. This does not, however,
prevent them from being lawlike or having counterfactualforce. The causal
homeostaticmechanismsof taxa license the prediction that a new bird will
detect its prey using visual cues or that in a new cephalopod, the blood
vesselssupplying the retina will lie under rather than over it. The causal
homeostaticmechanismsalso make it legitimate to extrapolate experimental
results to other membersof the sametaxon, espedally at the spedes level.
The fact that such predictions and extrapolations are not absolutely reliable
is simply beside the point . They are more reliable than chance, so unless
there is some other way to capture the sameregularities, eschewingthe use
of thesecategorieswould meandiscarding someof our understandingof the
structure of nature.
Parts and processes defined by evolutionary homology can be used for
explanation and induction for the samereasonthat historically defined taxa
can be used: phylogenetic inertia licensesthe extrapolation of morphological
.
and physiological properties in categories defined by common ancestry.
Also, among theseproperties are the very developmentalprocesses that are
likely to explain the phenomena of phylogenetic inertial Developmental
processes, as much as other anatomical or physiological kinds, can be
expec~ed to reflect phylogeny. What lies at the bottom of all thesephyloge-
222
III. Re~
Natural Kinds
netic patterns is, after all, the fact that related organisms inherit similar
developmentalresources. Plant physiology, for example, does not converge
on animal physiology whenever it would be adaptively useful for it to do so
becauseplant cells inherit a range of membranetemplates, organelles, genes,
and so forth that are fundamentallydifferent from those inherited by animal
cells.
If historical kinds are natural becauserelated individuals inherit similar
, it might seempossibleto define the kinds in terms
developmentalresources
of the developmentalresourcesthat underly them. This proposal would treat
the shared developmental resourcesof a taxon as the causal homeostatic
mechanismof that taxon, a mechanismthat takes the form of a traditional
microstructural essencepossessedby all and only the membersof the taxon.
I suspectthat this thought is at the back of severalstructuralist criticisms of
the use of historical kinds in biology . The structural or developmentalbiologist
seesthat the processes they are investigating explain the fact that members
of a taxon sharea rich cluster of properties, which suggeststo them that
the real essenceof the taxon is not its sharedhistory, but its shareddevelopmental
processes. In the next section, I show that this very natural line of
reasoning is mistaken becauseof the original, Darwinian considerations
againstessentialismoutlined at the beginning of the paper.
WHYGHISELIN
ANDHULLWERERIGHT
Darwinians will resist the suggestion that taxa be defined developmentally
becausethey expect developmentalprocesses to be just one more product
of evolution. As such, they expect developmentalprocesses to display variation
between individuals in natural populations, just as other charactersdo.
Empirically, they do not expect to find a list of developmental properties
possessedby all and only the members of a speciesany more than they
expect to find lists of phenotypic or genotypic characterspossessedby all
and only membersof a species.Conceptually, even if sucha list of properties
existed for a species, it would be an accidentalnot an essentialmatter. An
individual united in a genealogicalnexus with the existing membersof the
, but lacking someproperty on the list, would still function as amember
species
of the sameevolutionary unit. The purpose of the speciesconcept for a
Darwinian is to describethe units of evolution, and essentialistspeciesconcepts
fail to do this.
There are a number of reasonswhy Darwinians have wanted to take a
more developmentalperspectiveon evolution. A developmentalperspective
highlights the problems with an atomistic approachto the evolution of characters
, in which eachcharacteris assumedto be optimized independently of
the others (Gould and .Lewontin 1979, Lewontin 1983). A developmental
perspective can also draw attention to the wide range of developmental
resourcesother than genesthat can be the subject of evolutionary explanatio~ (Griffiths and Gray 1994). Perhapsmost importantly, a developmental
223
Griffiths
: ( uaringtheCircle
THESTRUCTURAL
HOMOLOGY
CONCEPT
I have defended the view that historically defined taxa are natural kinds and
the corollary view that evolutionary homologues are also natural kinds . I
have defended these views against some arguments associated with structur alist approach es to biology . In this closing section, I want to consider two
other , recent arguments that biology needs a structural homology concept .
The first argument suggests that the evolutionary homology concept is
. somehow unworkable without a prior conception of structural homology .
This argument is mistaken , but a second, better argument points to the
' potential value of a structural homology concept , including its value in il Iu
. minating the basis of the evolutionary homology concept .
The mistaken argument , which we can perhaps regard as put forward by
a hypothetical structuralist strawman , is that because candidates for evolu -
224
./
RethinkingNaturalKinds
Griffiths~~
g the Circle
the body are passedon from generation to generation for millions of years
as coherentunits of evolutionary change" (p. 279).
It is this sort of question that has been the focus of Wagner' s more recent
work on the evolution of modularization and canalization of development
( Wagner1996, Wagner et al. 1997). Wagner (1994) rejects an analysis of
vertical characteridentity basedon identical developmentalorigin, a modem
derivative of the traditional practice of judging homology from the relative
'
position of parts in the embryo: Too often do we find substantial developmental
variation among structurally, and presumably phylo genetically,
identical body parts" (p. 276). He would presumably reject an account of
vertical character identity based on identical genetic causesfor the same
reason: homologous characterscan persist through substantialchangesin the
genetic inputs to their development. In place of such ideas, Wagner sets up
the goal of understandingwhy organismshave come to have discrete, reidentifiableparts. A theory of why there are parts will tell us how those parts
can be naturally taxonomized. Wagner's researchprogram is thus (quite selfconscious
) a searchfor natural kinds construed as the objects of lawlike
generalizations.
"
Wagner' s work is an instance of what I have described as Darwinian
"
developmentalism becausehe looks for the origins of these units of
structural homology in the evolutionary processrather than in a system of
ahistorical biological types like the system postulated by the process
structuralists Goodwin and Webster. Wagner and his collaborators have
tried to model selective processes that favor the emergence of discrete
"
"
developmental modules that are stabilizedagainst various perturbations of
the developmental system. Although these modules function as developmental
invariants in at least some ti mescal
es, persisting with apparentdisregard
for the " conditions of life," they have themselvesemergedas a result of
the evolutionary process, and they will possessa phylogenetic signature---an associationwith a particular lineage.
ACKNOWLEDGMENTS
I have bene6ted horn the comments of Rob Wilson , Gunther Wagner , and
Karola Stotz on an earlier draft of this paper .
NOTE
1. Shecouldexistevenif domesticcatshadspeciated
somegenerations
ago- makingher, on
cladisticprinciples
, a memberof oneof two newspecies
(LaPorte1997
).
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, H. (1975). The meaningof "meaning
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, E. (1980, reprint1994). Evolution
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pointof view. Cambridge
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. In B. K. Hall, ed., Homology
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, naturalkindsand the evolutionof modularity
Wagner
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, G. P., G. Booth
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, B. C. (1997). A populationgenetictheoryofcanaliza
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, G., andB. C. Goodwin(1996). Formandtransformation
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andrelational
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.
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Wimsatt, W. C. (1976a
). Complexityand organisation
. In M. GreneandE. Mendelsohn
, eds.,
Boston
studies
in philosophy
, vol. 27: Topicsin philosophy
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of biology
: Reidel.
Wimsatt,W. C. (1976b
). Reductionism
, levelsof organisation
andthemindjbodyproblem
. In G.
Globus
, G. Maxwell, andI. Savodnik
. eds., Consciousness
andthebrain.NewYork: PlenumPress
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Wimsatt, W. C. (1986). Developmental
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MartinusNijhoff.
228
1lI. RethinkingNaturalKinds
IV
' ~les
pe
232
/
in Folkbiological
Taxonomy
Atran: TheUniversalPrimacyof GenericSpecies
three and six ranks. Taxa of the samerank are mutually exclusiveand tend to
display similar linguistic, biological, and psychologicalcharacteristics
.
Ranksand taxa, whether in folkbiological or scientific classification, are of
different logical orders, and confounding them is a category mistake.
Biological
ranks are second-order classesof groups (e.g ., species
, family, kingdom)
whose elements are first-order groups (e.g., lion, feline, animal). Folkbiological ranks seemto vary little , if at all, acrosscultures as a function of
theoriesor belief systems; in other words, suchranks- but not the taxa they
contain- are universal. Ranks are intended to representfundamentally different
levels of reality, not convenience}
The most general folkbiological rank is the folk kingdom- for
example,
plant and animal. Such taxa are not always explicitly named and represent
the most fundamentaldivisions of the biological world . Thesedivisions correspond
to the notion of " ontological category" in philosophy (Donnellan
1971) and in psychology (Keil 1979). From an early age, it appears, humans
cannot help but conceiveof any object they seein the world as either
being
or not being an animal, and there is evidencefor an early distinction between
plants and nonliving things (Hickling and Gelman 1995, Inagaki and Hatano
forthcoming). Conceiving of an object as a plant or animal seemsto carry
with it certain assumptionsthat are not applied to objects thought of as
belonging to other ontological categories, suchas the categoriessubstance
or
artifact (Keil 1989, Mandler and McDonough 1996, Hatano and Inagaki
1998, seealso Keil and Richardson, chapter 10 in this volume).
The next rank down is life fonn. Most taxa of lesserrank fall under one or
another life form. Life-form taxa often have lexically unanalyzablenames
(simple primary lexemes), such as tree and bird, although some life-form
namesare analyzable, suchas quadruped
. Biologically, membersof a life-form
taxon are diverse. Psychologically, members of a life-form taxon share a
small number of perceptual diagnostics: stem aspect, skin covering, and so
forth (Brown 1984). Life-form taxa may representadaptationsto broad sets
of ecological conditions, such as competition among single-stem plants for
sunlight and tetrapod adaptation to life in the air (Hunn 1982, Atran 1985).
Classifying by life form may occur early on: two -year-old children distinguish
familiar kinds of quadruped(e.g ., dog and horse) from seaanimalsand
both of those from air animals(Mandler, Bauer, and McDonough 1991).
The core of any folk taxonomy is the generic-specieslevel. Like life-form
taxa, generic speciesare often named by simple lexemes, such as oak and
robin. Sometimes,generic speciesare labeledas binomial compounds, suchas
, they may be optionally labeled as binomial
hummingbird. On other occasions
such
as
oak
tree
.
In
both cases
,
, the binomial makesthe hiercomposites
archicalrelation apparentbetween generic speciesand life form.
Generic speciesoften correspond to scientific genera (e.g ., oak) or species
(e.g., dog), at least for the most phenomenally salient organisms, such as
larger vertebrates and flowering plants. On occasion, generic speciescan
correspond to local fragments of biological families (e.g ., vulture), orders
234
/
Speciesin Mind andCulture
(e.g., bat) and, especially with invertebrates, even higher order biological
taxa (Atran 1987, Berlin 1992). Generic speciesmay also be the categories
most easily recognized, most commonly named, and most easily learnedby
children in small-scale societies(Stross 1973). Indeed, ethnobiologists who
otherwise differ in their views of folk taxonomy tend to agreethat one level
best captures discontinuities in nature and provides the fundamental constituents
in all systems of folkbiological categorization, reasoning
, and use
.
1993
Ellen
Hunn
1982
1974
Bulmer
)
,
,
(
'
The term genericspeciesis used here, rather than folk generalfolk generic or
me, for four reasons:
specie
folk specieslfolk
1. Empirically, ethnobiologists and historians of systematics (as well as
"
tolerably well
working biologists) mostly agree that speciescome to be
" Darwin 1883
time
,
at
and
(
defined objects . . . in anyone region
anyone
heart
the
are
137) and that such local speciesdefined by ordinary people
of any natural system of biological classification. Whereas zoologists and
ethnozoologists generally refer to such common groups as species(or spe
ciemes
) in focusing on reproductive and geographical isolation that is more
readily identified in terms of behavior (Mayr 1969, Bulmer 1970, Diamond
and Bishop 1998), botanists and ethnobotanistsrefer to them as genera(or
) in focusing on easeof morphological recognition without technical
generics
aids (Greene 1909/ 1983, Bartlett 1940, Berlin 1992). As working concepts,
either term alone is likely to be more confusing for historians of systematics
than the term genericspecies(see Stevens 1994), as when the zoologist
George Gaylord Simpson declaredthat "the hallmarks of priority attributed
by some of his colleaguesto the genus are characteristicof the . .. species,
"
not genus (Simpson 1961, 189).
2. Perceptually, a principled distinction between biological genus and species
is not pertinent to most people around the world . For humans, the most
vertebrates,
phenomenally salient species(including most speciesof large
and
cacti
as
such
) belong to
isolated
and
trees,
palms
evolutionarily
groups
related
.
locale
Closely
speciesof a polytypic
monospecificgenera in any given
no
and
readily perceptible
locally,
genusare often difficult to" distinguish
" canbe discernedbetween them (Diver
or
morphological ecological gap
1940).3
3. Historically, the distinction between genus and speciesdid not appear
until the influx of newly discovered speciesfrom around the world coma worldwide
pelled Europeannaturalists to sort and rememberthem within
.
system of generabuilt around mainly Europeanspeciestypes (Atran 1987) .
in terms of initially mono
Ti:te original genus concept was partially justified
other
which
to
speciesaround the world
typic generic European species
~ ght be attached(T oumefort 1694)
4. Onto logically, the term genericspeciesreflects a dual character. As salient
mnemonic groups, generic speciesare akin to genera in being those groups
"235
in Folkbiological
Taxonomy
Atran: TheUniversalPrimacyof GenericSpecies
236
/
IV. Spedesin Mind andCulture
237
/
Atran: The Universal Primacy of GenericSpeciesin Folkbiological Taxonomy
The Itzaj are Maya living in the Peten rainforest region of Guatemala. Until
recently, men devoted their time to shifting agriculture, hunting, and silvi culture, whereas women concentrated on the myriad tasks of household
maintenance
. The Itzaj were the last independent native polity to be conquered
.
the
by
Spaniards(in 1697), and they have preservedvirtually all ethnobiological knowledge recorded for lowland Maya since the time of the
initial Spanish Conquest (Atran 1993, Atran and Ucan Ek' forthcoming).
Despite the current awesomerate of deforestation and the decline of Itzaj
culture, the languageand ethic of traditional Maya silviculture is still very
much in evidenceamong the generationof our informants, who range in age
from fifty to eighty years old. The midwesternersin the United Stateswere
self-identified as people raised in Michigan and recruited through an advertisement
in a local newspaper.
Basedon extensive fieldwork with the Itzaj, we chose a set of Itzaj folk biological categoriesof the kingdom (K), life-form (L), generic-species(G),
folkspecmc(S), and folk -varietal ( V) ranks. We selectedthree plant life forms:
che' = tree, ak' = vine, pok,..."che' = herb; bush. We also selectedthree animal
life forms: b'a'al,..."che' kuxi'mal = " walking animal" (i.e., mammal), ch'iich' =
birds, including bats, kay = fish. Three generic-species taxa were chosen
from each life form so that eachgeneric specieshad a subordinatefolk specific
, and eachfolk specifichad a salient varietal.
Pretesting showed that participants were willing to make inferencesabout
. The propertieschosenfor animalswere diseasesrelated
hypothetical diseases
'
to the heart (pusikal), blood (k 'ik'el), and liver (tamen). For plants, there were
diseasesrelated to the roots (motz), sap (itz), and leaf (Ie'). Properties were
chosen according to Itzaj beliefs about the essential, underlying aspectsof
'
life s functioning. Thus, the Itzaj word pusik'ai, in addition to identifying
the biological organ heart in animals, also denotes essenceor heart in both
animals and plants. .The term motz denotes roots, which are consideredthe
initial locus of the plant pusik'al. The term k'ik 'el denotes blood and is conceived
as the principal vehicle for conveying life from the pusik'al throughout
the body. The term itz denotes sap, which functions as the plant's k'ik 'el.
238
/
IV. Speciesin Mind and Culture
'
'
'
"
"
The tamenor liver helps to center and regulate the animals pusikal. The Ie
'
or leaf is the final locus of the plant pusikal. Properties used for inferences
"
" is
have the form,
susceptibleto a diseaseof the ( root ) called ( X ) . For
" "
eachquestion, X was replacedwith a phonologically appropriatenonsense
'
name(e.g., eta) in order to minimize the task s repetitiveness.
All participants respondedto a list of more than fifty questionsin which
they were told that all membersof a category had a property (the premise)
" "
"
" " "
and were askedwhether all, few, or no membersof a higher-level category
(the conclusion category) also possessedthat property . The premise
category was at one of four levels, either life form (e.g ., L = bird ), generic
species(e.g., G = vulture), folk specific (e.g ., 5 = black vulture), or varietal
(e.g ., V = red-headed black vulture). The conclusion category was drawn
from a higher-level category, either kingdom (e.g., K = animal), life form (L),
generic species(G), or folk specificS ). Thus, there were ten possible combinations
of premise and conclusion category levels: L - + K , G - + K , G - + L,
5 - + K, 5 - + L, 5 - + G, V - + K, V - + L, V - + G, and V - + 5. For example, a
"
folk specificto life form (5 - + L) question might be, If all black vultures are
?" If
called eta, are all other birds susceptible
susceptibleto the blood "disease
"
"
a participant answered no, then the follow -up question would be, Are
'
someor a few other birds susceptibleto diseaseeta, or no other birds at allf
The corresponding life forms for the midwesternerswere: mammal, bird,
fish, tree, bush, and flower (on flower as considereda life form in the United
States, seeDougherty 1979). The properties used in questionsfor the Michigan
"
"
" "
"have
protein X, have enzymeY , and are susceptible
participants were
"
to disease2. Theseproperties were chosento be internal, biologically
basedproperties intrinsic to the kind in question, but abstractenough so that
rather than answering what amounted to factual questions participants
would be likely to make inductive inferencesbasedon taxonomic category
membership.
ExperimentResults
Responseswere scored in
Representativefindings are given in figure 9.1.
"
"
all
or virtually all responses
of
the
we
totaled
.
First
two ways
,
proportion
for eachkind of question (e.g ., the proportion of times respondentsagreed
that if red oaks had a property, all or virtually all oaks would have the same
"
"
eachitem, counting a
responsescores for
property). Second, we calculated
" as 2 and " none or virtually
"
"
"
few
or
3
some
all
as
,
all
or
of
,
virtually
response "
in the strength of
confidence
more
reflected
score
none as 1. A higher
.
an i J1ference
summarizesthe results from all Itzaj informants for all life
9.la
Figure
" "
diseases
and
, and shows the proportion of all responses(black),
fol : ms
"
"
"
"
few responses(checkered
), and none responses(white). For example,
given a premiseof folk specificS) rank (e.g ., red squirrel) and a conclusioncategory
of generic-species(G) rank (e.g., squirrel), 49% of responsesindicated
239
/
Atran: The Universal Primacy of GenericSpeciesin Folkbiological Taxonomy
00
eo
60
40
2
8
.
o
OS
LEVEL
of
PREMISE
CATEGORY
~
E FORM
~
responses :
none- 0
-
few - .
-
811- .
GENERIC
SPECIES
OLK
C ~
JECIFIC
VARIETAL
00
80
60
40
2
tt
IC
FOLK
ENERIC
~NGDOM
~ liFE
~ -FORM
GSPECIES
LEVEL
of
~
CONCLUSION
CATEGORY
A
that Ilalll' squirrels and not just IIsomellor IInonell would possessa property
that red squirrels have. Resultswere obtained by totaling the proportion of
lIall or virtually allil responsesfor eachkind of question (e.g ., the proportion
.
of times respondentsagreed that if red oaks had a property, all or virtually
all oaks would hav~ the sameproperty). A higher score representedmore
confidencein the strength of the inductive inference. Figure 9.lb summarizes
the results of Michigan participant responsescoresfor all life forms and bio.
logic~ properties.
1.40
IV. Species
in Mind andCulture
LEVELof
PREMISE
CATEGORY
J.,
E- FORM
~
responses :
none 0
1
0
01
5
801
601
40
201
3
01
.
9
001
801
6
.
601
40
201
o
~
,
00
80
n
60
40
20
0LIINGDO
LK
FOLK
ENE
EC
II
FE
F
ORM
sP
rx
: ~G
SPE
-
few .
-
a11 .
GENERIC
SPECIES
OLK
C ~
JECIFIC
VARIETAL
LEVELof
-+
CONCLUSION
CATEGORY
B
)
Pigure9.1 (continued
1.41
/
in Folkbiological Taxonomy
Atran: TheUniversalPrimacyof GenericSpecies
4Uf
..~..
in
-(V
..
.-m
an8mspo
".V
I-or
.,
4Itt8ll~
mm
. - J-UPODStS
(n . -an or 8f. ~
1,->0
G
-) 0 -~
1-) o -~:].~
--:I -Q--o
v-)-D
)E~
)1, ->0 ->I
~-) 0
4jffeJllal In co!!!.~
I SCOJI
I UJOm
(" ' ~ ,
-r......z, ~ . . . 1)
1-) 0
-) o---~
G
G
-) o--'t\'"
ITZAJ:
a11
1lfe - forms
I) 0 ~
8-) 0 ~
combined
v-)-0 -0- -0 -0 v-)-0 -0- -0 --0
)E )1, )G >I
)E )1 >0 ->I
1,-) 9
,-)
-)
G
-)
G
1-)
I-) I
I
-)-0- -0--0
v-)-~
v
0)K-)1, -)G-0
)1 )E )1. -)G-0>I
1.-)
G
-)
MICHIGAN
:
1-) . .
a111ife- forms
v-)-L)K
}-.Q
-)1,.-o
-)G-0
)8 combined
:0
KE
.
[ad
}p
-O
O
G
2
>
.
01
nw
-;voo
a
nI
t.fo
M
_.~
betweenadjacentcategories
in the rankandinferencetask
Figure 9.2 Significantcomparisons
for Itzaj Maya and MichiganstudentsResultsincludeall life formsand biologicalproperties
,
" "
"
"
" "
"
" " "
, andcombinedresponses
showing all (versus few" or none), none (versus all or fe W1
" "
"
"
"
inferences
Horna givenrank(premise
( all = 3, few" = 2, none = 1). Maindiagonals
represent
to
the
)
adjacenthigher-orderrank (conclusioncategory
): V(arietal) - S(pedfic),
category
S(pedfic) - G(enericspecies
), G(enericspecies
) - L(ife form), L(ife form) - K(ingdom). Moving
to holdingthe premiseconstantand varying the
horizontallywithin eachgraphcorresponds
conclusion
: e.g., V - S, V - G, V - L, V - K.
" "
, the overall Itzaj and Michigan patterns
strong. Notice that for all responses
are nearly identical.
Moving horizontally within eachgraph in figures 9.1 and 9.2 corresponds
to holding the premise category constant and varying the level of the conclusion
.4 Here, we firid the samepattern for " all" responsesfor both Itzaj and
Michigan participants as we did along the main diagonal. However, in the
combinedresponsescores(" all" + " few" ), there is now evidenceof increased
inductive strength for higher-order taxa among the Michigan participants
versus the Itzaj. On this analysis, both groups show the largest break between
inferencesto generic speciesversuslife forms, but only the Michigan
subjectsalso show a consistentpattern of rating inferencesto life-form taxa
higher than to taxa.at the level of the folk kingdom: G - + K versus G - + L,
5 - + K versus5 - + L, and V - + K versus V - + L.
Finally, moving both horizontally and along the diagonal, for the Itzaj
there. is some hint of a difference between inductions using conclusionsat
242
IV. S~
.
j
'
.
~
;
~
b
C
u
0
>
.
"
t
I
C
c
c
0
O
dU
d
U
d
s
from inferencepatterns for the Itzaj tree life form. There is evidencethat the
Itzaj confer some preferential status upon trees at the folkspeciflc level (e.g.,
savannanance tree). Itzaj are forest-dwelling Maya with a long tradition of
agroforestry that antedatesthe SpanishConquest(Atran 1993).
Experiment Discussion
243
colleagues). Yet like the Maya, they prefer generic species. On the other
hand, objective reality- that is, the actual distribution of biological species
within groups of evolutionarily related species- doesnot substantiallydiffer
in the natural environments of midwesternersand the Itzaj. Unlike the Itzaj,
however, midwesterners perceptually discriminate life forms more readily
than generic species. True, there are more locally recognized speciesof tree
in the Maya area of Peten, Guatemala, than in the Midwest United States.
Still, the readily perceptible evolutionary " gaps" between species are
roughly the samein the two environments (most tree genera in both environments
are monospecific). If anything, one might expect that having fewer
treesin the U.S. environment allows eachspeciesto stand out more from the
rest (Hunn 1976). For birds, the relative distribution of evolutionarily related
speciesalso seemsto be broadly comparableacrosstemperateand rainforest
environments(Boster 1988).
An inadequacyin current accountsof preferred taxonomic levels may be a
failure to distinguish domain-general mechanismsfor best clustering stimuli
&om domain-specific mechanismsfor best determining loci of biological
.
information. To explain Rosch's data, it may be enough to rely on domain. Suchmechanismsmay generatea basic
general, similarity-basedmechanisms
level in any number of cognitive domains, but not the preferred level of
folkbiology . Consider:
In striking contrast to the rich debate over the descriptive adequacyof
accountsof folkbiological taxonomy, little attempt hasbeenmadeto provide
an explanatory account of the psychological mechanismsand processes that
'
actually producefolkbiological groups. A notable exception is Hunn s (1976)
"
"
perceptualmodel, arguably the most influential proposal in ethnobiology
'
(Berlin 1978). This model accordswith Roschs (1973, 1975) generalaccount
of the cognitive structure of perceptual and semantic categories in hierarchical structures. These accounts are variants of what psychologists call
"
"
similarity-basedmodels (Smith and Medin 1981), which organize perceptually
identifiable categories on the basis of correlation or covariation of
stimulus attributes. With such models, one learns to recognize a particular
instance of a category by being exposed to multiple instancesof the category
"
, which implies that, as Boster (1991) puts it , the source of biological
similarity judgments is in the world , not in the brain."
To illustrate the story &om a similarity-basedpoint of view: becausethe
attributes of having a bark, large canines, and a terrestrial habitat usually
co-occur only when a dog is present, then their co-occurrencewill probably
figure in all and only those feature sets generally associatedwith the category
"
"
dog. The mind will automatically tend to cluster perceptible features
.
"
"
into gestalts of maximally covariant attributes, or basic-level categories,
becauseof the " objective" discontinuities that exist in nature. Notice that for
the model to work, it is not imperative that any particular feature always be
necessaryfor defining category membershipor that a given set of features
always be sufficient. All that is required is that the exemplars exhibit a
244
"
"
readily apparent family resemblance among a community of attributes
(Roschand Mervis 1975, Hunn 1982).
Becausethe processing mechanismis a general-purpose device that can
pick out perceptualstimuli &om whatever source, it should operateacrossany
cognitive domain that involves separatedclusters of perceptual attributes,
including categoriesthat occur naturally in everyday biological and social
contexts, as well as constructedcategories(e.g. artifacts). Later researchhas
tended to confirm the findings of Roschand her colleaguesfurther showing
that the basic level extends to artificial and natural categoriesas the level
that people most readily recognize and that children most easily name and
learn (Lassaline, Wisniewski, and Medin 1992).
The sameattribute-clustering strategy can be applied recursively at higher
and lower levels (Hunn 1976). Thus, the simultaneouspresenceof fur and
live-born offspring might figure in the feature set that distinguishes the category
mammal&om other categoriesof superordinate-level life forms, suchas
bird, fish, and so forth. Similarly, a high body length to body height ratio,
when added to tn'efeature gestalt for dog, might figure in the feature set that
distinguishes the subordinate":level category dachsund&om other types of
dog. The basic level, then, is that level without which relatively much information
is lost and below which little information is gained. That is, there is
a large gain in information when going &om the superordinateor life-form
level to the basic level and there is only a slight gain in information going
&om the basiclevel to the subordinateor specificlevel.
Thus, both anthropology and psychology suggest that privilege or
"basicness" could be a function of correlated featuresor
properties producing
natural clusters that are psychologically salient. These salient chunks
should organizeboth category organization and reasoninginvolving categories
(Anderson 1990). Compelling as this view is, however, it is inadequate
to describeour findings. The challengeis to explain why the generic-species
rank is preferred both by Maya, who have relatively extensive contact with
the natural environment, and by Michigan students, who have relatively little .
"
"
The key problem is that the linguistic and perceptualcriteria for basicness
used by Rosch and her colleaguespoint to the life-form level as preferred,
but as we have just seen, the break point in induction appearsat the more
specificrank of generic species.
To explain our data may require, in addition to domain-generic perceptual
heuristics, domain-specific mechanismsfor the formation of biological
"
categories that are not similarity-based. Along these lines, a living -kind
module" would involve a domain-specificsort of causalreasoning that may
"
"
be called teleo-essentialist (Atran 1995, Keil 1995). The idea is that univers
"ai and possibly innate principles lead people to believe that visible
morphotypical patterns of each readily identifiable generic speciesas well
as.~onobvious aspectsof biological functioning are causally produced by
an ~ derlying essence
. The nature of this essenceis initially unknown, but
.
The
learner
(e.g., a child) then attempts to discover how essences
presumed
/
Atran: The Universal Primacyof GenericSpeciesin Folkbiological
Taxonomy
govern the heritable teleological relations between visible parts , how they
link initially ill -perceived internal parts to morphotypical parts through
canonical patterns of irreversible growth and how they determine the stable
and complex functioning of visible and nonobvious parts . Virtually all people
in all cultures cannot help but follow through this spontaneously triggered
"
"
research program , which compels them to deepen and extend the
domain of information relevant to living kinds within a taxonomic framework
that focuses attention on generic species.
Notice that although a generic species may fail to be "basic " in Rosch' s
sense of a maximally rich cluster of readily available perceptual information ,
it may still be preferred as a maximally rich bundle of anticipated biological
information . In other words , domain -specific constraints on categorization
and category -based reasoning may diverge from domain -general constraints .
When and where they do , the expectation is that domain - specific constraints
are paramount .
In small- scale societies, adults as well as children learn about generic
species just by being told about them or by seeing a single instance. In our
society , one need only describe a single instance in a picture book or point
to an isolated example in a zoo or museum to have an adult or child
instantly extend that poor and fragmentary instance of experience to an
indefinitely extendible category . The taxonomic position of the category is
"
"
immediately fixed as a generic species. This fixture automatically carries
with it a complex internal structure that is partially presumed and partially
inferred , but by no means directly known .
How can people conceive of a given category as a generic species without
primarily relying on perception ? Ancillary encyclopedic knowledge may be
often crucial . Thus , one may have detailed perceptual knowledge of dogs but
not of oaks. Yet a story that indicates where an oak lives or how it looks or
grows or that its life is menaced may be sufficient to trigger the presumption
that oaks comprise a generic species just like dogs do . But such cultural learning
produces the same results under widely divergent conditions of experience
in different social and ecological environments , which indicates that the
learning itself is strongly motivated by cross-culturally shared cognitive
mechanisms that do not depend primarily on experience .
In conjunction with encyclopedic knowledge of what is already known
for the natural world , language is important in targeting preferred kinds by
triggering biological expectations in the absence of actual experience or
knowledge of those kinds (Gelman , Coley , and Gottfried 1994 ). Language
alone , however , would not suffice to induce the expectation that , for people
who live in urban areas in the United States, little or poorly known generic
species are more biologically informative than better known life forms . Some
other process must .invest the generic - species level with inductive potential .
Language alone can only signal that such an expectation is appropriate for a
given lexical item ; it cannot determine the nature of that expectation . Why
"
"
presume that an appropriately tagged item is the locus of a deep causal
246
IV. ~
Summary
Our findings suggest that fundamental categorization and reasoning processesin folkbiology are rooted in domain-specific conceptualpresumptions
and not exclusively in domain-general, similarity-based (e.g., perceptual)
heuristics. People in either subsistenceor industrialized cultures may differ
on the level at which they most easily identify organisms, but still prefer the
sameabsolutelevel of reality for biological reasoning- namely, the genericon
speciesrank. Michigan college students have greater secondaryreliance
life forms becauselife forms are what the studentsmost easily recognizeand
know from experiencein urbanizedenvironments, whereasItzaj Maya have
greater secondary reliance on folk specifics becausetheir silvicultural life
depends on experienceat that level. Despite the compelling needs established
by lived experience, both the u .s. students and the Maya over
wheimingly and in nearly equal measuresubordinate such influences to a
preferencefor generic species. I have argued that they show this preference
beCausethey presumethe biological world to be partitioned at that rank into
, or inherent
nonoverlapping kinds, eachwith its own unique causalessence
not readily
visible
whose
nature
may
mayor
,
products they
underlying
247
GENERIC
SPECIFS
ELECn ON, AND THEEVOLUTION
, NATURALS
OFHUMANCOGNm ON
248
IV. Species
in Mind andCulture
individuate the membersof other speciesin this way, but only as examplars
of the (generic) speciesthat identifies them as causallybelonging to one and
only one essentialkind.
Natural selectionbasically accountsonly for the appearanceof complexly
well-structured biological traits designed to perform important functional
tasks of adaptive benefit to organisms. In general, naturally selectedadaptations
"
'/
are structures functionally perfected for any given habit (Darwin
I'
1883, 140) and that have very much the appearanceof design by an intelligent
designer . . . on which the wellbeing and very existenceof the organism
"
depends ( Wallace1901, 138). Plausibly, the universal appreciation of
generic speciesas the causalfoundation for the taxonomic arrangementof
biodiversity and for taxonomic inference about the distribution of causally
related properties that underlie biodiversity is one such functional evolutionary
adaptation.
TURALEV0 LUTION
~ ANDCUL
SPEC
GENERIC
Folkbiology in general and generic species concepts in particular represent a
stable knowledge structure that is supported by high interinformant agreement
and that regularly and recurrently (within and across cultures ) serves as
a principled basis for the transmission and acquisition of more variable and
extended forms of cultural knowledge . Consider , for example , the spontaneous
- the
correspondence of social groups with
emergence of totemism
- at different times and in different
parts of the world . Why ,
generic species
"
"
as Levi -Strauss (1963 ) aptly noted , are totems so good to think ? In part ,
totemism uses representations of generic species to represent groups of people
; however , this pervasive metarepresentational inclination arguably owes
its recurrence to its ability to ride piggyback on folkbiological taxonomy .
Generic species and groups of generic species are inherently well structured
, attention arresting , memorable , and readily transmissible across minds .
As a result , they readily provide effective pegs on which to attach knowledge
and behavior of less intrinsically well -determined social groups . In this
way , totemic groups can also become memorable , attention arresting , and
transmissible across minds , which are the conditions for any meme to become
culturally viable (see Sperber 1996 for a general view of culture along the
"
"
lines of an epidemiology of representations ). A significant feature of
totemism that enhances both memorability and its capacity to grab attention
is that it violates the general behavior of biological species: members of a
totem , unlike members of a generic species, generally do not interbreed , but
only mate with members of other totems in order to create a system of social
exch' ange. Notice that this violation of core knowledge is far from arbitrary .
'
In fact , it is a pointed violation of human beings intuitive ontology , and as
su~ it readily mobilizes most of the assumptions people ordinarily make
about biology in order to help build societies around the world (Atran and
Sperber 1991).
249
GENERIC
SPECIFS
SENSE
ANDSCIENCE
, COMMON
Much of the history of systematics has involved attempts to adapt to a more
global setting the locally relevant principles of folkbiology - such as the
taxonomic embedding of biodiversity , the primacy of species, and the tel eoessentialist causality that makes sense of taxonomic diversity and the life
functions of species. This process has been far from uniform (e.g ., initial
rejection of plant but not animal life forms ; recurrent but invariably failed
attempts to define essential characters for species and other taxa; intermittent
attempts to reduce teleological processes to mechanics; and so forth ).
Historical continuity should not be confounded with the epistemic continuity
or use of folk knowledge as a learning heuristic for scientific knowledge
. Scientists have made fundamental onto logical shifts away from folk
understanding in the construal of species, taxonomy , and underlying causality
. For example , biological science today rejects fixed taxonomic ranks, the
primary and essential nature of species, teleological causes of species existence
, and phenomenal evidence for the existence of taxa (e.g ., tree cannot be
a scientifically valid superordinate plant group , but bacteria almost assuredly
should be).
Nevertheless , from the vantage of our own evolutionary history , it may
be more important to the everyday life of our species (or at least to the
aspects of everyday life that we became sensitive to as we evolved ) that our
ordinary concepts be adaptive than true . Relative to ordinary human perceptions
and awareness, evolutionary and molecular biology ' s concerns with
vastly extended and minute dimensions of time and space may be of only
marginal value . The onto logical shift required by science may be so counterintuitive
and irrelevant to everyday life as to render inappropriate and maladaptive
the use of scientific knowledge in grasping and responding to
everyday circumstances. This situation makes untenable any uniform application
of the doctrine of externalism for living kind concepts (i .e., the belief
that understanding the ordinary meaning and reference of living -kind terms
necessarily involves commitment or deference to scientific knowledge when
available - in other words , to a likelier nomological account of the world ).
Scientific knowledge cannot wholly subsume or subvert folkbiological
knowledge .
Reliance on folk concepts rather than on scientific concepts may depend
on c.ontext . Belief in essences, for example , may greatly help people explore
250
251
/
Atran: The Universal Primacy of GenericSpeciesin Folkbiological Taxonomy
252
/
IV. Speciesin Mind and Culture
these alternatives , however , severs any connection with the specific com monsense intuitions that universally allow humans beings to believe in the
organic world as a causally related totality in the first place and that were a
necessary (if not sufficient ) condition for the development of biological and
evolutionary thinking ( by contrast , even without notions of representation
or of the ether , human belief in the mind or the world of substance is conceivable
, as are the sciences of cognitive psychology and physics ).
Perhaps the species concept , like teleology , should be allowed to survive
in science more as a regulative principle that enables the mind to establish a
regular communication with the ambient environment than as an epistemic
principle that guides the search for nomological truth . Once communication
is established with the world , science may discover deeper channels or more
significant overlapping networks of causality . The persistence of a species
concept would function to ensure only that these diverse scientific explorations
are never wholly disconnected or lost from one another or from that
aspect of phenomenal reality that will always remain as evident to a Maya as
to a modem scientist .
A SYNOPSIS
OFFOLKBIOLOGY
Folkbiology (FB) may be canonicallydescribedas follows:
1. FB has a cognitive structure that is culturally universal and that places
a priori constraints on the ways human beings ordinarily categorizeand
reason inductively about the properties and relationships of organic
objects. This structure consistsof
1.1. Categorical distinctions and principles in the intuitive ontology of
humanbeings, suchthat
1.1.1. Every natural object is either a living kind or not.
1.1.2. Every living kind is either an animal or a plant; that is, it
belongs to a folk kingdom.
1.1.3. Each animal or plant belongs to one and only one essential
.
grouping, or genericspecies
1.1.4. In addition:
253
254
255
J Taxonomy
Atran: The' Universal Primacy of GenericSpeciesin Folkbiologica
ACKNOWLEDGMENTS
The comparativestudiesreported here were funded by the National Science
Foundation (SBR 93- 19798, 97-07761) and the FrenchMinistry of Research
and Education (Contract CNRS 92-C-O758). They were codirected with
Douglas Medin. Participants in this project on biological understanding
across cultures include Alejandro Lopez (psychology, Max Planck), John
Coley (psychology, Northeastern University
. ), ElizabethLynch (psychology,
Northwestern University), Ximena Lois (linguistics, Crea-Ecole Polytechnique
), Valentina Vapnarsky (anthropology, Universite de Paris X) , Edward
Smith and Paul Estin (psychology, University of Michigan), and Brian Smith
( biology, University of Texas, Arlington ).
NOTFS
1. Thus, comparing constellationsin the cosmologiesof ancient China. Greece, and the Aztec
Empire shows little commonality. By contrast, herbalslike the ancient Otinese ERH YA, Theo'
phrastuss PeriPutonIstorias, and the Aztec BadianusCoda, shareimportant features, suchas the
classmcationof generic speciesinto tree and herb life forms (Atran 1990, 276).
2. Generalizationsacross.taxa of the samerank thus differ in logical type from generalizations
that apply to this or that taxon. Tentaite
, pig, and lemonfreeare not related to one another by a
simple classinclusion under a common hierarchicalnode, but by dint of their common rankin this case, the level of generic species. A system of rank is not simply a hierarchy, as some
suggest ( Rosch1975, Premack1995, Carey 1996). Hierarchy- that is, a structure of inclusive
- is common to many cognitive domains,
classes
including the domain of artifacts. For example,
chair often falls under furniture but not vehicle
, and car falls under vehiclebut not furniture. But
there is no ranked systemof artifacts: no inferential link or inductive framework spansboth chair
and car or fu~ iture and vehicleby dint of a common rank, such as the artifact species
or the artifact
family.
3. For example, in a comparativestudy of Itzaj Maya and rural Michigan college students, we
found that the great majority of mammal taxa in both cultures correspond to scientific species
and that most also correspondto monospecificgenera: 30 of 40 (75%) basic Mid1igan mammal
256
IV. ~
CFS
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Atran: The Universal Primacyof GenericSpeciesin Folkbiological
Taxonomy
1 0
artifacts and that this distinction could explain the results of severalpriming
studiesthat revealedlatency differencesbetween the two kinds. Furthermore,
they claimedthat an attrador network that had distributed knowledge of the
empirically derived feature norms could simulate the samesort of behavior
in priming tasks. Of course, priming results alone are hardly enough reason
to supposethat the correspondingliving kind conceptswere fundamentally
different in structure, but the work done by McRae and his colleaguesillustrates
how suchan argumentmight proceed.!
Alternatively , it might be exceedingly difficult to specify any objective
informational patterns that set aside plants and animals as a distinct kind
with its own rich internal structure; instead, humans may carry with them
certain cognitive blasesto interpret information about speciesin highly distinctive
and unique ways. Thus, extremely simple perceptualfeaturesshared
by animals and plants (perhaps a kind of &actal structure seen more commonly
in living things) might trigger a cascadeof predeterminedcognitive
blasesthat make learning and the resulting conceptsof biological kinds radically
different &om oth~r conceptsof kinds. There may be some real informational
differencesas well, suchas differing densitiesof featureclusters, but
this view would argue that those differencesfar underdeterminethe nature
and kind of specializationsseenin biological thought.
We argue that there may be misleading assumptions and misconceptions
about the nature of the distinctive information and about the blases
that color our notions of what biological thought is like; also, the ensuing
misconceptionsabout biological thought, especially as a kind of intuitive
theory, may obscure the true nature of speciesconceptsand how they are
embeddedwithin the broader system of biological knowledge. Severalnew
lines of empiricalwork are suggested.
Much of our discussionexamineshow biological thought in general and
speciesconcepts in particular develop in the child. We take such a developmenta
perspectivebecause(1) it tells us what sorts of information might
be most salient to the naive mind, and (2) it suggestswhat might be the
most fundamentalblasesthat we all have in thought about the living world.
Between the empiricist and nativist extremes there are many gradations
and combinations. Our goal here is not to allocate responsibility to these
two extremesas much as it is to explore what eachmay contribute to a more
complex interactive model of where speciesconceptscome &om and what
makesthem special. We do so by trying to clarify what informational patterns
might be both distinctive to living kinds and salient to humans, and by
asking what sorts of cognitive blasesmight interact with that information,
even if those blasesthemselvesare not always reservedexclusively for biological
phenomena.
THE GENERAL NATURE OF FOLKBIOLOGICAL mOUGHT
"
Any notions of speciesare embeddedwithin broader systemsof belief about
biological kinds. One cannot understandwhat a dog is without also knowing
264
something about animal life cycles, nutritional needs, and the like. For.that
reason, it is helpful to consider how views of intuitive biological thought
have changed in recent years (e.g., Medin and Atran 1999). It has become
commonplaceto argue that lay people throughout the world possessrichly
structured beliefs about the living world that might be thought of as intuitive
or naive theories about biological processes and systems- such as
, and death ( Medinand Atran 1999).
growth , reproduction, digestion, disease
beliefs
as
theories
or mental models, however,
Characterizationsof these
may carry with them somewhatmisleadingideasabout how that knowledge
is representedin the mind of the individual. It might seemthat the knowledge
must be an explicit set of beliefs connectedtogether in the tightly coherent
manner of a formal scientific theory and that the models must contain concrete
, imagelike components whose interactions can be clearly visualized.
Folk theories would then be said to differ from formal scientific ones only in
terms of the particular setsof beliefs they embraceand not so much in terms
of their generalformat. Thus, a belief that demonic possessioncausesdisease
contagion might have very much the samekind of mechanisticset of lawful
relations as a belief that germs causediseasecontagion. (Keil et al. 1999).
But a closer look at intuitive biological theories and perhapsat many theories
in the more formal sciencesreveals something quite different from an
explicit set of propositions all linked together in a tightly connected, logically
consistent, and coherent set of inferences. Most people have strikingly
little knowledge of the detailed mechanismsat work in their own bodies, let
alone in other animals and plants. An exceedingly simple gloss may be all
that is known, such as food contains energy and that the body uses that
energy as a kind of fuel to power muscles, which makeus move. This simple
functional schematamay then occasionallyget filled with local mechanisms
and gradually becomeinterconnectedin somewhatlarger and more coherent
structures, but only the smallestpercentageof people in any culture can tell
you much of anything about the full causalchain that goes from the ingestion
of food to the production of a motor movement basedon the energy in
that food.
. They
Yet people seemto have far more than a set of functional schemata
classes
of
of
abstract
and
often
seemto have general
ways choosing among
explanations about biological phenomena, even though these explanations
may be equally satisfactoryfrom a functional point of view. An explanation
of mechanismsof digestion in "terms of a mechanismin which food particles
are converted to light that is routed around the body before being transformed
into muscle energy would be vastly less plausible to most adults
than a mechanismthat invoked transformation of food into a kind of fluid
" fuel " even if both mechanismswere
,
ultimately wrong .
above
the level of specificmechanisms
of
sorts
,
are
several
There
things,
kinds
about
and
believe
that adults and possibly children know
biological
and. that nonethelessmight be distinctive to those kinds and thereby make
biological thought different:
265
/
K~il and Richardson: Species
of Causation
, Stuff, andPatterns
266
/
IV. Speciesin Mind and Culture
267
/
: Species
Keil and RiChardson
, Stuff, and Patternsof
268
IV. Species
in Mind andCulture
269
/:
Keil and RiChardson
: Species
, Stuff, and Patternsof Causation
270
/
Speciesin Mind andCulture
THEFSSEN
CEOFFSSEN
CE
All of us may succumbto essentialistblasesthat compel us to assumeand
look for those critical aspectsof kinds that allow us to make powerful inferences
. Gelman and Hirschfeld (1999) contrast three types of " psychological
" : sortal ideal and causal. Sorial essence
essentialism
refers to critical defining
,
,
featuresor, in other words, singly necessaryand jointly sufficient setsof features
for determining category membership- an accountthat seemsto work
for only a small set of real-world concepts, however. Ideal essence
refers to
nonexistent perfect cases. The ideal essenceof parallel lines has no real
counterpartsbecauseno physical system can perfectly embody parallelism.
refers to something about a kind that results in its
Finally, causalessence
of
its
most
having many
typical and stable properties. The nature of that
"
" is critical. Most
commonly, it seemsto be thought of as a fixed
something
inner entity that has multiple causaleffects. That entity might be a kind of
substance(the essenceof gold being atoms with gold' s number of protons)
or an informational code (the DNA sequences] correspondingto tigers) or a
process.
Despite much talk about essencein recent years, it still is not clear how
most lay people actually conceive of essencesor whether the blases are
much stronger in younger children and in thought about somesorts of kinds
rather than about others. Medin and Ortony (1988) have suggestedthat we
can often believe in an essencewithout any idea of what the essenceactually
"
is- that we have an essence place holder" concept without the concept of
the essenceitself. This senseof essencemakesclearerhow it could be a bias
without specificcontent. But the place holder notion may overlook a related
set of beliefs, for even as people commonly believe in essenceswithout
, they still might prefer some sorts
knowing any details about those essences
of future details to others. Thus, there might be a physicalist bias, whereby
"
"
people prefer essencesto be seenas objects or stuff rather than kinds of
processes. Teaching that photosynthesisis the essenceof green plants might
be less compelling than teaching that it is a certain DNA sequence
, even
though the processof photosynthesismay be much more directly connected
causallyto a far greater range of phenomenalproperties of plants.1For speciesconcepts, then, notions of essencemay be absolutely essential,
yet we have little idea of what sorts of constraints there might be on
. Sometheorists have argued that notions of essencehave
notions of essence
had extraordinary limiting effectson how we think about speciesin the context
of evolutionary thought. Hull (1965), for example, talks about how
"
"
Ari .stotelianessentialismcauseda 2,000 year stasisin evolutionary thought
becauseit assumedthat specieshad fixed essencesand thereby could not
explain how new speciescould evolve through natural selection. The notion
of speciesas a probabilistic concept, a distribution of types, seemedto be
foreclosedby the essentialistbias even though that notion of distribution is
271
/
Keil andRichard
son: Species
, Stuff, andPatternsof Causation
272
/
IV. Speciesin Mind and Culture
273
Keil and .
dson: Species
, Stuff, and Patternsof Causation
274
275
SPECIES
CONCEPTS
AS A DISTINCTKIND OF CATEGORIZADON
A speciesconceptis a kind of categorization
. It treatsa classof living things
asequivalentin importantrespects
, andthat equivalence
thenlicensespowerful
inductions
, whichis presumably
why species
conceptsaresouseful.But
inductivepower is said to be a key motivationfor almostall
cognitively
naturalcategories
; what else, beyondessence
, is distinctiveabout species
? Thereappearto be manyqualitative
conceptsasopposedto otherconcepts
distinctionsbetweenliving kinds and artifacts
, includinghow taxonomies
,
teleology, and exemplarsare construed
, yet the questionremainswhether
or not thesedifferences
aredue to quantitativedifferences
in the spreadof
causalhomeostatic
.
patterns
Living kinds are saidto be muchmore deeplyembeddedin taxonomies
thanareothersortsof kinds, andthroughoutthe world, all
peoplesseemto
realizethis taxonomiccharacter
on
in
very early
(Atran, chapter
development
9 in this volumeand 1998). Abundantevidencenow showsthat this taxonomicassumption
is very powerfulfor living kindsandprobablynot nearly
aspowerfulfor othersortsof kinds(althoughthe latterhavenot been
nearly
ascarefullyor systematically
studied). However, whatis it aboutliving kinds
that leadsto this taxonomicassumption
? As with essences
, it is not so clear
what "triggers" the assumption
. Thus, it is not so objectivelyobviousthat
living kindsoccurin moredeeplyembedded
hierarchies
. Thereare, afterall,
quite deephierarchiesfor many naturallyoccurringcompounds(kinds of
rocks, soils, gems, etc.). For example
, the UnitedStatesDepartmentof Agriculture
a
four
basic
gives
layered
taxonomyof soil types- categorizing
, to
takeone instance
from
histolsolsto fibriststo sphagnofibrists
,
that areper, depth of hierarchyis not evidentlya cue. Similarly, there
gelic. Therefore
arevery rich hierarchies
for manyclasses
of artifacts.The U.S. PatentOffice
has more than five hundred classesof artifacts
: meanderingdown the
" has
scheme
, of thesefive hundred
, the class"Surgery
forty divisions,within
" has
which "Prosthetics
within which "Leg" hasnine, and of these
,
twenty
"Socket
" hassix
. Evenif the hierarchies
of living kindsaregenerally
categories
, it is not at all clearthat their depth in itself is responsiblefor
deeper
their being seentaxonomically
. Thus, thereis no evidenceto supportthe
intuition that deeperhierarchies
aremoretaxonomicallycompelling
. If depth
doesn't predicta senseof taxonomyfor the nonlivingworld, why shouldit
be a factorin theliving world?
-relatedtaxonomies
Do thesespecies
provide the only apparentways to
kinds
eventhoughtherearemultiplewaysfor other sortsof
classifyliving
kinds? No. Animalscanbe classified
aspredatorsor prey, asdomesticated
or
wild, or asedibleor not, or they canbe classifiedwithin theoriesthat focus
on phylogenetics
, ecology, and so on. It is difficult to know how to count
. the ways, but it is not obviousthat
.
living kindsaredifferentin this respect
.
Evenin folk biology, we seetreesand nontrees
asone way of sortingplants
276
N . Sp~
277
bestexemplarbecomes
moredifficultwhenthe functionis not easilyhamed:
What is the one-line functionaldefinitionfor churchor theinternet
?
How do we think of the exemplarsof living kinds? In somecases
, living
kindshavefunctionalvaluefor humanbeingsand canbe evaluatedon that
dimension
: the besttreesarethosewith the highestfruit yield, for
.
example
" : the
Or we might hamethe bestexemplarasthat whichfulfills its "
purpose
bestbutterfly is camouflaged
Hornpredators
, can extractnectarefficiently
,
lay a largenumberof eggs, and so on. But this hameworkbringsin all the
problemsof teleologydiscussedabove. In addition, how do we evaluate
how "doglike" a dog is? It couldbe the casethat we havesome
prototypeor
idealizedaverageof all dogs. Doesthis averageequatewith a
?
dog essence
Alternatively, we may havesomenotion that goesbeyondthe typicality of
of a homeostaticclusterof causal
perceptualfeaturesto a representation
. It becomesdifficult, however, to
propertiesthat are seento be significant
drawa firm distinctionbetweenliving andnonlivingkindson this criteriaof
well definedness
because
thereis considerable
variationwithin both classes
.
Well definedness
in itself may thereforenot be the cut betweenartifacts
andliving kindsbecause
they both havefuzzyaspects
, but the fuzzlness
may
occurin differentwaysfor the two kinds. Furthermore
, it seemsthat when
morerigid criteriaareappliedin the evaluationof
living kinds, thosekinds
arebeingmeasured
in relationto somefunctionalgoal, eventhe seemingly
hollow purposeof winning a dog show. In short, species
conceptsmay be
importantlydifferentHornother sortsof conceptsthat categorizekinds, but
the dimensions
anddegreesof differencearestill unclear
.
It seems
, therefore
, that speciesconceptsmay be distinctin severalways
that go beyondnotionsof essence
andthat arenot closelyrelatedto patterns
of causalhomeostasis
. The detailsof the psychologicaldifferences
between
thesewaysarejust beginningto be uncovered
, however.
CONCLUSIONS
Speciesconceptsseemindeed to reflect a specialkind of categorization. Several
cognitive blasesmay interact with some very distinctive informational
patterns of living kinds, such as a disposition toward certain types of properties
and relations, as well as a tendency to discretize these homeostatic
patterns into an essenceof some sort. But it is clear that all of thesepsychological
contrasts are just beginning to become apparent. The nature of a
speciesconceptis mostly a placeholder at presentand is &amedby only the
softest and vaguest of constraints. We therefore do not really know much
about how it is that speciesconceptsarise, but this lack of
knowledge should
.
not be discouraging becausefor the first time we are now in a
position to
learn a great deal more about how those concepts
emerge in development
and become used by adults. The rapid growth of work on
biological
thought, especially in cross-cultural and developmental perspectives, has
helpe.d set up a &amework in which it is now possible to pose highly
278
ACKNOWLEDGMENTS
therein
of partsof thispaperandsomeof thestudiesdescribed
Preparation
to
HD23922
ROI
Health
of
grant
weresupported
by anNationalInstitutes
FrankKeil.
NOTES
'
)
1. As an aside
, it is interestingto note the similaritiesbetweenMcRaeandcolleagues(1997
concepts
representing
of anattradorspaceasa way of describinga cognitivemechanism
discussion
' s (1996) claimthattheconceptof spedesshouldbeunderstood
andGoodwinandWebster
" seeGriffiths,
"
Geld
(
chapter
in biologytheoryin termsof an attradorspaceor a morphonogenic
.
8 in thisvolume)
2. Somesort of cognitivebiasmaybe favoringthenotionof DNA in lay thinkingaboutliving
in this volume,
of living kinds. As arguedelsewhere
kindsandappropriatingit as the essence
more
be
can
there
because
variability
DNA is not a sufficienttool with whichto dividespecies
, it appearsasif the scientifictermDNA hasbeen
within speciesthanbetweenthem. However
andDNA divideup
, eventhoughessence
equatedwith someintuitive, folk notion of essence
groundsfor this bias,
the world in very differentways. We discussthe possiblepsychological
the notion of
but for now we might makethe observationthat whenpeoplemisunderstand
an
belie
make
mistakes
underlying
the
do
commonly
few
a
all
but
they
DNA asprobably
belief.
essentialist
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281
: Species
Keil andI ((chardson
, Stuff, andPatternsof Causation
Species Begone !
11
Hierarchy
Marc Ereshefsky
For Linnaeus, the fundamental divide in his hierarchy lies between genera
and all other higher taxa. Classificationsof speciesand genera reflect real
groups in nature, whereas classificationsof classesand orders are artificial
(Cain .1958, 148, 152- 153). This distinction stems from a central tenet of
286
/.
Speaes
Begone!
. According to Unnaeus,
Linnaeus's biological theory: Aristotelian essentialism
the membersof speciesand genera are endowed with essentialnatures created
by God. The job of the biologist is to discover those essencesand their
, on the
associatedspeciesand genera (Ereshefsky1997). Orders and classes
.
artmcial
constructs
and
are
on
other hand, are constructed pragmaticgrounds
"An order is a subdivision of classesneededto avoid
more
placing together
" Unnaeus
in Mayr 1982, 175).
follow
the
mind
can
than
,
(
quoted
genera
mind
have
and
,
Unnaeus
to
,
independentessences
genera
species
According
us.
to
their
existence
owe
and
classes
whereasorders
'
When we dive a bit deeperinto Unnaeuss biological theory, in particular
his sexual system, we see more clearly why he thought that speciesand
. Following Cesalpino,
, have essences
genera, but not orders and classes
Unnaeusbelieved that plants have two vital functions: nutrition , which preserves
the individual, and reproduction, which preserves the kind (Larson
"
1971, 146). The function of reproduction in plants is found in their fructifi "
cation structures - namely, their flowers and fruits. Fructification structures
are found at the level of genera, and are the essencesof genera, according to
'
Linnaeus(Larson 1971, 74 ffj . Moving down a level, a species essenceis a
'
combination of its genuss fructification structure and those parts involved in
the function of nutrition (Larson 1971, 115 ff.). The essencesof speciesand
they are responsible
genera are particularly important for Linnaeusbecause
's
God
of
taxa
for the continued existence
beyond
original creation. Fructification structuresallow the membersof a speciesor a genusto reproduceand
thus allow taxa to continue. Classesand orders, on the other hand, are not
definedby fructification structures. They are merely aggregatesof organisms
.
containing different fructification structures
' s divide between
In the last hundred years, Linnaeus
genera and higher
taxa has fallen on hard times. One reasonis the rejection of essentialismin
Darwinian biology . Unnaeusthought that speciesand genera, but not orders
and classes
; thus, the former are real, but not the latter. In
, had essences
. (See
Darwinian biology , no taxon of any rank has a taxon-specmcessence
Hull 1965 and Sober 1980 for arguments against taxa, especially species,
; d . Wilson, chapter 7 in this volume.) The relevancehere is
having essences
, so
that speciesand generano more have essencesthan do orders and classes
' essentialist
and
lower
between
the
distinction
for
Unnaeuss
holding
ground
higher taxa no longer applies.
Unnaeus also based his distinction on the assumption that only species
and generahave generic-specificfructification structures. Orders and classes
are artmcial groups of genera containing different fructification structures.
Bio}ogists no longer believe that fructmcation structures are responsible
for the existence of taxa. Instead, many hold that taxa are the result of
interbreeding among conspecific organisms (see, for example, Mayr 1970,
3-73- 374, Eldredge and Cracraft 1980, 89- 90). Speciesare populations of
organismsthat exchangegenetic material through interbreeding. That process
causesthe local populations of a speciesto evolve as a unit. In contrast,
287
/
: Speciesand the UnnaeanHierarchy
Ereshefsky
the speciesof a higher taxon do not exchangegenetic material, and the evolution
of a higher taxon is merely the by -product of evolution
occurring
within its species. Thus, instead of Linnaeus's ontological divide between
generaand other higher taxa, the authors of the Modem Synthesis, aswell as
many contemporarybiologists, hold that the ontological divide lies between
speciesand all higher taxa.1
288
/
V. SpeciesBegone!
tions that exchangelittle or no geneticmaterial. Here, the claim is not that the
local populations momentarily fail to exchange genetic material, but that
they fail do so for a significant amount of time. If thesebiologists are right ,
then many speciesof sexual organismsare bound by processes working at
lower levels of biological organization than at the level of the entire speciesd
. Sterelny, chapter 5 in this volume). The unity of such speciesmay be the
result of interbreedingwithin local populations, or their unity may be due to
processes that independently affect organisms, such as selection or genetic
. Either way, somespeciesof sexualorganismsare akin to higher
homeostasis
taxa: they are bound by causalmechanismsacting at lower levels of biological
,
organization. If such sexual speciesexist, or if there are asexualspecies
from
not
then the processof gene flow does
universally distinguish species
higher taxa.
Another argument for the speciesfhighertaxa distinction highlights the
processof speciation. This argument can be found in Eldredge and Cracraft
(1980, 327) and Mayr (1982, 296), and it runs as follows. Speciationis the
primary causeof change in evolution. It occurs in speciesbut not higher
taxa. Therefore, species,but not higher taxa, are the units of evolution. Once
again, speciesare the active agents of evolution, whereas higher taxa are
merely passive results. The trouble with this line of reasoning is that the
locus of speciation is neither the species nor the higher taxon, but the
founder population. According to Mayr' s (1970) allopatric model, speciation
begins when a small population of organisms becomes isolated" and is
. The population undergoes a genetic
exposed to new selection pressures
" and becomesthe
revolution
founding population of a new species.
The point here is that the processof speciation occurs in founding populations
, not in entire species. So if higher taxa are not evolutionary units
becausespeciation occurs in only a portion of them (founder populations),
then by parallel reasoning, speciesare not evolutionary units either.
Now one might counter that it is inaccurate to assert that speciation
occurs in founder populations rather than entire speciesbecausethere is
nothing more to an incipient speciesthan its founder population. Suppose
we grant that point . Then it applies to incipient higher taxa as well: at some
stage in their developmentthey are nothing more than founder populations.
Thus, if speciesare evolutionary units becausespeciation occurs in their
incipient forms, then higher taxa are evolutionary units as well. In brief, the
processof speciationdoes not separatespeciesfrom higher taxa.
Stepping back, we seethat the distinction between speciesand higher taxa
is problematic. Linnaeus drew the distinction along the lines of essential
and fructification systems. Modem biology has rejected essentialism
natures
'
'
and Linnaeuss sexualsystem. Alternatively , the authors of the Modem Synthesis
and many contemporaryauthors basetheir distinction betweenspecies
and higher taxa on the processes of speciationand interbreeding, but those
processes do not adequately distinguish speciesfrom higher taxa, either.
1.89
/
: Species
andthe Unnaean
Ereshefsky
Hierarchy
SPECIES
PLURALISM
Reasonsfor doubting the existenceof the speciescategory come from other
quarters as well. The current taxonomic literature contains no less than a
dozen speciesconcepts(seeEreshefsky1992b and recent issuesof Systematic
Biology). Biologists and philosophershave respondedto this wealth of concepts
in two ways. Monists believe that biologists should settle on a single
correct concept. Pluralists suggest that a number of speciesconceptsshould
be accepted as legitimate (cE. Dupre and Hull, chapters 1 and 2 in this
volume).3
Undoubtedly, a number of currently proposed speciesconcepts will be
found wanting and relegatedto the history of science. However, two major
- the
approaches to species
interbreeding and phylogenetic- are currently
well entrenchedin biology for good theoretical and empirical reasons(de
Queiroz and Donoghue 1988, Ereshefsky 1992a). The interbreeding and
phylogenetic approaches highlight noncomparabletypes of speciestaxa, so
if one acceptsthese two approaches, then there is no single unitary species
category, but a heterogeneouscollection of basetaxa referred to by the term
. Speciespluralism, in other words, poses a threat to the existenceof
species
the speciescategory. The aim of this section is to display that threat and to
highlight the disunity of the speciescategory. But before getting to that, we
need a quick introduction to the interbreedingand phylogenetic approaches.
The interbreedingapproachis typified by Mayr ' s (1970) biological species
concept. Speciesare gene pools held together by interbreeding and protected
. Examples of reproductive
by various reproductive isolating mechanisms
isolating mechanismsinclude courtship behavior that prevents the
mating of two interspecific organismsand hybrid inviability if such mating
does occur. Other speciesconceptsthat fall under the general interbreeding
290
v . Sp/ aesBegone
!
'
approach include Ghiselin s (1974) reproductive competition concept and
'
Patersons (1985) mate recognition concept. Though theseconceptsdiffer in
"
important respects, they agree that a speciesis a field for genetic recombination
" Carson
1957).
(
The phylogeneticapproachis found in the work of cladists (for example,
Cracraft 1983, Mishler and Brandon 1987, Ridley 1989). Propinquity of
descentis the operative notion here. A taxon must contain a single ancestral
speciesas well as all and only its descendantspecies. Any taxon meeting
. The founder of cladism, Willi Hennig, did
that requirement is monophyletic
not intend the notion of monophyly to apply to species
, but reservedit for
higher taxa. Recent cladists have extended its use, however. Mishler and
"
Brandon(1987, 46), for example, define a speciesas the least inclusive taxon
recognized in a classificationinto which organismsare grouped becauseof
"
degreeof monophyly (seealso Mishler, chapter 12 in this volume). Phylogenetic species are base monophyletic taxa maintained by a number of
forces- including selection, interbreeding, genetic homeostasis
, and developmental
canalization.
Given thesetwo approaches to species- the interbreeding and the phylo genetic- one might wonder what feature unifies phylogenetic and interbreeding
speciesinto a single categoryd . de Queiroz, chapter 3 in this
volume). A simple requirement for the existence of a category is that its
entities sharea feature that distinguishes them from entities in other categories
. If phylogenetic and interbreeding speciesare both species, then they
should share some common and distinctive feature. If they lack such a feature
, then the species category consists of noncomparable entities.4 The
remainder of this section examinesthose features that might render interbreeding
and phylogenetic speciescomparable.
A good place to start is with the processes that maintain the existenceof
interbreedingand phylogenetic species.In interbreeding species,gene flow is
the primary unifying force. How much and how often genetic material must
be exchanged varies from interbreeding species to interbreeding species
, the situation is quite
( Templeton 1989, 165). For many phylogenetic species
different. Some phylogenetic species contain sexual organisms living in
isolated populations. In such species, selection, genetic homeostasis
, or
flow.
not
forces
are
the
canalization
,
gene
primary unifying
developmental
Similarly, phylogenetic speciesconsisting of asexualorganismsare bound by
forces other than interbreeding, so different types of speciesare unified by
different types of processes. Consequently, no single unifying process(or set
of processes) servesas the common feature of all speciestaxa.S
P~rhaps we would be better off looking at the structuresof interbreeding
and phylogenetic species.Perhapsa significant similarity can be found there.
So~ e interbreeding speciesconsist of a single local population. Other interbreeding
speciesconsist of a number of local populations connectedby inter
to
have
such
causes
of
the
species
bree4ing. Either way,
process interbreeding
the structure of causallyintegrated entities. Of course, interbreeding among
291
/
: Species
andthe Linnaean
Hierarchy
Ereshefsky
292
v . SpeciesBegone!
IF
IDE
ABC
X
(afterRosen1979,
Figure 11.1 A simplifiedcladogramof the swordBshgroupXiphophorus
276). C + F forms a spedeson the interbreedingapproachbut not on the phylogenetic
.
approach
, whereas C forms
, A + B forms one species
Figure 11. 2 On the interbreeding approach
another species.On the phylogenetic approach
, A, B, and C are three different species.
a species because it doe ~ not contain all the descendants of the common
ancestor X : A + B is missing the organisms of C. So, for many cladists, A
goes extinct at the time of speciation and two new species, B and C , take its
place.
Phylogenetic species often fail to be interbreeding species as well . Some
proponents of the phylogenetic approach believe that asexual organisms
fon. n species (Mishler and Brandon 1987 ). Supporters of the interbreeding
approach disagree and argue that asexual organisms do not belong to any
sP.ecies (Ghiselin 1987 , Hull 1987 ). For my part , I see no reason why the
term speciesshould be reserved for only sexual organisms (see Ereshefsky
1992b and 1998 for arguments ). If asexual organisms do form phylogenetic
species, then many phylogenetic species fail to be interbreeding species. This
293
~
.
Hierarchy
Ereshefsky:Speaesandthe Linnaean
294
/:
V. Species
!
Begone
jar. At this stage in biology , it is not clear what divides speciestaxa from
other taxa. As a result, it is not clear whether a distinctive speciescategory
exists.
Problemsfor the speciescategory come from another angle. The entities
of an existing category should have a theoretically important commonality.
The speciescategory seemsto fail that requirement. The problem is not simply
that speciestaxa vary, but that they lack a common and distinctive biological
feature. Speciestaxa are maintainedby different processes, they have
different ontological structures, and they carve the tree of life in different
ways. This disunity is not causedby a few speciestaxa lacking a biological
feature found in most speciestaxa, but by vast numbers of speciestaxa
forming one typeof speciesbut not another.
We now have two separatelines of reasoning against the existence of
the speciescategory. Together, they provide a strong casefor doubting the
6
reality of the category. Once again, the casehere is merely against the existence
of the speciescategory, not against the reality of those taxa we call
"
"
species. None of the argumentsgiven thus far should causeus to doubt the
.
existenceof such taxa as Homosapiensor Drosophilamelanogaster
The idea that the speciescategory does not exist is far from new. According
to Ghiselin (1969, 93 ff.) and Beatty (1985, 277 ff.), Darwin held this
'
view. In the Origin of Species
, Darwin writes, 1 look at the term speciesas
one arbitrarily given for the sake of convenience to a set of individuals
closely resemblingeachother, and that it does not essentiallydiffer from the
term variety" (1859, 52). In a letter to JosephHooker, he goes even further:
It is really laughable to see what different ideas are prominent in various
"
"
'
naturalists minds, when they speak of species; in some, resemblanceis
everything and descentof little weight in some, resemblanceseemsto go
for nothing, and Creation the reigning idea- in some, sterility an unfailing
test, with others it is not worth a farthing. It all comes, I believe, from trying
to define the indefinable. (December24, 1856; in Darwin 1887, vol . 2, 88)
Of course, Darwin did believe in the existenceof evolving lineages, including
"
"
those called species. He just doubted the reality of the speciescategory
and the other categoriesof the Linnaeanhierarchy.
If the speciescategory does not exist, then how should we treat the term
? Should it be dropped from the discourseof current scienceand relegated
species
? Perhapsin a world of purely rational agents,
to the history of science
this choice would be appropriate. We could, for instance, disambiguatethe
with terms that distinguish the various
languageof biology by replacingspecies
"
'
taxa we call ~species. Grant (1981, 36) suggestscalling interbreeding
"
for phylogenetic
species biospecies." We could add the term phylospecies
. In the abstract, this suggestion is attractive, but there are practical
speCies
is well entrenchedin contempomatters worth considering. The term species
ra:i Ybiology and everyday life, so eliminating it might causemore problems
than simply keeping it . Whether we should or can eliminate the term species
is not an .easy question to answer. I return to it in the last section of the
295
:{ pedesandthe Unnaean
Hierarchy
Ereshefsky
296
v . S~ esBegone
!
in the namesof taxa. The first problem of the Linnaeanrules governing the
'
names of speciesis therefore a semantic one: the designation of a taxon s
rank through the use of binomials may mark no distinction in nature. Similar
observationsapply to the use of suffixes in the namesof higher taxa and the
use of trinomials for subspecies(Ereshefsky1994, 1997).
There are other problemswith the use of binomials, and they arise regardless
'
of whether one is a skeptic of the speciescategory. One of Linnaeuss
motivations for introducing binomials was his belief that a biologist should
memorize the taxonomic positions of all the specieswithin the kingdom he
studies (Cain 1958, 156 ff.). Linnaeusthought there were too many species
to do that, more than ten thousand plant speciesby his estimation (Atran
1990, 171). But he believed that the number of generain the world was sufficiently
small- approximately three hundredplant generaand three hundred
animal genera(Mayr 1969, 344)- and that the number of generawould not
increasesignificantly. With not too much difficulty , a biologist could memorize
the namesand positions of all the generain a kingdom. Thus, binomials
served as handy guides for memorizing the taxonomic positions of all the
speciesin a kingdom.
However, Linnaeusdid not envision a world where evolution continually
gives rise to new speciesand genera. As a result, he grossly underestimated
the diversity of the organic world . He believed that there were 312 animal
genera; more recent estimatescite more than 50,000. As Mayr (1969, 344)
"
observes, a generic name no longer tells much to a zoologist except in a
"
few popular groups of animals. Matters are even worse when we turn to
plants. Linnaeusthought there were, at most, 6,000 speciesof plants. More
recent studies estimatehundreds of thousands. Becausethere are too many
'
generic names and taxonomic positions to memorize, Linnaeuss original
motivation for assigning binomial names has been lost. Nevertheless, the
namesof speciesmust still include a generic name. Indeed, the genus of a
speciesmust first be determinedbefore a speciescan even be named, which
may seem like an innocuous requirement. According to Cain (1959, 242),
"
however, the necessity of putting a speciesinto a genus before it can be
named at all is responsiblefor the fact that a great deal of uncertainty is
." Simply put, some
wholly cloaked and concealedin modern classifications
- a
species are placed in genera without adequate empirical information
practicethat is the result of an outdated rule of nomenclature.
Another problem with binomials involves taxonomic revision. Classifications of the organic world are constantly revised. For example, a species
assignedto one genusmay be reassignedto a different genusas the result of
a n~w DNA analysis. The assignmentof binomials further complicatessuch
revisions by requiring that a species be given a new name when it is
assigned to a different genus. Consider a simple case. When the species
, its name was
Cobitis heteroclitawas found to belong to the genus Fundulus
changed to Fundulusheteroclita(Wiley 1981, 399). As a result, all previous
classificationsof that speciesbecame outdated. Taxonomic revision is an
297
Frph~~kv~Species
andtheLinnaean
Hierarchy
298
v . S~
es Begone!
only those problems concerning species, yet many other aspectsof the Linnaean
system are problematic as well (Griffiths 1976, Ereshefsky1994, de
Queiroz and Gauthier 1994). Given the problems facing the system, perhaps
we should consider replacing it . The next and final section of this chapter
briefly introduces somealternative systems.
ALTERNA
TIVFSTO THELINNAEANSYSTEM
A comprehensiveintroduction to alternative systems of classifications
, not
to mention a proper comparisonbetween them and the Linnaeansystem, is
beyond the scope of this section. Its aim is more modest: (1) to show that
alternative systemsof classificationdo exist, which meansthat the abandonment
of the Linnaeansystem would not leave biological taxonomy in a vacuum
, and (2) to show that there are promising non-Linnaeansystemsworthy
of attention and further development.
One aspectof the Linnaeansystem that should be replacedis its hierarchy
of categoricalranks. The challengehere is not to the assumptionthat life is
hierarchically arranged, though some theorists have questionedthat assumption
(Hull 1964). The challengeis to the continued use of the Linnaeancategories
to capture that hierarchy. For Linnaeus, taxa of a particular rank
should be comparable. All speciestaxa, for example, should have some common
feature that distinguishes them from all other types of taxa. This
assumptionwas also held by the proponents of the Modem Synthesisand is
held by many contemporary biologists. Yet species taxa may be noncomparable
units: some are interbreeding units, others are phylogenetic (see
the secondsection of this chapter). If such taxa are noncomparable
, then it is
not very meaningful indeed, it is misleading to say that they belong to a
single, unified category. In addition, the distinction between speciestaxa and
generahas been called into question (seethe first section). If that distinction
lacks a basisin nature, then it is misleading to designateone taxon as a species
and another as a genus.
For these reasons, and many others, some biologists have suggestedthat
the categorical ranks of the Linnaean hierarchy lack an ontological basis
(Hennig 1969, Griffiths 1976, Eldredge and Cracraft 1980, 168). Some have
even suggestedthat we completely abandonthe Linnaeancategoriesbecause
not only do those categorieslack independentexistence, but their continued
use misleads biologists into thinking ~hat there are essential differences
among taxa of different Linnaeanranks (Griffiths 1976, Ax 1987, chap. K).
Two non-Linnaean systems for indicating the subordination of taxa have
.
been.offered: an indentationsystemand a numericalsystem
The indentation system indicatesthe relative taxonomic positions of taxa
by il\denting subordinatetaxa below their higher taxa (Farris 1976, Ax 1987,
Gauthier et al. 1988). Sister taxa have the same indentation. Here is an
examplefrom Gauthier and colleagues(1988, 65):
299
: Species
andthe Linnaean
Ereshefsky
Hierarchy
Amniota
Synapsida
Reptilia
Anapsida
Diapsida
Mecopteroidea
)\ nnphiesmenoptera
Trichoptera
2.2.2.2.4.6.1.2
Lepidoptera
2.2.2.2.4.6.1
The use of numericalprefixes avoids Wiley 's objection that the indentations
of different taxa are difficult to compare, and like indented classifications
,
numerical ones indicate hierarchicalrelations without using Linnaeanranks.
Supporters of the Linnaean system, however, suggest that the numerical
alternative is inferior because" numerical prefixes are not the languagesof
"
ordinary use and are foreign to our efforts to communicate ( Wiley 1981,
202; seealso Eldredge and Cracraft 1980, 224- 225). In addition, the numerical
prefixes for lower taxa in large classificationsmay be inordinately long .
Anyone who desires a non-Linnaean system can avoid these problems by
adopting the indented approach. Alternatively , they can use the numerical
approachand argue that its inconveniencesare slight comparedto the costs
of using the Linnaeansystem.
300
V. SpeciesBegone!
301
/
: Speciesand the LinnaeanHierarchy
Ereshefsky
ACKNO
IWLEDGMENTS
I thank David Hull , Jay Odenbaugh, Tony Russell, and Rob Wilson for their
helpful comments on earlier drafts of this chapter. Financial support was
generouslyprovided by the Killam Foundation.
NOTFS
'
1. One might wonder whether Unnaeuss genera/higher taxa distinction, and the speaesj'nigher
'
taxa distinction of the Modern Synthesisare one and the samedistinction. After all, Unnaeuss
fructification systemsare the reproductive organs of plants, whereasthe speciesof the Modern
Synthesisare groups of organismsthat can success
fully interbreed. Thesedistinctions, however,
are quite different. Unnaeus's sexual system classifiesorganisms according to qualitative similarities among fructification struCtures
, but with the advent of Darwinism, qualitative similarity
. Furthermore, many characteristicsof fructificatook a backseatto genealogicalconnectedness
tion structures(numbersof stamensand pistils, for example) are of little functional significancein
conspecificreproduction (Mayr 1982, 178).
2. I am not denying that the membersof asexualspeciesare causallyconnectedin somemanner;
after all, they are connectedthrough parent- oft'spring relations. Of concern here, however, are
302
V. SpedesBegone!
to distinguishspecies
connections
, suchasinterbreeding
, whicharesupposed
intragenerational
.
fromhighertaxabut aremissingin asexualspecies
3. Species
pluralismis a growthindustryin thephilosophyof biology, with severalversionson
. Thetype of pluralismadoptedin this chapterdiffersfrom otherprominentversions
the market
to be spatio' s (1984) formsof pluralismallow species
in two ways. Dupre's (1993) andKitcher
form
the
used
here
that
classes
of
unrestricted
;
requires species
organisms
pluralism
temporally
's
an
that
entities
.
Mishler
and
Brandon
(1987
)
organismbelong
pluralismrequires
genealogical
to only onespecies
; thepluralismadoptedherepermitsan organismto belongto two different
speciesat the sametime. For full illustrationsof the versionof pluralismused here, see
1992aand1998.
Ereshefsky
4. One could avoid this conclusionby denying that both phylogeneticand interbreeding
in Ereshefsky
that is considered
1992aand1998.
arespecies
, a response
species
taxashare
is a unifyingprocessthat all species
S. Onemightrespondthat inheritance
, whichis
fromothertypesof taxa. Recallthat we
esspecies
correct
, but it is not a processthatdistinguish
.
arelookingfor a unifyingprocessthat occursin all andonly species
6. One might try to savethe speciescategoryby employingBoyd's (1991) "causalhomeo
'
" accountof naturalkinds. However
stasis
, I dont think that this particularphilosophical
will help. ConsiderGriffiths's (1997, chapter8 in this volume) applicationof the causal
approach
offeredagainstthe speciescategoryin this
homeostasis
approachto taxa. The ho arguments
- do not distinguish
. The naturalkind traits Griffithscites- homologies
chapterremainunaffected
fromhighertaxa
for
thus
no
new
from
,
taxa
;
grounds distinguishing
species
species higher
areoffered
. Furthermore
, Griffithsbuysinto a pluralisticaccountof speciesvery similarto the
.
in thischapter
onearticulated
, sothedisunityof the species
categoryremains
es to
betweenthe interbreedingand phylogeneticapproach
7. For examplesof discrepancies
causebiologiststo
wheretaxonomicdisagreements
. Forexamples
, seethesecondsection
species
1994.
, seeMichener1964andEreshefsky
assigndifferentnamesto the sametaxon
rulesof nomenclature
,
8. Foradditionalproblemswith thebinomialruleandtheotherUnnaean
seeEreshefsky
1994.
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Ereshefsky
Hierarchy
12
The debateabout speciesconceptsover the last twenty years follows acuri ous pattern. Rather than moving toward some kind of consensus
, as one
an
the
trend
has
been
toward
ever
might expect,
increasingproliferation of
concepts. Starting with the widely acceptedspeciesconcept that took precedence
in the 1940sand 1950s as a result of the Modem Synthesis- the biological
speciesconcept (BSC) we heard calls for change from botanists,
behaviorists, and others. Despite the babel of new concepts, the BSC continues
to have fervent advocates(A vise and Ball 1990, A vise and Wollenberg
1997) and hasitself spawnedseveralnew variants. A recentpaperby Mayden
'
(1997) lists no fewer than twenty -two prevailing conceptslWe cant seemto
eliminate any existing concept, only produce new ones.
Why ? The obvious conclusion one might draw- that biologists are contrarians who want to make their own personal marks in a debate and thus
coin their own personalconceptsto defend- is really not the case; this is no
debate about semantics. The conceptualdivisions are major and real. In my
opinion, the plethora of ways in which different workers want to use the
speciescategory reflects an underlying plethora of valid ways of looking at
biological diversity . The way forward is to recognize this view and face its
implications: the basis of the confusion over speciesconceptsis a result of
heroic but doomed attempts to shoehorn all this variation into an outdated
and misguided classiAcation system, the ranked Linnaeanhierarchy. Most of
the confusion can be eliminated simply by removing the ranks. The issues
that remain can then be dealt with by carefully considering what we want
formal classiAcation to represent as the general referencesystem and then
by carefully specifying criteria for grouping organisms into these formal
.
classmcations
To develop this argument, I mst make the casefor generalizingthe species
problem as a specialcaseof the taxon problem. For a consistent, general reference
classiAcationsystem, all taxa must be of the sametype; speciesshould
be regardedas simply the least-inclusive taxon in the system. Then, I review
the' reasonswhy phylogeny provides the best basis for the general purpose
classmcation
: speciesshould be consideredas just another phylo genetically
based taxon. Next , I addressthe recent calls for rank-free classiAcation in
general and pursue the central thesis of this paper: the speciesrank must disappear
along with all the other ranks. Finally, I explore the practical implications
of eliminating the rank of speciesfor such areasas ecology, evolution,
and conservation.
SPECIFS
ASJUSTANOTHERTAXON
308
V. Species
!
Begone
level, and large guilds of organisms from divergent speciescan act as one
group ecologically in somesituations.
Finally, there have been attempts to distinguish speciesfrom all other
taxa phylo genetically (Nixon and Wheeler 1990, Graybeal 1995, Baum
1992). In this view, speciesare the smallest divergent lineage, inside of
which there is no recoverabledivergent phylogenetic structure (only reticulation
). Again, nice in theory, but unsound empirically, at least as a general
principle (Mishler and Donoghue 1982, Mishler and Theriot 1999). Some
biological situations At the model well (e.g., in organisms with complex
and well-deAned sexual mate recognition systemsand no mode of asexual
propagation). However, in many clonal groups (e.g., aspen trees, bracken
fern) discernible lineages go down to the within -organism level (the problem
"
"
of too little sex ; seeTempleton 1989). On the other hand, occasional
"
"
horizontal transfer events ( reticulations ) occur between very divergent
"
"
lineages (the problem of too much sex ; see Templeton 1989). In all
such cases
, a large gray area exists between strictly diverging patterns of
gene genealogiesand strictly recombining ones (d . Avise and Wollenberg
1997).
To sum up, we have no and are unlikely to have any criterion for distinguishing
speciesfrom other ranks in the Linnaeanhierarchy, which is not
to say that particular speciestaxa are unreal. They are real, but only in the
sensethat taxa at all levels are real. Speciesare not special.
THE NECESSITYFOR PHYLOGENETIC CLASSIFICADONS
The debate over classiAcation has a long and checkeredhistory, but this
essay is not the place to detail the history fully (see Stevens 1994 and
Ereshevsky, chapter 11 in this volume). I want to begin with the conceptual
upheavalin the 1970s and 1980sthat resulted in the ascensionof Hennigian
phylogenetic systematics(for a detailed treatment, seethe masterfulbook by
Hull 1988). Many issueswere at stakein that era, foremost of which was the
nature of taxa. Are they just convenient groupings of organismswith similar
features, or are they lineages, marked by homologies? A general, if not completely
universal, consensushasbeen reachedthat taxa are (or at least should
be) the latter (Hennig 1966, Nelson 1973, Farris 1983, Sober 1988).
A full review of the argumentsfor why formal taxonomic namesshould
be used solely to representphylogenetic groups is beyond the scopeof this
paper, but they can be summarizedas follows. Evolution is the single most
powerful and general process underlying biological diversity . The major
outcome of the evolutionary processis the production of an ever-branching
phylogenetic tree, through descent with modiAcation along the branches.
This results in life being organized as a hierarchy of nested monophyletic
gr.oups. Becausethe most effective and natural classiAcation systems are
"
"
those that capture the entities resulting from processes that generate the
309
. Mishler:
?
c. IiIns Rid of Species
A number of calls have been made recently for the reformation of the
Linnaeanhierarchy (e.g., de Queiroz and Gauthier 1992). Theseauthors have
emphasizedthat the roots of the Linnaeansystem are to be found in a nonevolutiona
worldview - a specially createdworldview . Perhapsthe idea of
fixed ranks made some senseunder that view, but not under an evolutionary
worldview . Most aspects of the current code, including priority , revolve
.
around the ranks, which leads to instability of usage. For example, when a
change in relationships is discovered, several names often need to be
changedto adjust, including the namesof groups whose circumscription has
not changed. Authors often frivolously change the rank of a group even
though there is no change in postulated relationships. Although practicing
310
v . SpidesBegone
!
systematists know that groups given the same rank across biology are
not comparablein any way (i.e., in age, size, amount of divergence, internal
diversity, etc.), many users of the system do not know this. For example,
ecologistsand macroevolutionists often count numbersof taxa at a particular
"
"
rank as an erroneousmeasureof biodiversity . The nonequivalenceof ranks
meansthat at best (to those who are knowledgeable) they are a meaningless
formality and perhaps not more than a hindrance. At worst, formal ranks
lead to bad sciencein the hands of a user of classificationswho naively
assumesthat groups at the samerank are comparablein someway.
It is not completely clear at this point how exactly a new code of nomenclature
should be written , but the basicsare clear. Such a new code should
maintain the principle of priority (the first name for a clade should be followed
) and other aspectsof the current code that promote effective communication
of new namesto the community. However, the major changewould
be that the Linnaeanranks (e.g. phylum, family) should be abandonedfor
more efficient and accurate representation of phylogenetic relationships.
Instead, namesof cladesshould be hierarchicallynesteduninomials regarded
as proper names. A clade would retain its name regardlessof where new
knowledge might changeits phylogenetic position, thus increasingnomenclatorial stability . Furthermore, becauseclade nameswould be presentedto
the community without attached ranks, users would be encouragedto look
at the actual attributes of the cladesthey compare, thus improving research
in comparativebiology .
It is important to emphasizethat despite misrepresentationsto the contrary
, theorists who advocate getting rid of Linnaean ranks do not at all
advocate getting rid of the hierarchy in biological classification. Nesting of
groups within groups is essentialbecauseof the treelike nature of phylo
genetic organization. Think of a nonsystematic example: a grocer might
classify table salt as a spice, and group spicestogether under the category
food items. This simple hierarchy is clear, but requires no named ranks to
be understood. In fact, all human thought is organized into hierarchies, and
becoming educated in a field essentially means learning the hierarchical
arrangementof conceptsin that field. Taxonomy is unusualin the assigning
of namedranks to its hierarchies; they are superfluousto true understanding.
RANK
GE1TINGRID OF THE SPECIES
Curiously, so far in this debate, even the advocatesof rank-free phylogenetic
classi6cationhave retained the speciesrank as a specialcase. All other ranks
are to be abandoned, but the speciesrank is to be kept, probably becausethe
speciesconcept is so ingrained and comfortable in current thinking . However
, all the arguments that can be massedagainst Linnaeanranked classi6c.auon in general can be brought to bear against the speciesrank as well. As
difficult as it is to overthrow ingrained habits of thinking, logical consistency
demandsthat all levels in the classi6cationshould be treated alike.
311
?
Mishler: G.tt ;ngRidof Species
Given the background developed in the previous three sections, the conclusion
seemsinescapable
: the speciesrank mustgo the way of all others. We
must end the bickering over how this rank should be applied and insteadget
rid of the rank itself. This solution is truly the " radical solution to the
species
"
problem sought unsuccess
fully by Ghiselin (1974). Biological classification
should be a set of nested, named groups for intemested clades. Not all
cladesneed be named, but those that are should be named on the basis of
evidence for monophyly (see further discussion of the meaning of monophyly in Mishler and Brandon 1987). We stop naming groups at somepoint
'
approachingthe tips of the phylogeny becausewe don t have solid evidence
for monophyly at the present stage of knowledge. This may be due to rampant
reticulation going on below some point or simply to a lack of good
markersfor distinguishing finer clades. We shouldn't pretend, however, that
the smallestcladesnamedat a particular time are ontologically different from
other, more inclusive named clades. Further researchcould easily result in
subdividing thesegroups or lumping severalof them into one if the original
evidencethat supported them is discoveredto be faulty .
Given the redundancynow presentin speciesepithets(e.g., californicais used
in many genera), there needs to be a way to uniquely place each smallest
named clade in the classification. My recommendationfor nomenclatureat
the least inclusive level under a totally rank-free classificationwould be to
regard namesin a similar way as personal namesare regarded in an Arabic
culture. Each clade, including the least inclusive one named, has its own
uninomial name; however, the genealogicalrelationships of a clade are preserved
in a polynomial giving the lineage of that clade in higher and higher
.
groups Therefore, the familiar binomial, which does after all present some
grouping information to the user, could be retained, but should be inverted.
Our own short clade name thus should be SapiensHomo. The full name for
our tenninal cladeshould be regardedas a polynominal that gives the names
of the more and more inclusive cladesall the way back. To use the human
example, this full name would be something like: SapiensHomo Homidae
PrimateMammalia Vertebra
ta Metazoa EucaryotaLife.3 Again, as in a traditional
Arabic name, this formal and complete name would be used only
and
for the most formal purposes (although it would be very useful
rarely
behind the scenesfor data-basing purposes); the everyday name of the clade
would be SapiensHomo.
PRACn CALIMPLICATIONS
"
rid of species" has another, all too ominous meaning in today's
. Getting
world. Named speciesare being driven to (and over) the brink of extinction
at a rapid rate. What will be the implications of the view of taxa advocated
in this paper? If we get rid of the speciesrank, with all its problems, will we
hamstring conservationefforts? I tend to think not ; scienti6chonesty seems
312
v . Sp~ s Begone!
313
1991, Martins 1996) and the rapid proliferation of ever-improving dadograms for most groups of organisnis, this changecan only be for the best.
, RIP.
Species
NOTFS
1. In Hennigian phylogenetic systematics, homologyis defined historically as a feature shared
by
two organismsbecauseof descentfrom a common ancestorthat had that samefeature.
2. A strictly monophyletic group (a clade) is one that contains all and only descendantsof a
common ancestor. A paraphyletic group is one the excludes some of the descendantsof the
common ancestor.
3. Note that some of the nested cladeswill have formal suffixes
indicating their previous rank
(e.g ., -idae for family). Although these endings would be retained for existing clade names in
order to avoid confusion, there would be no meaning attached to them, and newly
proposed
cladenameswould have no particular suffix requirement.
REFER
-ENCFS
Avise, J. c ., and R. M. Ball (1990). Principlesof genealogical
concordance
in speciesconcepts
andbiologicaltaxonomy
. OrfordSuroeys
in Evolutionary
Biology7, 45- 67.
Avise, J. C., andK. Wollenberg(1997
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). Phylogenetics
. Proceedings
of the
NationalAcademy
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Baum
. D. (1992). Phylogenetic
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in Ecology
andEvolution
7,' 1- 2.
species
concepts
BrooksD. R., andD. A Mclennan(1991). Phylogeny
. Chicago
, ecology
, andbehaoior
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of ChicagoPress
.
de Queiroz
, K. , andJ. Gauthier(1992). Phylogenetic
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and
taxonomy
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Systematics
Faith
, D. P. (1992a
evaluationandphylogenetic
). Conservation
Consm
Jation
diversity. Biological
61, 1- 10.
Faith
, D. P. (1992b
andconservation
: On predictingthefeaturediversityof subsets
). Systematics
of taxa. Cladistics
8, 361- 373.
Farris
, J. S. (1983). The logicalbasisof phylogeneticanalysis
. In N. PlatnickandV. Funk
, eds.,
Adoances
in Cladistics
, vol. 2. NewYork: ColumbiaUniversityPress
.
. Brooks(1990). Phylogenetic
Funk
, V. A., andDR
asthebasisof comparatioe
.
systematics
biology
InstitutionPress
.
WashingtonD.C.: Smithsonian
Ghiselin
, M. T. 1974.A radicalsolutionto thespecies
. Systematic
23, 536- 544.
problem
Zoology
methodin eoolutionary
Harvey, P. H., andMD . Pagel(1991). Thecomparatioe
. Oxford:
biology
Oxford UniversityPress
.
Hennig, W. (1965). Phylogeneticsystematics.Annual Reviewof Entomology10, 97- 116.
. Urbana: University of nlinois Press.
. Hennig, W . (1966). Phylogenetic
systematics
as a process
Hull D. L (1988). Science
: An evolutionaryaccountof the socialand conceptual
development
. Chicago: University Chicago Press.
of science
Levin, D. A (1979). The nature of plant species. Science
204, 381- 384.
314
v . SpeciesBegone!
315
?
: GettingRidof Species
Mishler
Index
Accommodation, 141
conditions, 149
demands, 148- 150
, 148, 176
disdplinarymatrices
empiricistview, 151
, 176
equifeltilityprinciple
. 176- 178
highertaxa
historicity, 154
homeostasis
, 157
inexactandmessy
, 151- 156
kindsandlaws, 151
, 176
methodology
naturalkinds, 146
nonetema1156
, 151- 156
parochialsciences
.
relativismandrealism
, 159- 161
reliableinduction
. 146- 148
., 165
, 161
species
153
,
vagueness
&dencysyndrome
immunode
Acquired
(AIDS), 1.8
. 181, 1.48
Adaptation
, 194
Adrenergiccells, neuraltaxonomy
), 37
AJoant ImInfof SciIncI, TIll (Kitcher
, 94
, ciliates
Aging controversy
model
transformation
. speciation
Anagenetic
56
, 1.1.5
AnalogiesVB.homologies
. 149
Analyticalfunctionalism
Ancestralspecies
, 1.91.
. about, 183
, realism
Ancestry
, 16, 20ft
, subspecies
Angiosperms
., 1.34
Animals
, folkbiology, 1.31
, 1.45
,
Anthropologytaxonomy
, 1.11, 1.1.1.
Antiessentialism
, biologicaltaxa
Antirealism
, 74
. philosophy
1
.5
.
pluralism , 74
Aristotle
essentialism
, 1.87
1
.1
.0
,
physics
Artifacts
causalfacton, 1.75
taxonomy, 1.34, 1.37
Asexual species
bacteria, 7
highertaxa, 1. 88
monism
. 6, 19n
organisms, 1.93
reproduction, 55, 66, 81.n
Attractor space, cognition, 1.64, 1.79n
Autapomorphic species, 78
Bacteria
.1
geneRow, 11
killer, 106
, 14, 19n
taxonomy
, 104
Bacteriophages
BelLG.. 94
theGene(Sapp
), 93
Beyond
, 56, 69
BifurcationmodeLspedation
names
Binomial
, 296
genericandspedfic
revising
, 297
, 296- 298
spedescategory
Biologicaldiversity, 64, 311, 313
.2
, 78, 11
Biologicalspecies
asexua
L6
, 77
conceptVI. de6nition
307
debate
,
exoticvariants
, 134
view, 210
genealogical
geneRow, 7
interbreeding
, 290
dassi&cation
Linnaean
,9
, 74
terminology
misleading
(cont.)
Biologicalspecies
naturalkinds, 166
kinds, 168
paradigmatic
Biology. SeealsoEvolutionarybiology
functional
, 13, 166
naturalkinds, 212- 214
, 212
processstructuralism
, 36, 60, 65
species
concepts
structuralhomology
, 224
170
,
Biospecies
(Rubus
), geneRow, 8
Blackberry
Blastationmodel
, speciation
. 56, 69, 81n
inertia, 220
Burden
. phylogenetic
. species
, 276- 278
Categorization
concepts
Causalhomeostasis
, 217, 272, 295, 303n
, 147
Causallysustained
generalization
Causation
, 28- 31
analysis
, xv, 263
patterns
, 28- 31
pluralism
Cells,generaltaxonomy
, 193
Chemistry
, 148
disdplinarymatrices
elemental
andfundamental
, 158
reliableinduction
, 147
, 35
species
concepts
Children
, biologicalconcepts
, 268- 270
cells
neural
,
, 194
Cholinergic
taxonomy
Chromosomes
, 108
Chronometrics
, 111
Ciliates
. SeealsoTetrahymena
clonalaging
, 94
evolutionarytree, 101
history, 107- 109
, 98
matingsystems
, 103
taxonomy
Ciliatology, 93- 95
dadesandCladism
, 51, SOn
65
,
biology
extreme
, 179
modest
, 182
, 291
monophyly
312
, 314n
naming
,
dadistic species
, 78, 131, 219
model speciation
, 56
Cladogenetic
Clans
, biology, 51, 65, SOn
. Classes
, species
, 32, 67. SeealsoKinds
Classification
. SeealsoTaxonomy
alternativesystems
, 299- 302
35
,
generalpurpose
maingoals,5
, 36
theory- dependent
318
Inde~
Climatechange
, ecologicalfracturing
, 128
Clonalaging
, 94
, ciliates
Clones
, 51, 65, 8On
dusters. SeeHomeostatic
propertycluster
( HPq
Cognition
bias, 271, 279n
children
. 268- 270
folkbiology, 231
, 248
genericspecies
hmnanevolution
. 248
Cohesionspecies
, 10, 78, 122
, realism
, 188
Commonality
Commonsense
, genericspecies
, 250
semantics
157
,
Compositional
Connectionism
, 263
Conservatism
, taxonomic
, 15- 19
149
Continuum
, deAnitions
,
, tacitjudgments
, 177
Conventionality
, N., 252
Copernicus
Corliss,J., 97
Counterfactua
I force, 216
' s model 27
-law analysis
, Hempel
,
Covering
Creationism
, 36, 38
Culturalevolution
folkbiology, 231
, 249
genericspecies
, 231
taxonomy
Darwin, C. R.
historicalessences
, 220
, 76
lineageconcept
, 29S
species
category
, 3, 18
taxonomy
Darwinism
, development
aL 223
Dawkins
, R., 119
Definitions
continuum
, 149
, 141, 168
explanatory
, 141, 168
programmatic
Demographicexchangeability
,9
Descent
, realism
, 183
, canalized
, 224
Developmentalism
,
Developmental
psydtology, genericspecies
236
of Ta.ronomic
Development
TheorySince1851
(Gilmour), 3
Dick, S. J., 23
Difference
matrix, molecularanalysis
, 112
Disciplinarymatrices
accommodation
, 148, 176
semantics
, IS7
compositional
homeostasis
, IS7
),
Disco
O" Y of Time(TouJminandGoodfield
111
Diversity. SeeBiologicaldiversity
, T., 108, 124
Dobzhansky
taxa
, 189
), species
Dogs(Canis
Dupre,J., 30, 160, 202
, 78
species
Ecological
fractured
. 124, 128
, 74
terminology
misleading
mosaics
, xiii, 119
. 13
pluralism
taxa
, 308
Economic
theory
feudalandcapitalist
, ISS
inexadlaws, 154
naturalkinds, 216
, 170
Ecospecies
Eliminativepluralism
. 24
. 142, IS1
Empiricism
, 310
Entomology
, 127
Environment
, physicalandseledive
149
access
condition
,
Epistemic
, naturalkinds, 169
Epistemology
, 176, accommodation
Equifertilityprinciple
178
. 12S
, punctuated
Equilibrium
, M ., S
Ereshefsky
Essence
, 271- 273. SeealsoHistoricalessences
causalhomeostasis
, 218
childview, 270
realismandkinds, 187
sortalandcausal
, 271
272
or
token
,
type
Essentialism
. xiv, 188
Darwinian
, 209, 287
individualism
, 209- 212
Unnaean
, 287
&pedes
naturalkinds, 196- 201
, 271
psychological
understanding
, 145
. Su alsoFolkbiology
Ethnobiology
, 1.35
genericspecies
, 94
Eukaryotes
, 16
Eurocentrism
, taxonomy
Evolution
biologicaL231
cultural
, 1.31
de6ning
, 65, 82n
sdenti&c, 1.31
.Evolutionarybiology
accommodation
, 166
ciiteria, 37
.319
Index
, 13
pluralism
33
,
species , 166
, 78
Evolutionaryspecies
, 77
lineageconcept
, 130
speciation
, 108
synthesis
, 101
tetrahymenines
, 56
theoryandoperations
5
n
units
,
Evolutio
.
. 1.88- 1.90
highertaxa
, 1.87
speciation
, geneticvs. demographic,9
Exchangeability
de6nitions
, 141, 149, 168
Explanatory
), 119
&tendedPhenotype
, The( Dawkins
, 70
Extinction
. species
Extremecladism
, highertaxa, 179
Feudaleconomy
, 155
Fixation,organisms
, 132
119
,
Folkbiology
canonical
, 253- 256
synopsis
, 231
cognitivenature
culturalevolution
, 249
231
,
genericspecies , 233
, 238- 247
Mayataxonomy
, 232, 234
plantsandanimals
, 244
psychology
scientificsystematics
, 232
, 231, 233
taxonomy
es, 264- 268
thoughtprocess
vivid illusion, 273
Force
, 217
, taxonomy
FormandTransformation
(Goodwinand
Webster
), 213
FrankelJ" 93
Frudi6cationstrudures
, 287, 302n
Functional
biology, 13, 166
, 149
Functionalism
, analytical
andorganisms
, 70
Fusion
, species
Genealogical
spedes
. 75
.antirealism
, 33
imperialism
, 78
lineageconcept
, 191
category
species
GeneRow, 8- 11
bacteria
. 121
interbreeding
, 291
monism
, 7
potential,8
, 54
process
,7
sharpdiscontinuities
, 147
Generalizations
, causallysustained
Generallineageconcept
, 49- 53
history, 76- 78
, 61
ontologyandtaxonomy
, 64- 78
philosophy
Generative
entrenchment
, 220
Genericspedes
, 235
, 248
cognition
commonsense
, 250
culturalevolution
, 249
folkbiology,231
, 238- 247
Mayataxonomy
naturalselection
, 248
inertia
, 221
phylogenetic
science
, 250
universalprimacy
, 231
Genes
, 222
exchange
pools, 107, 110
, 104- 106
tetrahymenines
Geneticalspedes
, 77
Genetics
, 9, 109
Genetics
andtheOriginof Species
(Dobzhansky
), 108
homeostatic
,
, 292
Genotypes
Genusvs. spedes
, 235
Ghiselin
, M. T., 222
Gilmour,J. S. L , 3
God
, spedesconcepts
, 23
Good6eld
, J., 111
Goodwin
, B., 213
Grade
, species
, 119- 123
Gradualism
. phyletic, 128
, biologicalspecies
, 210
Groupingcriteria
'
-law analysis
, 27
Hempels model,covering
, 78, 309
Hennigianspecies
, 112
Heterochronidty
, 190
Heterogeneity
Highertaxa
accommodation
, 176- 178
binomialnames
, 296
, 181
evolutionarysystematism
evolutionunits, 288- 290
extremecladism
, 179
homeostasis
, 141- 146, 180- 182
individuals
, 175
Unnaean
, 286- 288
spedes
, 182
. modestcladism
naturalkinds, 174
realism
, 176- 183
. real
kinds, 182
, 141, 173, 286
spedes
Histones
, 104
320
Index /
Historicalessences
, 219- 221
cladistictaxa
, 219
naturalkinds, 209
. 154
Historicity, accommodation
Homeostasis
217, 272, 295, 303n
causaL
semantics
, 157
compositional
, 157
disciplinarymatrices
, 141- 146, 180- 182
highertaxa
, 164
phenomena
propertytenns, 143
realism
, 180- 182
, 141- 146
species
Homeostatic
) , 142- 144
propertycluster( HPC
accommodation
, 165- 167
essentialism
, 145, 196
andstrategy
, 144
examples
view
203
,
integrationist
naturalkinds, 164- 169
realistview, 200
, 165- 167
species
, 310, 314n
Homologies
, 225
analogies
, 34
homoplasies
sb"uctural
, 224- 226
Hull, D. L. , 38, 222
Humanimmtmode6ciency
virus(fDV), 29
Hume,D., 151, 164
Huxley, T. H., 3
, geneRow, 122, 136n
Hybridspecies
Idealessence
, 271
.
, 33
Imperialismgenealogical
indentationsystem
. taxa
, 299
Individualism
, 1.1.2
biologicaltaxa
essentialism
. 209- 212
realismandpluralism
. 193- 196
taxa
, 189- 191
species
Individualspecies
, 162- 164
Inexactitude
, accommodation
, IS2
Inference
, Mayataxonomy
, 240, 242
Inheritance
, taxa
, 291, 303n
, 290. SeealsoSexual
Interbreeding
species
reproduction
, 292
causallyintegrated
, 291
phylogenetic
structures
, 291
Introduction
to theClassification
ofAnimals
( Huxley
), 3
Isolationspecies
, 78
, tetrahymenines
Isozymeanalysis
, 99
Itzaj. SeeMayataxonomy
Killerbacteria
. 106
Kinds. SeealsoNaturalkinds
empiricistview, IS1
historical
, 123
realismandessence
, 187
L 33
spatiotempora
types, 33
Kinetics
, dllary units, 97
Kitcher,P., 37
Language
, 246
biologicalexpectations
monisticconcept
, 1.6
Lawlessness
, 152
, accommodation
Lawsof nature
, 151, 187
Uk cycles
, 68- 71
, species
Uk forms, Mayaexperiment
, 238- 1,47
natural
161
Uli
R
Ctae
kind
.
),
Uly (
. SetalsoGenerallineageconcept
Uneage
, 10
evolutionary
, 192
genealogical
-level, SO
, 63
population
SO
84n
,
,
species
unication, 68
Unnaean
hierarchy
binomialnames
, 296- 298
, 286- 288
highertaxa
reforming
, 310
, 285
category
species
Unnaeanspecies
alternativesystems
, 299- 302
, 286- 288
highertaxa
sexualsystem
, 287
Linnaeus
, C., 286
Uving kinds
causalfadon, 275
, 276
taxonomy
Locke
, J., 142, 174, 215
, smoking
, 30
Lungcancer
Margulis,L , 107
Matingsystems
ciliates
, 98
, 132
recognition
Mayataxonomy
, 238- 247
experiment
folkbiology, 231
, R. L., 42
Mayden
E.
,
Mayr , 77, 108
, 125, 127
Mayr' s Brake
MendeLG., 108
M~ aphysics
biology, 173
321
Index
thesis
innocence
, 178
muddled
, 31- 34
naturalkinds, 169
, 31- 34
pluralism
taxa
, 173, 184ft
Microbialgenetics
, 109
ModemSynthesis
, 76
evolutionunits, 288- 290
.
, 108
participants
andtaxa
, 288
species
Modestcladism
, 182
, highertaxa
Moleculargenetics
, 109
Monadicpropertyterms, 143
Monism
asexualspecies
,6
geneRow, 7
, 26
language
minimal
, 4
, 73
philosophy
, 14, 2S
pluralism
, 3- 5
species
,4
taxonomy
troubleswith, 5- 11
, 78
Monophyleticspecies
, 40
concepts
entities
, 50, &On
gene8ow, 9
, 310, 314ft
groups
taxa
, 291
Monophyly
cladism
, 17
highertaxa, 40
.
, 11
pluralism
, 57, 81n
spedation
, 235, 1056n
Monospedficgenera
Re
Idt , 1013
Morphogenetic
.7
Mosaics
, 123- 11
, species
., 175
Naturalindividuals
, 161
Naturalkinds
accommodation
, 146, 174
, 168
biological&pedes
.- 1.14
biology, 1.11
constitutiverelations
, 143
force, 1.16
counterfactual
de6nitions
, 149
disdplinerelative,148
empiridstview, 151
for, 1.16
enthusiasm
essentialism
, 196- 1.01
andstrategy
, 144
examples
189
,
hierardty
historicalessences
, 1.09
Naturalkinds(cont.)
homeostasis
, 157
Humeanreconstructions
, 164
vs. individuals
, 145
monadicpropertyterms
, 143
, 168
paradigmatic
reality, 158
relativismandrealism
, 160- 162
scientificrealism
, 187- 189
~ es, 141- 146, 164- 169
structuralhomology
, 224- 226
Naturalselection
, 125
adaptation
cognition,248
, 248
genericspedes
Nestedboxes
, tetrahymenines
, 100- 103
Neuralcrestcells, 193, 198, 2O5n
Neuraltaxonomy
, 193- 196
Neuroscience
, 202
Neurotransmitters
, taxonomy
, 195
Niches
, 124, 136n, 222
Nomenclature
, taxonomic
, 15
Nominalism
, 142
Nonetemalde6nitions
, accommodation
, 156
Nonintrinsicdefiningproperties
, 153
Numericaltaxonomy
, 36, 300
Oak(Quercus
), species
, 7, 9
concept
Objectivestructures
, 170
Objectivistphilosophy
, 37
OnthePluralityof Worlds( Whewell
), 2.3
, 61
Ontology, species
problem
, 189
Ordering
, realism
Organisms
classifying
,3
environment
, 127
fusion
, 70
, 7, 19n
lineage
materecognition
, 132
ReIds, 213
morphogenetic
, 130
populations
species
individuality,67
Originof Species
( Darwin
), 76, 295
, 65
Paleontology
naturalkinds, 168
Paradigmatic
Paramecia
. SeealsoCiliates
. matingsystems
, 98
Paramebic
individuation
, 202
\
, 3141
Paraphyletic
groups
, 57
Paraphyly
, 51, SIn
lineages
pop~ tions, 59
322
Index /
Parochial
sciences
, accommodation
, 151- 156
Particulars
, naturalkinds, 191
Pattern
Formation
: CiliateStudies
andModels
(Frankel
), 93
Periodictable, 35
Pheneticism
, 38
monism
, 5
similarity, 190
Pheneticspecies
, 78
criteria
, 36
, 308
taxonomy
theoryandoperations
, 55
, 35, 119
Phenomenological
species
, speciation
, 129
Phenotypes
Philosophy
biology, 252
, 64
generallineageconcept
monism
, 73
, 37
objectivism
ontology, 234
, 27
pluralism
issues
, 64
species
, species
Phyla
, 57, 81n
concepts
Phyleticspecies
, 128
gradualism
stasis
, 127
transformation
, 56
, 122, 136n
Phylogenetic
species
cladism
, 291
classmcation
, 309
inertia
, 220
interbreeding
, 291
, 74
misleading
terminology
, 12
pluralism
, 40, 309
systematics
trees
, 35
, 18, 8On
Phylogeny
, 295
Phylospecies
Physicalenvironment
, 127
, Jo
, 268
Piaget
Plants
folkbiology, 232
geneBow, 7
Plato, 215
Pluralism
antirealism
, 25, 74
casefor, 11- 15
causation
, 28- 31
aiteria. 36- 38
eliminative
, 24
Godconcepts
, 23
,
individuality 193- 196
, 26
language
, 31- 34
metaphysics
monism
, 14, 25
, 73
philosophy
, 23- 25
questioning
realism
, 193- 196, 202- 204
reflexivity, 26
, 11, 31, 290
species
, 191- 193
category
species
Worlds
(Dick), 23
Pluralityof
cells, taxonomy
, 195
Pluripotential
Polyphyly, 57, 81n
, 202
Polytheticindividuation
165
,
Polytypicspecies
Populations
essentialism
, 209
, 50, 53, 81n
lineages
Mendelian
, 108
neuraltaxonomy
, 202
, 130
organisms
, 59
polyphylyandparaphyly
views
192
,
reproductive
Positivismvs. postmodemism
, 43
Preference
, 237- 247
, taxonomy
, 42, 44
Primaryspecies
, 188
Priority, realism
Process
es, unifying
, 54
st Ncturalistbiology, 212
Process
definitions
, 141, 149, 168
Programmatic
, 215
Projectable
properties
, 94, 110
Prokaryotes
Promiscuous
realism
, 160
Properties
monadicterms, 143
-level, 129
population
, 215
proledable
, 273
Prototypespecies
. SeeCiliates
Protozoa
, 150
Psychofunctionalism
essentialism
, 271
Psychological
Psychology
folkbiology,244
ontology, 234
, 245
taxonomy
Punctuated
, 125
equilibrium
Racism
, 251, 2S7n
Rand,Ayn, 37
Rank
criteria
, 210
folkbiology,234, 2S6n
inference
, 240, 242
, 2.38- 247
Mayataxonomy
Rank-freetaxonomy
, 310
"
31.3
Index
Realism
accommodation
. 159- 162
descentandancestry
, 183
andkind, 187
essence
highertaxa, 176- 183
homeostasis
, 180- 182
HPCview, 200
individuality, 193- 196
naturalkinds, 170, 187
, 74
philosophy
. 25, 169, 193
pluralism
, 159- 162
promiscuous
sdentific
, 142, 183n
, 78, 131
Recognition
speaes
. 26
, pluralism
Reflexivity
Relationalspedes
, 71- 73
, 159- 162
Relativism
. disdplinary
Reproduction
ftssionandbudding
, 70
sexualandasexual
, 55
, 191
species
Reproductive
Retinalganglioncells, neuraltaxonomy
, 195198
add ( RNA), 112
Ribonucleic
Ribosomes
, 104, 112
, 108
Ribozymes
, J., 93
Sapp
Sdence
folkbiology, 231
, 250
genericspecies
naturalkinds, 187- 189
, 27
pluralism
, 232
systematics
, 231
taxonomy
wars, 26
, 217
, taxonomy
Scope
environment
Selective
, 127
Semantics
, 157
, compositional
Semantic
trickery, 83n
Settheory, 32
: TheHistoryof an
Su andDeathin Protozoa
Obsession
(Bell), 94
Sexualreproduction
, 55, 68
, geneRow, 7
Sharpdiscontinuities
71
,
Siblings
, 35
Similarity
, G. G., 77
Simpson
Smith
, j . M., 93
, 30
Smoking
, lungcancer
Societies
, 246
, small-scale
, 26, 218
Sociology
Sonneborn
, T., 98
Sortalessence
, 271
, 37
Sortingprindples
Spedation
evolutionunits, 289
Mayr' s theory, 125
models
, 56
, 127- 134
phyleticevolution
taxa
, 289
Spedes
accommodation
, 165- 167
biologicaldivenity, 64
causation
, 263
patterns
ciliates
, 93- 95
classi&cation
, 34
aiteria, 36- 38
Darwinian
, 76
de6nitions
, 63
, 19
earlyusage
eliminating
, 307
essentialism
. 187
fusion
, 70
geneBow, 7- 11
VI. genus
, 235
grade, II9 - 123
, 141, 173, 286
highertaxa
homeostasis
, 141- 146
individuals
, 67, 162, 201
life cycles
, 68- 71
50
,
lineage
, 31- 34
metaphysics
monism
, 3, 73
mosaics
, I2.3- 127
naturalkinds, 141, 164
ontology, 31
, 67
organisms
, 64- 78
philosophy
11
.
pluralism , 2.3, 31, 169, 290
rank-free, 311
realismandantirealism
, 74
setsVI. classes
, 32
, 8- 11
sharplydifferentiated
taxa
, 169, 189, 199
vivid illusion
, 273- 275
Spedescategory
binomialnames
, 296- 298
causalhomeostasis
, 295, 303n
, 191
genealogical
, 292
heterogeneous
, 294
implications
Unnaean
, 76, 285
hierarchy
neuralaest cells, 193, 2O5n
ontology, 61
. 191- 193
plw:aiism
31.4
Index
, 49, 63
properties
, 191
reproduction
, 61, 171
taxonomy
, 1.77
teleology
, 1.51- 1.53
Spedesconcepts
, 39, 41
applicability
, 1.76- 1.78
categoriation
children
, 1.68- 1.70
aiteria, 60
debate
, 307
, 53
lineages
,
plethora34- 36
es, 54
process
, 55
reproduction
structural
, 34
, 55
theoryaM operations
unity anddiversity, 53- 60
Spedesproblem
causes
andsolution
, 61- 64
Unnaean
hierardty, 173
, 61
ontologyandtaxonomy
taxa
, 171- 173
.
, 131
Sped6cmaterecognitionsystem
Stasis
, phyleticevolution
. 7- 134
, 11
Stemspecies
,9
Structuralconcepts
, 34, 1.1.4
Substance
, ontologicalproperty, 1.34
Swordfish(Xiphophorus
), interbreeding
, 1.93
, 310
Symplesiomorphies
, 310
Synapomorphies
10
,
Syngameons
, evolutionary
, 108
Synthesis
, biology, 1.33, 1.50
Systematics
, 35
Systematism
Taxa
. SeealsoHighertaxa
binomialnames
, 1.96
indentationsystem
, 1.99
individuality,189- 191
life form, 1.34
nmnerical
, 300
system
1
.98
;
placement
, 169- 173
pluralisticrealism
, 1.98, 307
problems
ranking
, 301
,
spedes308
, 171- 173
spedesproblem
Taxonomy
bacteria
, 7, 14, 19n
basiclevels,1.37
, 61
categories
conservatism
, 15- 19
Eurocentric
, 16
, 11
evolutionary
folkbiology,231, 233
es, 16
historicalprocess
Unnaean
hierardty,62
, 238- 247
Mayaexperiment
196
193
neuraL
, 298
problem
placement
rankandpreference
, 237- 247
rank&ee, 310
-related
, 276
species
Teleology
essentialism
, 245
, 277
category
species
Telomeres
, 108
Tdrah,VmtnR
, 95- 103. SeealsoCiliates
strains
amiaonucleate
, 99
axenicaIlture, 106
clonalaIlture, 99
, l04
complementarity
corticalfeatures
, 96
evolutionarytree, 101
, 106
geneticissues
history, 107- 109
, 99
isozymeanalysis
, 98
matingsystems
molecular
, 111
analysis
nestedboxes, 100- 103
Thought
children
, 268- 270
folkbiology, 264- 268
es, 265
process
typology, 37, 209
Totemism
, 249
Toulmin
, S., 111
Turnoverpulsehypothesis
, 126, 128
, 37, 209
Typologicalthinking
Uni6cationism
, 188
naturalkinds
r,' 68
individualih
.
), s
UnitsofEvolution
, The(Ereshefsky
species
, 53- 60
Unity, species
concepts
, 39
, species
concepts
Universality
Universalprimacy
, folkbiology, 231
, 153
, accommodation
Vagueness
, 29
Vi ! logy , causation
Vivid illusion, species
, 273- 275
, G. P., 225
Wagner
Webster
, G., 213
Whewel
LW., 1.3
, L, 49
Wittgenstein
325
Index
Zoolo
~f., generic
species , 235