Contributions
in
Conceptual issues
Alexandr P.
phylogeny, taxonomy,
and
Keywords: Phylogeny,
nomenclature
Academy of Sciences, Profsoyuznaya
Russian
Street 123, J17647 Moscow,
taxonomy, phenetics, cladistics, phylistics, principles of nomenclature,
Abstract
On
les
compare
trois
Phylogenetic hypotheses
on
tested
order of
in
things
in
(usually
of a set of presumptions,
certain
a
and
designed
are
the basis
describing
that
of
is,
nature. These
tique
la
et
phylistique
dernier terme
evidence
sound
for them
evidence
accepted
but
exists,
practical
A
realization
comparison
is
à
explicite
représenter
à
hitherto
proaches
them. A set of the most current
approach
of
explicitly
and
taxonomy
from
both
and the
certains
features
of evidence
use
The
phylistic approach
of the basic
answering
aims of
has certain
s’appuie,
of the
few
basic
explicitly:
are
fond,
sont
miquement
Nine
principles.
six
of them
are
found
of these
rely
to
principles
practice,
continuum
a
class
and the
separating
found
to be
(a
a
notion
it
from
combining
of subtaxa
other
the best
such
a
class
to
The
type
within
bien
la
un
qu’un
et
de
type
de
travailler
le
est
présomptionscertain
ordre
de
être
acceptées
leur
appui,
des contre
en
of both
by
a
de réalisation
problème.
base
nature.
Ces
en
l’absence
On discute
plus usuelles,
en
pratique
une
et
de
de
et
the
on
série
décrivant
assertions
série
de
cette manière
réponses
aux
de
taxonomique
base.
trois
six
une
classe,
tels
taxono-
taxono-
seulement
pratique
combinant
de
étant
autres
compatibles
La
Neuf
sont
avec
taxonomique
ni
individu,
un
des traits distinctifs
subtaxons,
meilleur
le
lacune.
continuum
instrument
concept
On considère
à notre
que
le
séparé
disposition
concept
s’il
s’agit
taxon-continuum.
Introduction
is
scientists
Frequently,
de
un
methodological
is
one
preuves
assez
un
ditional
do
take
not
much
notice of
exemple
means
to
1970), i.e.,
and methods
and
to
thing
pass
rejecting
bloodless, in
certain
able, albeit in
the
d’aborder le
may
data
a
that
approaches
kind
to abandon those
along
scientific
they usually
that
means
meaningful
and
tra-
with the under-
them with
even
re-
of scientific
are
in
of these
some-
ones,
result in
fact work-
somewhat different domain,
obtained by
still be
a
replace
rarely
How-
and methods of the
yield satisfactory
to
raise
different. Revolutions,
à
soli-
tempted
lying methodology
of their work.
grounds
time fail
(Kuhn,
means
doivent
présomptions
présente
de
concept
vérifiées
d’une
preuves
leur
gap
concept
the
telles, indépendamment des
validité.
des
la
d’assertions
mots,
dans la
seulement
phylogénétiques
concret
choses
comme
mais
leur
d’autres
sur
les
(notion
une
par
avec
revolution
implicite)
ayant
current
sults,
de manière
de
dont
formulés,
taxon n’est ni
l’individu)
work time after
hypothèses phylogénétiques
taxonomic
et dans les relations
recherche
taxonomiques).
continuum
continua
du
and three
Résumé
(normalement
de la
preuves
nomenclature
(donc
ever, when the standard
élaborées
la
la
indépendants,
dépendants
classe
d’autres
the
sont
de
usage
a
of the taxon-continuum.
Les
Ce
de manière
individual,
is delimited
operate
on
formulated
from
nor an
cladis-
la taxonomie.
explicitement
the
only compatible
features
some
that
continua.
available tool
are
Judging
concepts.
taxon is neither
individual)
are
taxonomically independent,
taxonomic
particular
taxonomic
ultimately
de
que
taxonomy.
is
fondamentaux
son
plusieurs principes
sur
concepts
taxa
in
mais
taxonomically dependent, that is, they
with
but
nomenclature
essaie
approche qui
une
dans
avantages
miquement
de
Taxonomic
la
question. L’approche phylistique pré-
en
considère
au
principes
certains
advantages
phénétique,
dans la similitude
fondamentaux
courante montre
involved.
la
apand
cladistics,
relatedness
savoir
les traits
temps
taxons
L’auteur
taxonomic
the basic
its
des
problèmes
The latter term denotes
represent
particularly
similarity
a
to
parenté
même
example
presented.
main
developed, viz., phenetics,
that tries
traditional
derived
is
three
factual
a
en
source
sente
and
approach
of the
phylistics (= evolutionary systematics).
an
of reasonably
à
particulier
en
et
whatever
matter
no
the absence
discussed,
of the
made
in
only
is
such,
as
pleading against
phylogenetic presumptions
of a
type
systématique évolutionnaire).
(=
s’applique
traditionnelle
be
to
are
Russia
approches taxonomiques principales
développées jusqu’à présent,
statements
(1996)
Amsterdam
paleoentomology, Xyelidae (Vespida)
concept,
statements
3-41
(1)
Publishing bv,
Rasnitsyn
Paleontological Institute,
implicit form)
66
Zoology,
to
SPB Academic
while
approaches
worthy of being
reas-
4
A.P.
sessed
rather
preferable
ground
it, in
of
the
than rejected.
examine
to
scientific
a
hope
approach prior
and thus
odological deficiency
proach
and the results of its
objective
of the
seems
abandoning
to
Each
the ap-
cessive
present publication.
in
step
quired,
This is the
application.
Phylogeny
its meth-
safeguard
to
taxonomy, and nomenclature
Phylogeny,
back-
methodological
recognize and improve
to
it
That is why
the
-
Rasnitsyn
in
my
First,
unknown
On the basis of
understood) pattern.
rience and/or the results of
Two
separate
methodological
One
issues
was
the
forced
me
of
problems
to
the
approach
descriptive
biology.
long-lasting crisis of taxonomy
that
resulted in the continual conflicts between "traditional"
phenetic
an
and
taxonomy
and,
intuitive
Rasnitsyn,
1988b,
of
&
the
&
1987a,
Re-
1988).
are
arguments
chapters
devoted
to
regards
in that
ence.
it
is rooted in my
believe
I
menclature
are
ple
paleontological
the
basic
cation is
deficient in
to
classify
one
is often forced
rules of nomenclature. This is
in
paleontology,
or
An
escape
and
the
seated
ally accepted
nevertheless
the
objects
gives
sort
begins
rules. The
to
little
discern
not
their
1992b).
practice
in
a
stu-
taxonomie
satisfaction
more
deep-
followed the gener-
reason
why is that
are
taxonomically deficient
must
break the
rules of nomenclature.
why he has
renders these basic
1986;
appliof tax-
where
normal
violating
derlying principles of nomenclature
ible with the
to
common
stages of fungi,
of this
student
reasons
mitting
sim-
be classified.
This
the
incompat-
material that
controversy
principles discernible, thus
identification
and
un-
per-
study (Rasnitsyn,
In
permits
we
influence
found
otherwise
permit
of
structure
or
the
tirely
new
(I
a
the
cant
he
will
vide the taxon in
group
separately
with other
give preference
such
internal
features
creates
as
(except
and
an
en-
en-
also
must
of his
of that
a
In
to
proposal
not
the
In
this
taxon).
signifi-
how
colour
the
but
already
similarity
to compare the
addition he
it
or
must
many
quently allows
us
re-
will tend
to
over
patterns.
concerning
One
particular
under examination
patterns
that
the
previously
had been
question. The supposition ought
in
predict
classify each sub-
seemed to be dissimilar to the
mechanism
similarity,
or
subdi-
cognition begins here.
predictive, i.e., it should
particular
to
to
comes
morphological similarities
class of patterns
patterns in
when it
eventually
data).
hypothesis
a
studied
in
or
the
opinion
classification,
question and
similarity between
can
when
his
disregard features,
The second stage of
and
pro-
with,
that discriminate males and females,
per se,
adults and larvae
sults
a more
taxon, he
separate
an-
behaviour under
in
an
general taxonomie position
treatment
its
example,
classificatory aspects
only
mean
For
instate
to
as
group
deal with the
and
pattern
various circumstances.
wishes
or
In turn, the
appreciate and foresee,
to
us
our
corre-
way
correlated
are
the
that
is
(i.e.,
one
those that have
are
on,
the
to
context,
suppose
discern
to
us
this
unknown pattern
essentially
an
yet
expe-
these known
sort
otherwise reflects in
important features
study of immature parasitic
breaking
experience
until
the
of asexual
cannot
practice
formuso
areas
difficult
no-
taxonomically important characters (or
lacking them),
worms,
are
that their existence and
has
one
of
and
principles
perceptible only in peripheral
onomy. When
dent
experi-
principles
only partly understood
such. Some of these
as
and self-evident
even
different
Despite the long history of taxonomy and no-
menclature,
lated
is
cause
(i.e.,
previous
other) important features of the pattern.
tomologist
the
nomenclature,
then
important when
or
phylogeny and taxonomy.
As
considered
are
known
not
important (i.e., relevant
known elements of
suc-
preliminary confron-
a
we
unimportant.
as
lated with,
1983a,
my
or
knowledge
Dlussky,
develop
to
in
below
set),
goal
either
of traditional
controversy, al-
1972,
as
of it
(Ponomarenko
success
Rasnitsyn
the
-
elements
pattern,
Because of
ones.
superiority
Rasnitsyn,
sults of my
attempts
presented
a
much
1971;
1992a;
approaches
involved in this
was
without
though
rival
the cladistic
later,
feeling
I
taxonomy,
the
tation with the
ac-
four
seeking for
start
we
essentially
an
is
knowledge
opinion proceeds through
stages.
elements in
while
cognition,
to
that
is
clarify why
other
there
characters.
hypothesize
various environments and
for
responsible
be
is
a
the
also
a
This subse-
that the
similarity
the behaviour of the pattern in
tinue with the above
to
propose the existence of
question
conditions. To
example, the taxonomist
con-
will
Contributions
possibly
that the
pose
characters
could
pattern
that
pothesize
is
natural,
more
weight
much
tion of the
less
alternative
the
on
To
with
the
whose
to
of
details
formal
concerning
Karl
by
Moreover,
it is
result
arises
Farris,
is
by
your
black
swan
stained
always
of
a
giving
the
claim
sess
a
we
you
be
stage
of
consider
a
reply.
is
falsifying
due
to
claim that all
a
black
should
modified
setting
hypothesis
a
the verification
nor
is
swan,
reject
like
that),
black
a
and
skilful
your
possibilities
are
rather,
of
for
&
Rasnitsyn
justice
particular
in order
to
reach
is
ultimate verification and falsifica-
is
That is
a
why
is
vital,
necessary
it is
be
to
unless
is
accepted
how strong the
dangerous
generally
suspicion
of
a
between its
of
their
subgroups.
perception,
so
that
These results
of
concerning
consist
organisms,
and
attitude
too
commonly
not
group
of
relatedness
commonly
is perceptible
from the process
of traits
including
behavioural
of
understand
slow for imme-
phylogeny
from its ultimate
results,
no
described in
it is
addition,
proceeds
proved,
particular
genealogical
In
punish
may be.
history being
diminishing
that evolution
a
to
unpunished.
accusation is
history
evi-
considered to
is
the
no
only
that for the
to
as
to
presumption of
the above conclusions
living organisms,
function
tangible
study phylogeny. Phylogeny
can use
we
sound
criminal
a
not
the
de-
a
the alternative decision
more
leave
to
that
means
otherwise the safest.
the observation
on
criminal
interpreted
only
This
The well-known
suspect
terms
likely
or
reasonably
a
We
how
a
that is considered
decisions)
society
innocent than
matter
1988b;
there exists
problems
consequently
accepted.
health of our
be
decision
seeking
presumption.
most
innocence is based
the
a
against it. Otherwise
be
to
"correct"
class of
the
such decisions
has been sufficient. This "gener-
the absence of
dence
(or,
numerous
what tangible evidence for it exists, but
matter
itself.
a
the
class of
a
decision
diate
as-
(or
generally
in
nor
deci-
a
are
where information is often insuffi-
where
alization" is termed
be
they
(Rasnitsyn,
that the time needed for
cision
automati-
It has been found in
1988).
Dlussky,
given
a
but
make
we
time
long
a
but
for
everything
making
bad
not
cient
that
and
now)
and
are
mistakes,
of how
explanation
existed
courts
we
making
at
information
almost
act
we
us
be-
possible
enough
sufficient
to
to lead mankind to extinction.
physiological,
an
make
until
were
enough
reason
necessary
is
live and
we
insufficient
plain sailing,
do
We
agreed
decision.
Since both
sion).
just
appears
with
information
be
can
nothing definite
skilful
are
and do not admit that
cally
not
hypotheses
falsifying evidence
(e.g.,
(e.g., artificially
It is for this
cognition
to
possi-
(2)
likelihood and
competing
(cf.
prior
the
or
ne-
swans
not a swan
goose,
believe that these
1
we
suppose
to
how
unreliable? This is
so
decisions
(when
The
(circumstances)
swan
naturally
not
yes-or-no
relevant
pro-
can
rejected.
be
viz. that the
if you
something
or
matter of
fourth
when
is
black).
must
mean
chance
encounter
convergently
mystification,
supposition
falsify
to
influences
1983). Indeed,
a
of each
a
hypothesis definitively.
a
bilities that (1) your black
it
make
to
who clarified the fact that
possible
neither
white and yet
abandoning
is
has been well studied
cognition
hypothesis,
external
glected
are
not
I
cause
If so,
things.
seems
be certain of any
can never
and considerations concerning
world which offers
"generalization"
or
since this would
alternative
an
experi-
cognition
availability
verify
way to
completely,
of
the
Popper (1959),
no
and
discussion follow.
a
The third step of
there is
in
compare them
below).
further testing,
Examples and
the
implications,
to
a
and thus
has
assump-
propose
observations
whether any
posed hypothesis:
proposed
chain of
The fourth and last step of
retained for
cognitive
discuss these
not
should
we
opportunity
our
see
the
test
diverse
better
a
results
decision
cognition
of
nature
The
that end
most
have
(for
ments
more
we
observations
work in
here.
reach
and
the
impossible,
are
our
correct
personal experience and intui-
investigator,
third stage is
to
gives
of
stages
stages
order
homo-
tion
of
seems
is still obscure, and I shall
longest
that
one
hy-
classification
burdened with
in detail
tions.
species
taxonomist would
nature
The
the
(say,
one
above-mentioned
two
depend
generic)
the colour pattern.
to
The
morphological
inferred tentative
e.g.,
than the
plasies,
the
words,
the
in
5
while the similar col-
lower
a
1996
-
higher (say,
a
similarity,
concerns
In other
level).
(1)
similarity
indicate
level of the overall
our
66
Zoology,
to
of form and
morphological,
characters,
organism
to
biotic
those
and
A.P.
6
Rasnitsyn
abiotic factors, their distribution in space and time,
changes.
and the like.
amination, which,
The
first
and
It claims:
phylogeny.
sidered
as
"the
any
auxiliary
but
erally knowable. Indeed,
"if
dently gained similarity]
is
this
it
1983:
of
phylogeny
from
the
water
evident
not
can
hydrogen
not
be
an
obtained
the
different level of
in close
(Farris,
and
circumstances
engender
but
conditions,
either
by
two
burning
why living beings
their
on
while
history,
essentially
are
evidently
cases
complexity of chemi-
That is
dependence
in chemicals
changes
governed by
their his-
simple rules (not by
Living beings
are
acquired
independently
traditional methods
cerning
procaryote
Observations of this
strictions
on
of different
in
a
sort
help in dis1987).
quite evidently impose
taxa of
a
Generally speaking,
taxa.
place
Hence
it should leave clearer evidence of how such
has
vergence
feel
that the
especially
tionships
this
those
often
more
of,
say,
I
1986).
is
a
much
think.
similar
genera,
aware
or
organization
fashion
to
than rela-
that
(cf.
of
homo-
than students
concerns
highly
less
explanation
event
di-
species,
much
out
viz., the hypothesis
particularly
a
often
subfamilies
of the
common
we
of
are
to sort
families
am
and thus
almost identical
a
larger
difficult
more
This
closely related
in
in
phenomenon,
plasy
Nevertheless,
genealogical relationships
manifest and
Saether,
proceeded.
re-
analysis.
lower order takes
past than that of higher
remote
that
vast
(Zavarzin,
the domain of phylogenetic
divergence of
less
so
often of little
phylogeny
There is another restriction.
the
the amount of
is
similarity
are
It has
complexity.
that in procaryotes
suggested
groups of
similar forms whose
enables them
similar
to react
environmental
the
beyond
ex-
scope
tion
paleontological
in
resulting
sition
well
as
as
abundance of
an
often characteristic
from
the
contemporary
of
of
as
an exam-
parallel
organization
tran-
of
(grade)
amphibians (i.e., independent
groups of the characters
the level of
to
and mammals
ac-
char-
generally
and further
amphibians)
organization of reptiles
is
taxonomie lev-
serve
cases
evolu-
parallel
homoplasies,
higher
numerous
level
fishes to that of
quiring by
of both
Vertebrates may
species.
with
meticulously studied, both
it is evident that extensive
material,
(or birds).
In
the last two decades this old theme has been under
Russian students of evolution
close examination
by
(Tatarinov,
1987; Shishkin,
1992).
va,
born,
1976,
Under the
1934),
this
old
is
pattern
1987; Vorobye(Os-
aristogenesis
term
discussed by
Rautian
(1988).
Parallel
lar
described
tendencies
of
nomenon
the
1979,
lar albeit
not
understood
underlying
present
group,
the feature is
develop
ber of the group
acquisition
(the
of
to
its
a
resilient
has been the
general
strong
breathing,
pendent
cussed
by
capacity
in every
For
the
to
mem-
instance,
longitudinal
the
"string"
that
backbone)
to
common
obtain
swimmer
being
a
(whales,
we
they
cause
of
are
1979).
rarely
the
be-
ich-
dugongs,
case
is the
cos-
the
inde-
homoiothermy,
Iordansky (1977,
however,
independently
Another
remote
of
base for vari-
whenever
appearance
development
examples,
genetic
successor,
thyosaurs, tadpoles, etc.).
tal
many
the
group and
members of the
expressed
not
chordate animals
come
apomorphy
accumulate the mechanic energy of the
body bending,
fish-like
or
of simi-
that has been
function)
(cf. Sluys, 1989).
the
chord and
permits
ous
some
because
probably
group.
epiphe-
The notion of
an
ancestor of the
common
in
or
a
an
key apomorphy,
1977, 1979).
in structure
gained by the
as
a term
synapomorphy implies
(any change
that is
is
to
synapomorphy
1989;
meaning
proposed by Iordansky,
underlying
intrinsic
1986; Sluys,
identical
is
groups
manifestation of particu-
a
be
can
related
closely
as
tendencies
evolutionary
(Saether,
in
evolution
commonly
The
tory).
been
regard
group is
a
to
ple, for they show
lipids inside organisms.
between
living beings.
vs.
change
does
man
under any
ape,
equally
difference
A
at once.
by oxidizing
or
lies in the
When
is
charac-
lies
however,
of the
article.
present
els and
gen-
deserves close
problem certainly
phylo-
homoplasy [indepen-
universal,
The
acteristic of
and
man,
cals
is
14).
This is
The
con-
state-
far
imply nothing about the genealogy"
ters
of
reverse
is
dictum
central
is the claim that
genetic analysis
a
should be
( 1966) calls
principle",
The
phyloge-
"knowability"
inherited unless and until the
auxiliary.
being
of
similarity
reasonably proved. Hennig
ment
in
presumption
central
netic analysis is the postulation
taxonomy, and nomenclature
Phylogeny,
-
Unlike
aware
dis-
as
these
of
the
Contributions
concrete
that
causes
that
real
a
of
jority
results
cause
for
for
in
hence
-
a
re-
par-
I believe
underlies the
the
develop
to
the
apo-
"underlying
term
parallel evolution does
to think of taxa
vergence of their
tation of
related
could contribute
well,
as
Parallel
ancestor.
forms
if it
especially
(term proposed by Mamkaev,
1968).
of
can
parallel evolution
as
a
whole
combined with
changes
in the
anced (inadaptive
and
old
that
(1859)
(probably
the
com-
diversity"
The process
of
Rasnitsyn,
of
extant
due
large extent)
taxa
are
time. That is
long
in
of
their
evolution
have
to
seem
compared
as
earlier
an
The
note
how,
tree,
is
we
and
becomes
of
a
say,
can
for example
and
even
ancestor
scribed,
sils
to
albeit
a
at
never
pat-
proposed
are
important
seems
procaryotes.
a
of its
be
to
speak of
family .
Any-
phylogenetic
general validity
quite certain about
about its
1
the
phylogenetic study of
construe
sophisticated
the
ancestor of a class.
real
details,
ancestral
an
In
so
quite
a
that it
species
few cases,
research has failed
ordinal position
of the
And this may be
to
common
true even
if the
has
already
not
yet recognized among other fos-
(Vorobyeva,
little in the actual
Further
ranged
the
toward the
be
can
according
methods
of
but
phylogenetic study.
phylogenetic
analysis
clades,
change
presumptions
gross classes
two
respective
of
practice
phylogenetic
in
ar-
whether
to
directed
are
of either groups
history
or
2
1992).
been
In
Group analysis
The main method of group
tological
collected and de-
short, parallelisms
af-
one,
the
on
the
fossil
presented.
until sound
universal
not
record is
rather
is
and
stand
has
which
do
the
nor
pre-
fossil
this
chance,
to
special
a
paleonto-
fossilization
is
and
In
a
possible
to
a
instance,
fossil insects
and
environ-
this
in
informa-
infer with reasonable
case
or
of absence of
a
found
mostly
Paleozoic,
sources
became
the
not.
are
in the
deposits (other
relatively late stages of
most
to
Using
insects
information
later
of fossil
and
section of the fossil record
and, especially
low marine
mological
good
contain fossils than
to
non-existence
fossil
a
fossilized
deposits vary greatly
particular
particular
a true
lacustrine
a
have
excavated
of information.
often
and
what kinds
better chance
certainty whether
group from
organisms
be
it is al-
many cases,
buried
to
of organ-
considerable
a
those patterns (see e.g.,
to
Additionally,
degree
due
due
gathered
1990).
chance
not,
ones.
tion, it
why
is
and
becoming
stand
ments
For
vast
There
already
Crowther,
good
a
other
because
is
method
incomplete. However,
neither
of burial
it
of
chance
is
be
known what kinds of
ready
paleontological
paleontological
application
of data relevant
&
Briggs
their
to
regular.
patterns
isms,
ancestral
evidence
discipline called taphonomy that deals with
logical
the
in
known
incompleteness
but
earlier in
considered
contradictory
for groups. The
apparently
two
appearing
one
We shall call it the
sumption
as
It is based
1979, 1990).
should be
paleon-
redefined
recently
among
the
groups
record
unless and
is
that
presumption
related
closely
is the
analysis
which has been
stratophenetics (Gingerich,
amount
of
case
particularly
to
and
themselves
described above
assessment
pointless
thorough
identify
We
clear-cut
Here it is
1987b).
pattern
an
to
intermediates
still able
are
quite
later stages
history.
than in the
although
tree
of
(Rasnitsyn,
restricted.
the
or even
difficulties for the
eucaryotes
more
at
involved in explaining
extinction
less
cause
taxa,
in their
that the
taxa
easily discernible phylogeny
lower
stage
elsewhere
to
to
often
are
more
hypotheses
of
tern
a
part
extinction of in-
to
why higher
to
Darwin
termediate forms. This maturation often takes
a
as
1987b)
leads back
hypothesis
gaps between
a
is
elimination of unbal-
sense
well-known
to
adap-
be described
intermediate groups. This approach
the
but otherwise the above considerations
not
the accumulation of similar
maturation, i.e.,
taxon
unresolved
similar niches
to
reduction of the "archaic
a
with
dichotomous)
to
simple products of di-
as
common
closely
bined with
them
filling
opposed
as
.
Wide distribution of
us
by
cladograms
characters
synapomorphy".
permit
feet
(polytomous
ma-
evolution. This
parallel
tendency
a
the
by
yet unknown (but
suggestions)
of massive
character
morphic
7
feature
particular
a
cause, albeit
common
cases
1996
organisms. Nevertheless,
Sluys, 1989,
see
-
(1)
responsible
are
of
peated acquisition
ticular group of
66
Zoology,
to
of paleoento-
important only
group's history).
seem to
in
in shal-
be either
in
That is
flying
or
8
A.P.
aquatic/semiaquatic,
terrestrial, and
and lake
record
later
phalian
(1988)
be
to
of
are
based
fossils that
on
bristletails
from
taken
as
Pterygote
evidence
that
On
pre-Triassic
from the
that
ancestor
Triassic
method.
Groups
that
until
is,
hypo-
hymenopteran
a
before
the
methods that
rely
died
out
group in
a
evolving
are
on
distributional
a
way
terms
biogeographical
of its
to restore
distribution.
their distributional routes.
of
history
with its
benefit
may
the
all,
the aim of the method is
Indeed,
the
phylogenetic
first of
analysis,
correlate in
geny
in any
figure
from
group could
a
genealogy,
that
so
phylo-
There
biogeography.
are
problems, however.
The
classic,
considers
a
group
as
a
result of its individual
dispersal through various barriers. Major factors of
distribution
are
thus
supposed
structure of the space,
graphic
tential of the
group,
bution patterns
similar history,
of
be
to
the
(2)
(1) the
geo-
dispersal
po-
and (3) chance. Similar distri-
two
groups
could
imply their
including genealogy, depending
on
similarity of the three above factors. However,
seems
not
particularly
essentially
that the pattern
likely
similar,
is
similarity
the
biogeographical
doubtful value for
In
contrast,
phy (the
mary
the
that all three
so
Therefore, under the classic
phy,
etc.)
on a
proach
the
drift, but also of
ignores unique
that
pattern
the
frequented
gruent
of barriers
often due
to
chance.
method
appears
to
be of
phylogenetic analyses.
cladistic
by Humphries
of
factors
should suppose
concept of biogeogra-
concept
treatment here based
distribution
we
it
&
Parenti,
plants and
of
mostly
on
the
sum-
1986) considers the
animals
as
a
result,
both
climatic nature,
a
under
various
solely
The
dence of the
the
con-
with
the
as
the
of their
names
there
interpreted
are
on
that
groups
investigation.
replaced by
names
areas)
populated
in
repeats
areas
and relies
events
cladograms (the cladograms
area
animal/plant
evi-
respective area-dependent genealogy
of all groups concerned.
A cladistic
In
biogeography
of the
objectives
seeking
then
not
so
is known
be
to
on
an
existing
the
dispersal potential
but
not
in
chaotic
unique
may
be
to
as
area
easily
that could be
could
demonstrate
would
a
from
as
concern
fected
disperI
by extinction
For
example,
there
a
are
And
numerous
some
record,
enough
a
if it
method.
cladistic
Humphries
are
recon-
distribution
&
of
Parenti,
no
exists, quite
time in the past
to
af-
easily
pattern
the cladistic
at-
not
are
cladograms
that most of them
fossil
at
yet they
area
produce
see
as
(dis-
little
pay
("Gondwanian")
review,
I suspect
shows that
events.
can
of austral
because the
tool. Both classic
interprétable by
patterns (for
be cautious
biogeographical
biogeography
entire biota. The
wrongly
to
the
but often consistent
common
but
tral
for
biogeographical
reason
phylogenetic
a
the
structions
the
of
prospects
tention to extinction
very
research
cladogram patterns,
using
and cladistic
persalist)
in
interpreted
Unless
subordinate role
area
events
cladograms
restoring genealogy.
the
method
of area
so
producing
as
deceivingly
There is another serious
regards
out
on
vary
necessarily
not
biogeography.
of the
origin
refrain
method in
but
easily
of cladistic
and
Regular dispersal
produce repeating patterns
terms
only
is
of
de-
and,
of barriers
sorted
cladograms.
involved.
does
group (which
a
disorderly way),
a
area
Dispersion
common event
a
system
of
group
however.
easy,
of
hypothesize
simply
for each
cladograms
is
matter
organisms
should
the
ideally
of groups.
phylogenetic analysis
we
isomorphic
The
fit
to
seems
particular repeating pattern
a
cladograms,
1986).
biogeogra-
imposed
connected
(primarily
agents
continuous ancestral distribution. The ap-
sion in the
dispersalist biogeography
distribution of
will be
physical
with continental
pending
history of
Thus
absence
various
by
and nomenclature
taxonomy,
phylogenetic and geographic,
hymenop-
and their
should avoid
we
the
.
group
the
be
Phytogeny,
-
3
There exist other
on
seriously
not
presumption;
groups that
among
could
searching for
require
be
record should
strong evidence,
have
theses
al.
Carboniferous
cannot
fliers,
et
Absence
certainty].
they
the above
analysis using
de-
fragmentary
the other hand,
good
are
West-
at
findings
too
are
Lower
record
ancestors.
insects
terous
the
fossil
[that is,
sea
fossil
and Labandeira
(1984)
identified with sufficient
(Mississipian)
we
al.
et
along
entered the
older
time;
Namurian
The
wingless,
are
common
The Thysanura
Shear
by
s.l.)
than the Pterygota
vs.
scribed
especially
not
shores.
forms.
shore-dwelling
bristle-tailed insects (Thysanura
Rasnitsyn
use,
often
presumed
aus-
occurred in northern continents and begroups
came
extinct
there
(for details,
see
Eskov,
1987,
Contributions
66
Zoology,
to
(1)
is often less
1992). Austral paleontology
than the boreal one, and yet there
tral fossils known for groups that
a
boreal
strictly
for
stead of
being
(Humphries
likely
be
to
of
of
remnants
butions that have become
that
biota
probably
Zherikhin,
1978,
is
paleontology
austral
there
the
the
ones
1971;
known relevant and
only
Late
from
Burma),
vealed
stead,
are
Cretaceous
now
under
among
several
the
fossils
groups
rich
specimens
amber
Paleogene
has
forms.
tropical
common
in
rikhin,
of social
of
functioning
functional
did
they
It
follows
tiated
by
that
forests,
meaningless
was
these
and
are
in the
possible
nowadays,
nor
area
record.
still have much
formulate
to
of
pattern
unless it is substan-
complete fossil
we
fore it will be possible
a
That
do be-
to
workable
for
presumptions
biogeographical
As
ants.
1993),
repeated
the
is why I believe that
of
com-
past.
reasonably
a
is
play key roles
known
not
are
in the
is still
cladograms
rain
tropical
social
(1978,
groups that
analogues
occur
the
(Zhe-
Apennines
assemblage
higher
Zherikhin
by
the three animal
those from
as
isopteran family Termitidae,
and
bees,
demonstrated
well
as
Burmese
devoid of the
pletely
and
the
1993),
Unlike the tropi-
know it.
we
and northern
Miocene of Sicily
the
set
analysis
of taxonomie groups.
The
same
holds
evant
methods,
which
suggests
as
In-
vice
assemblages
as
from the later Paleogene Mexican
and Dominican ambers,
re-
there
rain forest
tropical
cal assemblages
fossil
buried in
already identified,
very
The
1987).
investigation,
my
characteristically
no
with the
comparable
earlier
or
that
Late-
equatorial
(Meyen,
200
the
of importance
comparatively
arthropod assemblage (ca.
the
tropical
than
developed
is
of the present
distri-
tropical
equatorial
Cretaceous non-marine biota
tropical
the
Unfortunately,
it
more
(Razumovsky,
sound indications of
no
are
zone
poorly
Nevertheless,
one.
are
1986),
world-wide
past
1993).
in-
distributions,
separated by
Eocene
more
of
hypothetical Pacifica
a
in
originated
during
responsible
distribution
Parenti,
&
references).
indeed
(bipolar)
heritage
a
continent
aus-
have
now seem to
amphitropical
These
taxa.
complete
finds of
are
(same
could be
amphitropical
the
many
distribution
extinction
Similarly,
9
1996
-
approach,
cladogram
of the host
of the
use
depending
versa,
an
specialized parasite,
which
on
Brooks &
promising (see
more
rel-
apparently
other
parasitological
model for that of the
a
for
the
e.g.,
the
for
true
or
looks
cladogram
McLennan,
1991,
overview).
from the Late-Cretaceous fossil resins of North Siberia
and
Canada,
1901
and
and
least
at
present
hymenopteran
latter genus is
there
bipolar). Additionally,
Scudder,
sonotus
distribution
known
(Cape
territories
Eocene,
as
the
Baltic
In
the
(the
more
contrast,
amber
suppositions
even
in
specimens
forest
stronger
that
differed
has
is
of
a
used
either
to
(morphocline),
unique origin
of
methods for
eral
(1)
The
polarize
the
falsify
to
or
similarity
(stratophenetics).
elevated
the
Late
witness,
pendent
fundamentally
the
the
fossil-
from
the
of
one
but
a
These
two
to
various contemporary
to
spermous
ples.
why
that
a
similar
a
The
groups.
many
are
are
com-
direction in
of
mam-
(Tatarinov,
the
gymno-
good
we can use
transformation
that
inde-
are
cases
reptiles
1991)
cases
to
analysis
only fairly
similar
angiospermization of
in
sev-
Homoplasies
be not
have
plants (Krassilov,
That is
sumption
and
are
group
applications
malization of the theromorph
1985)
for
however.
shown
also often
There
method is
method
another,
paleontologically
mon,
as
-
transformation series:
polarizing
paleontological
Cuban
lowlands
evidence that
yielded
transformation series.
Polarizing
1976,
some
amber
analysis
transformation series
described above
extended far into the northern extra-
Burmese
Character
South
(New
Cuba
now
Africa
South
because the
and
tropical
is
Phryssonotus
and
analysis
bipolar (Phrys-
zone.
There is
iferous
a
Australia
zonally,
subtropical).
Phryssonotus
past
1954).
Chili,
equivocal
are
or
northern (snake-
millipede) extratropical
a
South
Uruguay,
island
a
Mediterranean,
the
Province),
record is
tropical
1885,
(Condé,
from
Wales),
of the
Raphidiida),
fossil groups
are
Character
extant
predominantly extratropical
that have survived with either
flies, insect order
genera
Palaeomymar Meunier,
the
1980;
(Rasnitsyn,
at
the
e.g.,
1937 and
Serphites Brues,
series
exam-
the
pre-
should be
10
A. P.
according
polarized
the succession
to
in
character states
spective
the
less and until sound contrary data
This
will
presumption
that
groups
with
record
logical
well
are
teeming
or
ontogenetical
sumption:
fact,
Baer's
der
that
claiming
dem
"Aus
sive
das
bis endlich das
Speciellste
until the
morphies
of the terminal
There is
law" has
in
I do
any of its
secondarily
phosis,
it
or
not
(e.g.,
can
get
may
phylogenesis
of
It has
leading.
with other
rule but
be
no
as
a
1978;
and does
succession of
as
the
well
as
system
way
or
in
can
of the
not
biogenetic
often be
in
of
as
organisms.
some
another,
the
the
Indeed,
holds
fossils
hard
onto-
is
a re-
true
and
for
their
states
Each of these three
traces and thus
reflects,
process of descent of
Each of them deserves
mis-
a
for the pattern of character
of
the
comparison
work
same
in
or
deciphering
developmental stages
succession
is
paedomor-
to
the
result,
presumption.
a
stratigraphie
beings.
due
groups,
ar-
Ontogenesis
priori advantage
a
sets of data leaves
one
"direct
a
(streamlining)
sult of direct observation. The
over
"biogenetic
possibly modified
lost
effective
some
methods,
only
As
highly
so
[synapo-
Nelson,
believe this.
stages
path.
while
method,
traits,
ones
taxa] appear".
of rationalization
developmental
the
special
relationships
to
be used
a
ample
as
in
according
because
know the
we
from
cladogram
is
of the
we
from
patterns of various character states, par-
ticularly from the
structure of
ourselves
basing
DNA,
with
symplesiomorphic
se,
per
features
the
to
respect
while those of the
below).
see
ontogeny
on
could infer that the human skull
monkeys,
ex-
anthropoid
(primarily
sources
Otherwise,
we
(1984)
is instructive:
paedomorphic solely
topology
other
results
merits,
own
Bonde's
paedomorphosis
infer that the human skull
the
their
to
priori principle.
are
and
gibbons
great apes could be
synapomorphic.
The
(3)
next
important method
the method and
are
transformation
series
unless
group,
is
polarized
is
wing
lieve it
the
a
evolution of
However,
to
wings
ences
between
without
(not
insect
the
leg
cf.
disagrees. Likewise,
lian
lineages
body size
1904).
of
("law
contrary
wing
on
the
efficiency
presumption
character
states
it
1981)
be
most
one
the
that
(1994)
mamma-
increasing
growth", Cope,
same
no
holds
sound
true
for
laws of evolution.
method
formalized
of functional
the
not
argues
but Bitsch
that
polarizing
can
dis-
is
infer this direction of
empirical
well-known
functional
important
in mammals where
evidence is found. The
The
differ-
both modified
are
leg,
know
should
we
many other so-called
(4)
the
Nevertheless,
phylogenetic
everywhere
well.
as
great
evolved in the direction of
Therefore
evolution
wings
hypothesis concerning
parts of a biramous ancestral
we
be-
prevailed during
Rasnitsyn,
insect
are
that the
Kukalova-Peck (1978)
reason:
and the
group,
legitimately
and bird
groups and
wings.
absurd.
completely
can
a
the
for another
sure
is less evident because of
of their
similarity
we
similar
a
a
If
contrary data
of what
pterosaur
pose
insect
leg,
example
an
in
reasonably
we are
modified
be
to
presumption
valid
as
and until sound
if
and
presumption of analogy:
results should be considered
bat
inclu-
pre-adult adaptation (caenogenesis),
the process
genetic
less
the
phylogenetics
an
of human
[that is,
most
belief that the
widespread
a
1984).
flexible,
of the
extraordinary importance (as
gument")
Bonde,
special
most
after
not
and
inferences,
phylogenetic
should be evaluated
adduced. Indeed,
of the less inclusive taxa], and
[synapomorphies
on,
develop
for
taxonomy, and nomenclature
Phytogeny,
transla-
general relationships
most
AR],
-
weniger
italics). Or, in
from synapomorphies
taxa
through
Allgemeinsten
sich
,
Von Baer's
auftritt" (p. 224;
In
rewords the famous Von
bildet
allgemeine und so fort
beginning
polar-
decide otherwise.
to
reasons
Formverhältnisse
tion: "From the
the
pre-
succession
ontogenetic
"law" of the differentiation
(1828)
is
biogenetic
series should be
with the
presumption just
ontogeny,
method
the
on
character states, unless and until
serious
are
this
characters,
rial
-
taxonomie
similar
very
Transformation
the respective
there
paleonto-
morphology
and thus poor in
It is based
one.
ized in agreement
of
in
homoplasies).
Another and
(2)
useful
in the
inconclusive
re-
un-
presented.
are
particularly
represented
an
(either by being simple
with
be
of the
fossil record,
Rasnitsyn
as
using
based
efficiency: Of two
corresponding
to
a
more
efficient adaptation should be considered apomorphic,
ing
until and unless there is reliable contradict-
evidence.
considered
as
because it
living
ity
mate-
than the latter
For
example, viviparity
apomorphic
secures
the
in
is
comparison
safety of
(it permits viviparous
generally
to
ovipar-
progeny better
animals to
sur-
Contributions
to
vive
lower
despite
of
a
Subordinate
(5)
complexity:
wasteful of
is
the above
That is
should be considered
simple
a more
tures often tend
cient in the
primitive
indicating
viz.,
character
state
showing
state
is in fact
The last and
is the outgroup
for
gent, and
make
model,
state
only
there
are
that
could infer
This
only
the
phic,
state
we
tionary
other
is
one
a
can
ap-
see
a
group and
infer
(unless
that the first
the
should
changes
its
is
imply
-
at
least
limits. If
more
independently
hypothesis
we
is
the
we
less
should
unless
within
found only
apomorphic
and outside
within
a
in respect
the
evidence
contrary
are
the
to
apomor-
analysis
til"
ties that
we
acquired
first
char-
an-
that the inher-
all
than
other
conclude that
the
likely (or less parsimoni-
similarity
encounter
contradictory
similarities
we
cannot
and
un-
similariway that
be all
con-
con-
accept the similarity of
their
treat
we
is inher-
them. The distribution is
must
we
the
valid,
we
a
If
homoplasy.
other mammals in their
to
as
when
inherited: if
as
inherited,
tions
only
when the
tradictory
and
as
was
discrimination be-
until the "unless
necessary
analyse
to
analysis
similarity
pursuing
distributed in
are
ought
as
until...")
It is
applies.
The
-
presumption ("similarity
and
not
homoiothermy
as
swimming adaptaof
independently
fishes,
and vice
versa.
There
The
plasy: (a)
in
turn
one
main methods
two
are
implication
if
ought
necessity" (Farris,
homoplasies,
likely ,
4
But
even
a
law
a
many
because characters
similarity
each
8),
a
homoplasy,
postulated
and
between
are
and
the
then
beyond
most
likely
the least number
are
cladogram
presumption
as
first presump-
our
why again "unless and
parsimony only
or
as
be
implicating
another
homo-
which is
parsimony,
of
unless and until there
indicating
sons
to
detect
to
usually inherited
appears
not
1983:
is that
cladogram
is
similarity
commonly
"homoplasy
of
presumption
more
less
group, and
then,
is
only
so,
that took
identical evolu-
agree
one,
ited
we
doing
the
sub-
outgroup presumption,
state
similarity
first
our
Indeed,
likely (more common)
acquired
being equal,
two
within
one
inherited
consider
tion.
subclade of the group in
hypothesizing
reptiles, cf.
plesiomorphic.
analysis.
second
apomorphic and the second plesiomor-
beyond
second
when
of
known
are
typically
are
that both hairs and feathers
characterized above
whales
which
and
of the
by
re-
1971,
application of this presumption
and scales
a
arguments)
The
potheses
in
persist
a
cases
should be considered
appears.
which is found
one
should
because,
group
sidered
we
1987).
application
to
a
to
These
Sharov,
related
some
which claims: A character
acquired by
evolutionary change
origin of
similarity
things
state
plesiomorphic,
cf.
of phylogeny,
both within
we
to
as
genealogy:
chance
and another
question. Otherwise,
acter
a
diver-
abundant
group. When
group,
is
state
apomorphic.
at
same
strong contrary
character
place
has
test
basic
our
generally
so
knowability
represented
beyond its limits,
within
with
about
character
a
Kurzanov,
in
(and again
in all
some
homology
feathers
proved). Similarly,
in birds
Similarity
method
probabilistic
descendants than that
later in the
character
a
be
to
tween
is
not
are
of
ancestor
wider range of
peared
It is
only
phic
took for
we
hairs described
pterosaurian
is still
the
1976;
in
(and
the lat-
over
popular polarizing
nothing
say
The
modi-
strong contrary evi-
cladogram
a
pre-
vestiges:
apomorphic
Tatarinov,
known
are
above transformation series should result in the hy-
Anlage).
an
most
of
polarizing
of
the
reptiles:
scales
mammals
unless and until strong
group,
com-
of their lost
signs
lated
in
only
of that distributed both
hoof).
found that what
presumption
remote
more
the
is
homoplasies
In this
6).
effi-
hair
amniotes,
jected
struc-
more
signs of being
model (evolution
similarity
the first
to
have strong
structures
more
any
comparison.
conformity
phylogenetic
ited
other
presumption
until it is
(e.g.,
vestige
(7)
p.
or
one
unless and until there is
dence
the
the
another
fied from
ter,
we
complex
simplified
rudiments
sumption,
structure
of evolution (compare e.g., the
secondarily
complexity,
if it
in relation
therian paw and the horse's
monly display
more
Keratin
below).
see
ous;
is that
is
complex
become simpler and
to
course
Such
(6)
why
unless and until
one,
offspring).
originates only
apomorphic
as
11
device
decide otherwise. Indeed,
to
reason
1996
presumption
and thus
resources
-
complex
more
a
(I)
number of
mean
to
efficient.
more
66
Zoology,
and
as
the
until?"
not a
most
Why
equal.
If
we
is
hard rule,
students believe? It is
not
of
sound rea-
so
consider
mimicking butterflies and count
similar element in their colour pattern
sepa-
12
A.P.
rately, and then
icking species
will
result
we
be
always
relatedness.
count similarities
its relative, I
to
in
similarity
the
genetically than,
surface
the
here, for
even
the
and
procaryote
colour
pattern
hairs,
of differences
importance
plant
is
in
subsidiary phylogenetic
not
varies
similarity
Nevertheless,
considered
I
Therefore,
only of
as
Hard rules
plasy.
the situation,
we
have
are
quite
have
(Rasnitsyn,
However,
the
flicting
sidered
as
as
other
(when
inherited)
inherited
cases
groups■), unless
I
use
homo-
describe
to
That is
In
fact
why
the
there
and
&
some
are
not
Dlussky,
needed
to
only safely
can
In
1988).
the
clarify
propose
of
case
all similarities
known
(especially
to
in
be
more
closely
con-
can
those which should be
are
of
discussed elsewhere
definition:
similarities
accepted
in
detail,
generalized
a
by
presumptions,
efforts
so
of
taxa
presumption: (b)
more
Rasnitsyn
more
many
in
impossible
been
already
1988b;
problem
here
one
sort
discriminating
weighted similarity.
number of such
a
of them
on
of
The
be
con-
reliable
related
and until strong contra-arguments
for
this
using
in
given
are
tera
the
Triassic
been
the
to
(Fig.
both
illustrated
with
the above presumptions
examples.
I
think
evolution of the hymenopterous
ticularly
of the
the
problem
of the
phylogenetic
sawfly family Xyelidae,
since it is
a
difficult
case
that
insects,
seems
and the
is better
the
and
early
par-
position
appropriate,
subject
of
long-
(order Vespida;
1988a,
dozen
c,
the
1989)
sphere
as
family
relict
a
Cretaceous
did it
menopterous
and
survives
belong
antennal
one
5
not
genera
and
until the mid-
of the
least in
at
structure
in the northern Hemi-
but
species,
-
These all
comprising
become
groups
and
species)
indicated by their char-
as
group
three dozen
have
species
(one
species).
venation
wing
Triassic
Australia
leading
more
hy-
temperate
climates.
Other
the
hymenopterous
Xyelidae and
Jurassic.
This
groups
it
makes
possible
hypothesis
a
all other
to
that
that show this
family
within the order
only
xyelid
most
venation
number (the
highest
most
all its
hypothesis
plesiomorphic
is
the
richest,
Indeed,
the
having
the
of veins
complete set)
most
and cells, and resembles that of other insects
the
fore
with
wing
branches in
a
RS
its
way similar
to
with
characters,
discussed below.
exception
wing
be the
to
pro-
of the other observations
regards
as
the
to
be ancestral
insects. This
hymenopterous
is in agreement with
the
Xyelidae could
Early
apply
to
for groups and
paleontological presumption
pose
appeared later than
recorded before the
not
are
diverging
most:
into
that of various
two
primi-
tive insects and with supernumerary SC veinlets in
some
fossil
base. The
species,
same
and
supposed
Particularly
morphies
so
that
xyelid
we
disregarded
The
have
to
as
xyelid
apomorphic
in
the
regards
is
the
respect
(Figs.
to
1980).
the
character
a
2,
mor-
feathat
staminate
the
unimportant,
autapomorphy
antenna
suggestion
(Rasnitsyn,
scrutinize
body
bionomic
from
retained
only
with free Cu
the
gymnosperm
cladistically
antenna, the
wing
groundplan
inside
possibly
are
as
relevant
development
was
and the hind
holds true
holometabolic insects
use
1988a &
1983b,
1982,
Rasnitsyn,
(two
1). The
cones
can
the
previous relevant
instead of Hymenop-
name
family Xyelidae,
acteristic
their
Examples
in
Central Asia
tures.
we
1980,
period. These
found
phology
appear.
How
my
insects
hymenopterous
reasons
involved.
should operate
to be
phylo-
what
are
least for the present.
at
rely
to
presumption
same
on
characters
when
seem
the
1969,
(Rasnitsyn,
on
1990; and references therein).
about
depending
we
characters
weighted
and
one
what
believe,
a
importance.
greatly
and
compared
are
of
importance
for
sets
The above observations indicate that the
genetic
exist
to mention
pigment
taxonomy.
generally
details of
or
Unless stated otherwise,
disagreement.
papers
and nomenclature
taxonomy,
first appear in the fossil record in the second halfof
similar in absence
are
from their
pigments
of blue
gross
exceptions
However,
all mammals
the
important phylo-
more
colour pattern
a
say,
structure.
as
in
or
assess
Phytogeny,
discussion below is based
what kind
on
generally
type of symmetry
of basic organs
structure
We
true
how reliably
depending
is involved.
state
lasting
that the
sure
favour of
in
do vary
they indicate relatedness,
of character
in
pleading
Similarities
of each mim-
am not
-
Rasnitsyn
earliest
Plesio-
however,
nature of the
not
easily
to be
of the family. 5
3a)
is
evidently
normal insect antenna,
Contributions
1.
Fig.
to
Triassoxyela
Middle
Xyelidae);
66
Zoology,
(1)
1996
-
1964
foveolata Rasnitsyn,
Late
or
Triassic
of
13
(Vespida,
Ras-
Kirghizia (after
Fig.
2.
Anthoxyela turgensis Rasnitsyn,
Cretaceous
of Transbaikalia
1990
Lower
(Xyelidae);
(after Rasnitsyn,
1990).
nitsyn, 1969).
with
several
long
and
basiflagellar
thick
of normal (thin
apomorphic
and short)
The
sumptions.
is
groups
of
paleontological
non-hymenopteran
=
The
sect
antennae
The
same
is
holds
the
the
at
earlier
the antennal
segment
same
found
is
one
conclusion,
only
Some of the
the
same
seems
son,
to
be
living
especially
the
xyeloid
of
and
insects the
occurs
some
horntails)
presump-
more
stage
homo-
Equally important
because
the
simi3rd
is
certainly
presumption
suggests
3f)
xyelid
antenna
while the
type is
normal
in
This
Indeed,
in
that
this
type
is
not
inference is
is
antenna
family,
be described
It
wide-
the
case
probably
although among
hymenopterous
can
3. Antennae
stated
of various
otherwise,
of
(Blasticotomidae);
Caenolyda
gidae); d,
liidae);
e,
Xiphydria
f, primitive
1982).
Arge
c,
ustulata
reticulata
camelus
I—XII
=
after
(Linnaeus,
of
segments
antenna,
(unless
1818
Malaise,
(Linnaeus, 1758) (Ar-
(Linnaeus,
Rasnitsyn,
and flies
Brébisson,
filiceti pacifica
1767)
(Pamphi-
1767) (Xiphydriidae);
fly Glutops semiformis Nagatomi
(Rhagionidae) (a—e,
antennal
Xyela julii
a,
Blasticotoma
pupa); b,
(Xyelidae,
sawflies, horntails,
imago):
1968; f,
“III”
=
&
Saigusa,
after
1970
Nagatomi,
compound
“third”
segment.
outgroup compari-
antenna
order, and
xyeloid
yield
to
seem
applied intraordinally.
true
Fig.
1931
Vespida
normal
one.
only
lower
is
above presumptions
erroneous, however.
ance
in-
both within and outside this order.
within the
with the
pre-Triassic
developmental
(Fig.
for the
result when also
because
spread
flies
within the
common
anten-
for characters is
evidently compound)
not
The outgroup
apomorphic.
had normal
segmentation
of
and oldest
xyelid apomorphy.
than in the adult insect.
antennal
for
presumption
biogenetic
presumption of analogy,
larly inflated (though
the
the
The
extinct order Palaeo-
display
for
true
seg-
thread
a
is supported
closest
presumption
not
a
the above pre-
because all known
do
because
nomous
(the
group
respective
applicable,
(Fig. 3a)
the
Miomoptera, Fig. 4)
also
tion,
an antenna
relevant because
manteida
with
apically
number of
a
into
antennal
flagellar segments.
of such
nature
by the application
nae.
"third"
The latter is mounted
ment.
fused
segments
compound
in its full
also the
insects
as
the
appear-
antennae
(sawflies
variously
modi-
fied
are
xyeloid
the
antennae.
antennae
of
The least modified of them
the
tenthredinoid
Blasticotomidae and
Argidae.
the
even
"third"
pared
to
segment
that of the
following segments
some
(Figs.
hypertrophied
as
Xyelidae themselves,
are
Blasticotomidae),
3b-c).
families
Both of them have
reduced
or
to
became
a
single
com-
but
the
one
completely
(in
lost
A. P.
14
-
Rasnitsyn
long
sawflies and horntails (lower hymenopter-
to
insects
ous
enough
In
type possible.
long
the
to
4. Palaeomantina pentamera
order
tiscidae,
Lower
Palaeomanteida);
1977
Rasnitsyn,
Permian
(Palaeomanof Ural
have
Xyelidae
the "third" segment not
so
but
and
still
clearly thicker
distal segments. The
Another
of evidence is
source
sawflies
living pamphiliid
Pamphiliidae (except
provided by the
and siricoid
have the "third" segment subdivided but it is
paratively
fourth
and
of
primitive
1871
three
to
less
no
horntail
distal
more
malesof B.
is the
important,
4
1871
grandis Philippi,
Philippi,
that has
primary
of
antenna
with
phies
in
were
also have this
larged (Fig. 3e),
part
of the
The
distribution
segment
can
pendent development
number doubles,
reported by
Van
tables 1, 2,
ite
segment
this character
somewhat
is
state
en-
possibly
a
the
we
& Van
its
by
in five different
Achterberg
of the
compound
explained
however, when
fig. 366)
structure
be
of
3rd
inde-
groups. That
consider data
Aartsen
(1986:
and suppose that the
composwas
segment
never
lost (cf.
Fig.
1968
thus
cases
of
an
independent
we
more
of the
numerous
take fossils into consideration. The fossil col-
lections
of the
Academy
of
contain
rassic
an
and
Paleontological
Sciences,
demonstrative
sects
acquisition
even
in
respect.
overwhelmingly
Lower
collected all
hailing
this
from
Among them,
collections
number of Ju-
hymenopterous
Siberia,
specimens
Russian
particularly
the world, though
Kazakhstan,
515
are
These
large
Cretaceous
over
Institute,
Moscow,
and
so
their
sider
in-
mostly
Mongolia.
of the above age be-
is
and
for the
forming
No
(Fig.
the
form and
appearance and
that of the
thick
I
Xyelocerus
1990 and
Rasnitsyn,
extinct
an
in
con-
family
recorded
are
and
genera with
ovipositor
position
in the
wing venation
Mesolyda
pomorphies
other
1968,
xyeloid
syn-
characteristic
body.
The
also
general
similar
to
these fos-
Prolyda Rasnitsyn,
family Xyelydidae
1963
are
assigned
which lacks
supposed
Pamphilioidea.
a
Among
Rasnitsyn,
and hence is
of
are
living pamphilioids.
the extinct
the
important
superfamily Pamphilioidea, viz.,
sils, Xyelyda Rasnitsyn,
the
is
(supposing
1968,
show the
6)
the reduced claw-like
to
which
autapomorphies
extinct
five
antenna
apomorphy of
to
of
also
of the other Tenthredinoidea.
next
type of
1968,
genera
Pterygophoridae).
Rasnitsyn,
as
and
rest
family, which implies its ancestral position
respect
general
SC
secondarily multisegmented
Dahurotoma
1968,
Xyelotomidae.
four
flagellum,
some
Xyelotoma
Pseudoxyela
The
forms
Rasnitsyn,
free
number of segments
antenna
Rasnitsyn,
the
retaining
distally,
present,
with the Tenthredinoidea as
antennal
reduced
Diprionidae
when
(Pseudoxyela
clade).
synapomorphies
a
that the
forms the sister group of the
probably
with
in
branch,
in
exceptional
tenthredinoid
The
compound segment become
if
is
regards
11).
The
that its fore
intercostal cross-vein
an
that the
family ground plan.
above
antennal
so
other
so
RS
lost. Most
genera
R,
share
Xiphydriidae
enlarged
got
have the SC stock fused, albeit only
segments
all
or
with
and addi-
not
cell
mcu
also
tion,
if
1
These
meeting
rather
(specimens
Most
wing
the
synapomor-
(Fig. 5).
with 2r-rs
first abscissa of RS is short
the
D.C.).
fore
an-
so.
four genera
firstly,
Tenthredinoidea
distally of 2r-m, and
studied in the collection of the Smithsonian Institu-
Washington,
the
than the
have
displaying
important putative
living
genera have
fossils
type there are,
23
with
Xyelidae
nearly
length
a
in
segment (subequal
into
Less
3d).
segments combined),
subdivided
tionally
(Fig.
genus Brachyxiphus
"third"
long
width
the
those
longer
or
type
be-
fossils,
55
32 fossils
remaining
non-xyelid
antenna
antennal
corresponding
in the
as
much
clearly
specimens
the
large
of the normal
the
Among
xyeloid
are
that share several
com-
than
longer
Xiphydriidae)
(Siricoidea,
thick and
times
subequal
known, though
the
several
small,
tennae that
horntails.
species of Pamphilius)
some
have
xyeloid antennae, including
(after
1977).
Rasnitsyn,
also
and
type. Among the remaining
antennal
Fig.
identification of the
turn, 65 of these
Of
Siricina).
their antennal structure
display
render the
to
the suborder
comprising
120
these,
taxonomy, and nomenclature
Phylogeny,
to
auta-
be ancestral
Praesirex
Rasnitsyn,
Contributions
Fig.
1968
Kazakhstan);
Cretaceous
on
after
of
1968; b,
robusta
Rasnitsyn,
after
Transbaikalia (a—b
and
1969, fig. 82,
Rasnitsyn,
heteroclita
Jurassic
Upper
of
South
1990,
Lower
1969;
Rasnitsyn,
PIN
specimen
a,
ni-
Xyelotoma
Pseudoxyela
c,
the
Dahurotoma
d,
15
1996
-
sawfly family Xyelotomidae:
1968;
from
(all
(I)
Rasnitsyn,
Rasnitsyn,
Rasnitsyn,
d,
of the
admirandus
gricornis
66
Zoology,
Representatives
5.
Xyelocerus
based
to
c,
2997/650;
1990).
Rasnitsyn,
Fig.
6.
and
Praesiricidae
b,
Prolyda
Rasnitsyn,
stan); d,
1968
and
Turgidontes
with
synapomorphic
dontidae in
signed
was
extinct
probably ancestral
One
more
assigned
group
several
comprises
syn,
the
having wings
the
to
the
to
1993).
genera
very
narrow
narrow
(except
apical
xyeloid
or
are
wings
and
SC branch
antennae
are
(secondarily
with the SC fused with
the
and
costal
pomorphies of the living Cephidae
the
cephid
themselves,
(Early
1968,
the
ancestors.
as
As
to
lacking
space.
and
having
genus
xyeloid
were
the
observation of their
Cretaceous)
Both
1988c).
with the costal
space
only basally),
crossing
(all
from the
Praesirex hirtus
Rasnitsyn,
1990
holotype;
e,
after
families
Xyelydidae (a—c)
excellens
1968;
Upper
Rasnitsyn,
Rasnitsyn,
1968;
Mesolyda depressa
c,
Jurassic
of South Kazakh-
1968;
Turgidontes
e,
mag-
from the Lower Cretaceous of Trans-
(both
after
baikalia) (a, b, d,
1983b;
Rasnitsyn,
c,
original,
based
on
1990).
Rasnitsyn,
Mesocephus
antennae
the
The last group with
The
loosely constructed.
nitsyn,
1969
horntails
the
presence
represent
a
of
Protosirex
with
represents
cidae. Other siricoids
Syntexyela
media
prob-
Rasnitsyn,
1968
ancient
Rasnitsyn,
(Rasnitsyn,
siricoid
Siricoidea,
synapomorphy
In
particular
rather
to
belong
of its
plesiomorphies
a
to
both
extinct
with
a
Rasnitsyn,
the
ap-
(e.g.,
it
siricoids
few other Jurassic
the
to
general
may
and
fossils,
family Gigasiri-
xyeloid
1968,
antenna
and
S.
are
inversa
(Fig. 9a-b). They surely belong
witnessed
as
(a
they
Mesozoic family
is
SC). Alternatively,
ancestral
group
antenna
seems
because
free
a
erroneous.
xyeloid
which
some
cephoids. Along
the
a
(Siricoidea)
pearance and
be
to
I include here Protosirex Ras-
(Fig. 8)
syna-
Cephidae
most
1988c) has been found
now
synapomorphy
Sepulcidae lack the characteristic venational
ably
as-
in the
super-
placed
their
band-like
basally lost),
sometimes
are
family Sepulcidae (Rasnit-
living Cephidae (Rasnitsyn,
moderately
R
They
7) which
(Fig.
Sepulcidae
families have their fore
Megalo-
family
nus
Rasnitsyn,
karatavica
1968
sawfly
Xyelyda
a,
Megalodontidae.
by
family Cephoidea because of
with the
are
(d—e):
Praesiricidae which
the
marked
extinct
The
living
without SC.
family
to
1990
Rasnitsyn,
of the
Representatives
strong,
by
needle-like
represent
to
their characteristic
the
Anaxyelidae (with
ovipositor).
predominantly
a
single living
A.P.
16
Rasnitsyn
Fig.
8.
The
1969
primitive
7.
lula
hybrida Rasnitsyn,
syn,
of the
Representatives
1993;
sawfly family Sepulcidae:
aculeatus
Rasnitsyn,
minutus
South Kazakhstan
(a, c)
and Transbaikalia
(d) (all
1993;
Rasnitsyn,
from
1993;
the
Upper
after
Jurassic
Rasnitsyn, 1993).
9.
the
by
synapomorphic
re-
media
c,
micrura
holotype).
that these two
important
slightly
Xyelidae,
seems
modified
segments
anaxyelid
normal
in
genera
or
with
large
in
but
S.
display
different
seemingly
see
that
it
related
either
only slightly
Rasnitsyn,
we
eration for
can
en-
1969).
take all the above data into consid-
assessing
antenna with
regard
the
to
meaning
the
early
of the
after
all from the
Upper
Jurassic
Rasnitsyn, 1969; b, original,
menopterous insects. In fact,
conflicting
xyeloid
evolution of hy-
putative
mentioned genera
with the
Xyelidae
we
of South
based
on
phies shared with
tails.
These
above
are
supposedly synapomorphic
some
next
as
to
while
be apomor-
other sawflies
cannot
putative
falsified (reassessed
seem
and horn-
cannot all be correct,
hypotheses
similarities
conflicting
The above-
in their antennal structure,
phies. Therefore our
the
encounter here the
synapomorphies.
their various other characters
the
Now
d,
a,
Rasnitsyn,
the
ways:
are
c,
inversa
seg-
entire in S.
Closely
antennae
for details
to
It is
subdivided into
clearly
media.
the 3rd segment
larged (Fig. 9c-d;
3rd
comparison
two
diminished but
inversa,, while it is
primary
species have the
and modified in
simply
wing.
(a,
S.
1925; d, Brachysyntexis
Anaxyela gracilis Martynov,
1969;
family Anaxyelidae:
1963); b,
(Rasnitsyn,
Rasnitsyn,
in the fore
of lr-m in the hind
of the horntail
Representatives
Kazakhstan
ment
South
of
duction of the SC stock and of the 2r-m cross-vein
wing and
of
Mongolia (b)
1968;
confirmed
Jurassic
1969).
Xye-
Syntexyela
as
(after Rasnitsyn,
xyelopterus Rasnitsyn,
Upper
Rasnit-
Fig.
species),
the
Micram-
e,
and Lower Cretaceous of
a,
from
1968; d,
Parapamphilius confusus Rasnitsyn,
c,
Pamparaphilius mongolensis
philius
Onokhoius
1969; b,
horntail Protosirex
(Gigasiricidae)
Kazakhastan
Fig.
taxonomy, and nomenclature
Phylogeny,
-
all
task is
to
be
i.e.,
synapomor-
decide which of
synapomorphies
symplesiomorphies
can
or
be
ho-
Contributions
moplasies)
to
arrive
considered here:
is
tenna
of
being
of
groups
spective
the
a
an-
while the alternative
are
genera
sets
in
genera
above
synapomorphies (the
are
with
monophyletic
and
superfamilies
has been
antenna
and
the
indebut
above,
as
antenna
as
hypotheses (1,2)
similarity
belong
some remote
similarities,
implies
that
the
thus
For
ovipositor
in
in
it is
most
but
difficult
and
hardly
to
and
ones
supposedly
Rasnitsyn,
duced,
Other
is
a
the
to
flat leaf-
or
dle-like
ovipositor,
hemipteran
crickets,
are
involves
clearly
both
the
e.g.,
one
of
an
all
serrate
anaxyelid
apomorphy
of the
structure
some
(Fig. 4).
of the
members of the
apomorphic
true
which
nee-
nature.
in
This
presumptions,
of
analogy,
nature
The
for
is
or
same
the
of the
judge-
reduced
hardly accept-
be
to
homoplasy,
(Fig.
6
erations of taxonomists
families and
acters
for
acters
were
observed
homoplastic
and
as
This
diagnostic
char-
development
therefore agree with the
These
especially
in
un-
gen-
sound
not
seems
accepted by
superfamilies.
as
mem-
families
having originated
as
italics ).
10,
imply
with other
superfamilies
considered
groups,
it does
because the above-
since these traits have been
likely
seen
the homo-
non-xyelid
synapomorphies
independently
related
charto
prone
groups
of
presumption
a
and
weighted
similarity.
One
to
more reason
hypotheses
consider either of the above
is the incidence of in-
unacceptable
as
termediate antennal types described above for two
species
of
of S.
ture
mediate
tenna
xyeloid
several
segment,
both
relatives,
under the
nothing
cases,
in
the
all
large
struc-
inter-
an
normal
in S.
an-
inversa
basiflagellar
seemingly
en-
xyeloid segment (which is
above
common
hypothesis),
or
other-
with that of S. media.
species
two
their
but
as
a
the
congeners)
rather
reduced
a
and
the antennal
In contrast,
one.
other
which is
is either
tire,
In
a
Indeed,
easily interprétable
of transformation of
stage
into
in
(and
Syntexyela.
media is
wise has
unless
respective
respective
would have
re-
ovipositor,
plesiomorphic
wing venation,
bers
of
putative
impossible
Archescytinidae,
holds
listed
in-
ovipositor of the sawflies.
of
among the
the
amount
hymenopterous
paleontological
support the
sev-
vast
be
can
hypothesis (2), suggesting
and
or-
the
presumption
outgroup one, and
which
are
ovipositor
direction of
so
examples
in
family
discussed
as
of the similarities between the genera
nature
an
saw-like
(non-hymenopterous)
oldest
ment
This
incom-
hypothesis infers
presents fewer difficulties. However,
in other
so
(including Palaeomanteida,
1980), the
and
non-hymenopter-
particularly
ancestral
re-
pamphilioid
reverse
the
it
believe that the
modified
majority of
all
gen-
common
respect of the saw-like
winged insects,
sects;
the
to
presumptions).
presumptions suggest the
Paleozoic
der
applied
likely because
Xyelidae and Tenthredinoidea. Indeed,
evolution. In the
ous
some cases
(a)
of inde-
(b)
the strong needle-like siricoid
plesiomorphic
eral
family),
that which is
to
(i.e.,
relevant
reduced
or
when
1,
opposite
normal
instance,
specifically
of the
is
the
possible
seems
ones.
the
the
direction of evolution of the
as
satisfying
to
ancestor or
the least
seems
spective characters
erally accepted
with
other groups is either
to
pendent origin. Hypothesis
those
that all the
monophyletic
are
The second
plastic
illus-
cannot
ovipositor (see above).
a
mean
I
presumptions,
of the
more
relevant clado-
a
from the
case
pre-
a
patible polarization
families)
acquired
(4)
re-
many
plesiomorphic.
discussed with
just
one
again
Hymenoptera
hypotheses,
gram. This is because this
question
inherited from
Unlike the next three
alternative
Xyelidae (i.e., cladistically
while their
venation is
complete wing
complete xyelid and
more
because
confirm that in the
sumptions
sym-
hymenopterous symplesiomorphy.
two
the
to
venation,
the
similarity lies in having the xyeloid
The first
ancestral
as
pamphiliid
homoplasies, (3)
above,
homoplasy),
a
xyeloid
able
trate the
are
xyeloid
the
be
can
alternative
genera
pendently (as
in
listed
as
non-xyelid
while the
and consis-
hypotheses
the
of similarities
sets
17
but
(2)
plesiomorphies,
similarities
the
1996
likely
most
(1) similarity
synapomorphy
a
similarities
of
the
at
-
(1)
Four different
cladogram.
tent
66
Zoology,
to
similarity
cannot
be
should be
close
homo-
plastic.
So
we
now
move
hypotheses (3,4).
larity
the
on
to
the
last
of the above-listed genera
xyeloid antennae)
groups
is
two
to
synapomorphic.
the
(those possessing
respective
These
non-xyelid
similarity
the xyeloid antennal structure is
homoplastic
or
differ
hypotheses
from each other in proposing that the
possessing
alternative
Both of these claim that the simi-
symplesiomorphic
in
of
either
nature.
The
18
A.P.
Fig.
that
10.
Cladogram
in
similarity
descent
refer to
of the lower
the
the
Achterberg
of
1 — Synapomorphies
flight
specialized,
for
nitsyn,
1988a,
2 — 3rd
antennal
the
& Van
order
much
short
apex
abscissa of RS short,
Vespida
and with
with
male
1988a:
with
(up
to
and RS
5
lost
distant,
with
molar
lobes
8 — propleurae
anterior
arms
Ras-
for
rotated
wing
with
2r-rs
basicon-
with
1st
large;
one;
except
pronotum
crossvein-like
fore
are marked
or almost
with
the
(based
on
in
hypothesis [(2)
homoplastic. Figures
under
Rasnitsyn,
the
text]
the lines
1988a, and,
of
for
by italics):
all
length of mesothoracic
with
11 — fore wing
submarginal cells;
structures
180°;
larva
modified;
larva
1-segmented,
and with
items
with
with
suprapedal
further
SC
venter;
short, appressed
R,
to
lacking fore
with
eye
and antenna
mandible
and
costal
full
space
narrow;
ovipositor
needle-like
certainty);
13 — propleurae contiguous ventromesally; fore wing
male
lost;
described
mandible
in
tentorial
item
bridge intervening
with M+Cu
between
15 — tibial
from
spurs
as
hypostomae;
twisted into
moving
angulate; ovipositor small,
exophytic, silk-protected,
RS2
etc.
13);
of cutting type (with cuttingedge
larva
with
gonomacula subapically;
Rasnitsyn (1988a: 121,
plane); fore wing
like;
with
gonostylus
with
claw-
appendages setiform,
eye;
with
soft
tips;
inner
tooth
of
tarsal
claws
reduced; hypopygial depression large;
lacking
subspiracular
synapomorphies,
see
16 — membrane of fore wing
on
17
tentorium
bridge; fore wing
issuing
with
RS2
small, claw-like modified;
wings lacking SC, pseudosternum extending
all
less
folded apically;
larva
with SC
— fore wing
lacking fore branch;
femora
widened;
hypopygial depression medium-sized;
18 — vertex
with
with
19 — fore wing
very
short
claws
over
more or
Angiospermae;
tarsal claws
5, 6, 8, 9);
below tentorial
with
— fore wing
and
20 — malar
9 — ovipositor
are
follows
as
antenna distant
23, 3);
between
lost;
10 — both
taxa),
14 — lower
short
possible synapomorphies,
contiguous ventromesally;
from
terminal
12
far beyond 2r-m,
further
numerous
120,
(or
similarities
(not known
beyond composite segment; fore wing lacking
flange,
1988a:
lost
items
lost
with caudal
antenna
merged;
see
segments beyond
1mcu cell
SC
(for
120,
RS
genitalia
incisive
Rasnitsyn,
RS2 lost
stem,
7 — flagellum
2r-rs;
etc.:
enlarged (antenna xyeloid);
thick; fore
and with
medially; fore wing
fern
venation
(wing
constructed consistent
alternative
Aartsen, 1986; homoplasies
5 — single flagellomere beyond composite
boring
clades
respective
taxonomy, and nomenclature
Phylogeny,
-
branch;
rather
necting pterostigmal
Rasnitsyn,
and the
(basiflagellar) segment compound;
4 — antennal flagellum
see
(suborder Siricina)
details);
flagellar segment),
6 — fore wing
of
dipterous, haplo-diploidy,
3 — basiflagellar segment
branch,
insects
synapomorphic,
of synapomorphies
sets
on Van
Pamphiliidae,
hymenopterous
antenna is
xyeloid
Rasnitsyn
with
grooves
two
with
thin,
space
1st abscissa
and with
with
acute basal
21 — ovipositor
deep
and
subparallel anteriorly;
long bristles medially;
sheath
of RS
differentiated
short,
very
conspicuous
dark
setose
with
1r-rs
patches;
depression;
tarsal
lobe;
with large stylus
[possibly secondarily
Contributions
first
(3, Fig.
have
11)
this
supposes that
quired by
lost in
a
&
question
hypothesis (4, Fig.
(Linnaeus,
12)
uniquely
and later has
ancestor,
betulae
19
in
genera
character has been
common
Pamphilius
Pesarini
1996
"third" segment inde-
compound
while the second
pendently,
-
(J)
that the
means
the
acquired
66
Zoology,
to
and
1758)
ac-
been
festivus
P.
Pesarini, 1984];
22 — female
fore
with inner tooth
with distal third of costal
of hind
claw
cell
shorter
glabrous;
than
apical
24 — fore wing
with
costal
narrow;
space
needle-
ovipositor
like;
26 — head
and hind
with
capsule
elongate;
lateral
lines;
bearing
feeding
fungi,
and oblique sulci;
with
envelope,
tails,
see
sterna
larval
ductus
with
synapomorphies listed
short
28 — pronotum
pterostigmal apex
larva with
180°;
incisive
molar
scribed in
wood
at item
lobes
30 — pseudosternal
8
postero-
also
with
all
length of mesothoracic
with basal
on
herbaceous
by symbiotic
and
covered
other
22,
with
possible
with
connecting
1st abscissa
genitalia
rotated
lacking
synapomorphies
items
hind
2r-rs
with mandible
distant,
numerous
as
de-
2—3, 3, 5);
wing lacking
and with
sternum
prolegs 2-segmented,
with
salivary gland
with
subanal
margined
cross-
spurs
lost;
2
apically;
of
rows
cavity
and
occipital foramen;
below
mesally;
tentorial
male
abdominal
oblique sulci,
basally
toward
common
hypostomae contiguous
35 — ovipositor
lost in P.
—
sterna
base
with
subanal
32 — ovipositor small,
with
and
anterior
larval
ductus
claw-like
oral
arms
ventro-
larva
and
longitudinal
appendages segmented,
slit;
shifted
salivary gland covered
quadrangularin
modified;
larva
from
into
wing
and
feeding
lower
sheath
with
fore
mesofurca
with
tibia
with
of swellings;
specialized
sheath
with
hind
fore
costal
modified
larva
inner
with
exophytic
appendages
large stylus
makes
with
with
with
with
apical tooth; fore wing
on
setiform,
more
or
less
[possibly secondarily
(Gmelin, 1790)];
pair
larva
moving plane);
membrane of fore wing
eye;
arms
with
stylus
rudimentary (or lost);
spur
long, fused for
described
as
sheath
leafroll;
stylus large, glabrous;
narrow;
space
ovipositor
(outer)
larva
some
distance; fore
xylophagous
living
on
(1988a: 122,
by Rasnitsyn
item
19);
39 — fore wing
with
SC lost
and with
Cu
straight
within
1mcu
cell;
40 — mesonotum
costal
membranous
space very
in
narrow
41 — metanotum with
RS
very
fore
except laterally;
basal
section;
exophytic,
cenchri
short and with
third
wing
with
half; ovipositor long
or
with
A2
ditioned
silk-
boring
in
item
fore wing
hearing transscutal
suture,
abdomen
lost;
aspera
of
1r-rs
larva
feeding
on
dead
wood
in
con-
Rasnitsyn
with
single
postgenae
prepectus
cross-vein
contiguous;
r-m
each;
mesoscutum
concealed under
postero-
edge ofpronotum;
with
45 — mesoscutum
laterally;
fore
adlateral
modified
listed in
lines;
into
transscutal
with basal
wing
46 — tentorial bridge
as
area
1st abscissa
cross-vein
angiosperm plants;
wing
with
gum
with
with
22);
and hind
44 — head capsule
with
large,
by symbiotic fungi, modified as explained
(1988a; 122,
lateral
1mcu
and
straight,
larva
lost; fore wing with
cell
42 — ovipositor needle-like;
and
between
gonomacula subapical;
lacking prolegs
of anal
envelope,
issuing
than
silk-protected,
nemorum
36 — frons
38
shorter
larva
glandular
bridge; propleurae contiguous
gonostylus
with
tentorium
larva
medially; female
angulate; ovipositor small, claw-like;
Angiospermae,
43 —
with
claw
bristles
long
compressed;
cells;
31 — head capsule
fore
straight line,
aspera;
twisted
cutting edge
two
male
appendages lost,
with
eye;
extending;
m-cu
suprapedal and
excised
hardly visible, strongly
ductus
M+Cu
long,
tibial
hind
of
wing
male
large);
lost; preapical
with
tooth
(with
37 — ovipositor
de-
from
deep and subparallel anteriorly;
grooves
with
by
merged;
sulci
cutting type
tarsal claws
glabrous;
common
area
a
tentorial bridge intervening between hypostomae; mandible of
longitudinal
(for further
section
19 &
fore
in
loss of
angiosperm plants;
with
antenna distant
larva
pro-
extending over
pectinate;
entire
forming
behind
straight, running
A2+3
distant
antenna
venter; antenna
in
or
and pseudosternum
sections of RS and M
they
homoplasies
acquisition
appendages setiform,
folded apically;
21—22);
1-segmented,
with
by
protected,
spur
(outer)
crossvein-like
beyond 2r-m,
and antenna
with antenna
subspiracular
with
single
in
antennae.
comparison
because
numerous
(either
in
likely
more
hypotheses,
than
character
plants,
Rasnitsyn (1988a: 120,
larva
and
lost;
29 — fore wing lacking 2r-rs,
from
pose
of
pair
with normal
groups
seem
more
long, fusedfor
conditioned
items
122,
1mcu cell
flange;
hind
lacking prolegs
and R-RSfar
eye
except
medially; fore wing
and with
neck-
suture
under
arms
with
occipital
gonomacula subapical;
salivary gland
there
with RS 2
— fore wing
with
fore
quadrangular in
1988a;
Rasnitsyn,
of RS short,
with
angiosperm
abdominal
concealed
with SC lost
gonostylus
dead
on
with
transscutal
tibia
edge of pronotum; fore
male
and
and
cavity
each;
r-m
contiguous
oral
prepectus
distance; fore wing
some
cross-vein
medially, dorsally; propleurae
rudimentary (or lost); mesofurca
branch;
single
hypostomae
mesoscutum
adlateral
with
between
short
foramen; pronotum
incipient
wing
subcontiguous
postgenae
vein;
little
34 — vertex
25 — fore wing
27
hypotheses
with the first
with
tooth;
like
independently in
Both
33 — both wings lacking SC,
23 — female
wing
lost
narrow
fore
suture
abscissa
of RS
stripe-like,
n-like
wing
mesosomal
with
SC
lost;
propodeum;
bent
bent;
mesonotum
1st abdominal
larva
Rasnitsyn (1988a: 123, item 28).
cephalad
subvertical;
parasitic;
ter-
etc.,
A. P.
20
11.
Fig.
of the lower
Ciadogram
that
similarity
the
following synapomorphies
1 — Same
as
in
in
the
xyeloid
hymenopterous
antenna is
(suborder Siricina)
insects
and the
homoplastic,
alternative
Fig. 10;
and
very
with
further
large (for
120,
2r-rs
with
beyond 2r-m,
1988a:
item
connecting pterostigmal
1st abscissa
of RS short,
apex
possible synapomorphies
see
crossvein-like
Rasnitsyn,
short
to
5
segments),
medially; fore wing
structures
prolegs 2-segmented,
with
salivary
with
ductus
gland
and
flange,
for
120,
larva
one; fore
boring
larva
in the
text]
Fig. 10,
with
hypothesis [(3)
Otherwise
as
in
subanal
margined
appendages lost,
with
2
rows
of
and
glandular
with
with
and antenna
mandible
suprapedal
further
eye
and
lacking
incisive
subspiracular
synapomorphies
see
lobes
Rasnitsyn,
beyond composite
with
10 — hind wing lacking
11 — pseudosternal
from
2nd claw
m-cu
sulci
with
below
tentorial
male
toward
with
with
anterior
and
larval
slit;
with
and
shifted
salivary gland covered by
quadrangular in section;
13 — ovipositor small, claw-like modified; larva exophytic,
protected,
larva
longitudinal
appendages segmented,
ductus
oral
arms
ventro-
gonomacula subapical;
lacking prolegs
subanal
issuing
propleurae contiguous
with
base of anal
envelope,
tentorium
bridge;
sterna
between
hypostomae contiguous
gonostylus
abdominal
basally
with
occipital foramen;
appendages setiform,
antenna distant from
15 — both wings lacking SC; pseudosternum extending
or
almost all
of mesothoracic
length
silk-
eye;
very
segment;
and
very
larval
leg
modified;
16 — basiflagellar (3rd antennal) segment compound and
hardly visible, strongly
M
forming
behind
entire
area
18 — lower
tibial
all
very
fore
wing
straight line,
larva
aspera;
with basal
sections
and with A2+3
feeding
on
of RS and
straight, running
herbaceous
angiosperm
plants;
cross-vein;
lost; preapical
over
venter;
large;
17 — antenna pectinate;
lost
8
distant,
5, 6, 8);
secondarily 2-clawed
and
large;
with
large;
flagellum
with
wing
fern stem, with
(3rd antennal) segment compound
sternum
except
180°;
2r-rs;
numerous
items
12 — head capsule
cavity
oblique sulci,
SC lost
9 — basiflagellar
with
the
synapomorphic.
14 — basiflagellar (3rd antennal) segment compound and
1-segmented,
antenna
merged;
with
very
thick;
modified;
8 — fore wing lacking
1988a:
are
with
common
submarginal cells;
7 — male genitalia rotated
molar
rather
and
fore branch;
lost between
with
constructed consistent with
similarities
mesally;
6 — single flagellomere beyond composite
caudal
1mcu
23);
large; flagellum short (up
5 — pronotum
and RS
and with
4 — basiflagellar (3rd antennal) segment compound
RS
taxonomy, and nomenclature
cells;
large;
3 — fore wing
cell
Phylogeny,
-
inferred:
2 — basiflagellar (3rd antennal) segment compound
far
Rasnitsyn
spurs
excised
lost;
male
apically; larva
mandible
plane);
tentorial
of cutting
fore
wing
bridge intervening
between
type (with cutting edge
with
M+Cu
angulate;
hypostomae;
twisted into
moving
Contributions
66
Zoology,
to
the
hypertrophied
ply
any
"third"
1996
-
21
and do
segment)
falsification of the
well-based ideas
(1)
im-
not
existing and apparently
the
regarding
evolutionary
path-
The
of
terms
last
two
homoplastic
alternative
Figs.
11
and
of
loss
multiple
the
results
in
makes
it
the
possible
soft
more
loss of
to
apply
to
which will confirm
hypothesis.
applying
However,
weighted similarity
structures
inner
tips;
a
with the
captions
to
seems
generally considered
are
tooth
of tarsal claws
tibial
to
spurs
reduced; hypopygial
depression large;
20 — fore wing
less folded
more or
larva
apically;
on
SC
femora
lacking fore branch;
widened;
with
hypopygial
de-
claws
with
with
grooves
and
deep
bristles
long
two
1st abscissa of RS
medially;
short,
very
dark
conspicuous
with
1r-rs
short
wing
and
with
thin,
and
patches;
with
malar
claws
shorter
of swellings;
pair
than
larva
female with
apical tooth; ovipositor
makes
space
with
with
dif-
basal
acute
distal third
of costal
27 — female with
inner tooth
sheath
of hind
with
stylus
specialized leafroll;
26 — basiflagellar segment enlarged;
a
complex
asynchronous
The
plain.
of
hypothesis
antennal
pound
quite
homoplastic
multiple
and
remote
of
structure, and thus it
of
hypothesis
a
This is
consideration
needs
segment
event
the
cell
inner
fore
wing
of female
with
glabrous;
tooth
P.
sheath
nemorum
only
acquisition of
with
claw
shorter than
apical
large stylus (possibly secondarily
larva
makes
specialized leafroll;
31 — fore
mesofurca
tibia
with
xylophagous
segment
with
fore
on
hind
arms
living
Rasnitsyn (1988a: 122,
32 — fore wing
with
enlarged;
(outer)
long,
spur
with
large,
1m-cu,
with
repeated
gain
especially
some
modified
as
distance;
larva
described
by
if
so
larva
space very
extending;
narrow
in
takes
case
larva
boring
with
feeding
lost; fore wing
with 1st abscissa
1r-rs
in
costal
into
of
Syn-
and
very
with
costal
short
very
with Cu
area
space
and cell
straight
before
abdomen
lost;
aspera
in
conditioned
Rasnitsyn
needle-
ovipositor
narrow;
wood
explained
as
of RS
angiosperm plants;
space
dead
on
long,
and with
straight,
modified
by
symbiotic
(1988a:
122,
37 — basiflagellar (3rd antennal) segment compound
38 — fore wing
and hind
39 — scutellum
rounded
item
and
very
wing
with
cross-vein
single
to
except laterally;
basal
third
or
each;
r-m
basally;
entire
externally;
41 — basiflagellar segment compound, enlarged;
basal
42 — several
flagellar segments enlarged
about
to
form
extending;
bearing
with
transscutal
edge
of
with
46
tum
R, lacking
fore
fore
wing
half; ovipositor
with
postgenae
bent
suture
abscissa of RS
except
mesoscutum
under
concealed
postero-
pronotum;
transscutal
SC lost
postgenae subcontiguous;
suture; prepectus
44 — head capsule
contiguous;
mesoscutum
fore
wing
with
lines; fore wing
with
cephalad laterally;
subvertical;
with
incipient
crossvein-like
adlateral
fore branch;
basiflagellar segment compound, slightly enlarged;
short
medially,
feeding
on
stripe-like,
with
well
long
modified
developed;
listed in
into
dorsally;
prono-
propleurae neck-like elongate;
angiosperm wood;
47 — head capsule
narrow
membranous
one
to
obscure
22);
larva
19);
short, appressed
branch;
33 — mesonotum
enigmatic
of the
above-mentioned
cross-vein
A2
compressed;
like;
lost,
45 — mesoscutum
rudimentary (or lost);
fused for
plants,
item
SC
single and
a
the
superior compared
seems
with cenchri
with SC
narrow,
basal
29 — basiflagellar segment enlarged;
30 — basiflagellar
ex-
com-
large;
43 — head capsule
hind
of
(Gmelin);
to
34 — basiflagellar (3rd antennal) segment compound
lateral
28 — ovipositor
widely
difficult
origin
repeated loss of the
a
an
structure
function. This makes its
scure
is
and of ob-
compound, enlarged basiflagellar segment; ovipositor scarcely
tooth;
costal
compound
rather
mul-
a
antennal segment
40 — basiflagellar segment enlarged though
glabrous;
lost in
with
organ
a
reduction is
have occurred than
to
to
function,
large;
tarsal
depression;
setose
very
fore
lobe;
25 — frons
claw
tiple origin.
fungi,
24 — basiflagellar segment enlarged;
ferentiated
tarsal
subparallel anteriorly;
23 — basiflagellar segment compound, enlarged;
with
The
logical
seems
subsequent multiple
likely
more
36 — fore wing
pression medium-sized;
22 — vertex
generally
the
it
structures
compared
as
structure of enigmatic
a
and
unique origin
1mcu
Angiospermae;
21 — basiflagellar segment compound, enlarged; fore wing
multiple origins
multiple loss. Equally
35 — metanotum
membrane
have
to
that for
adaptation.
19 — basiflagellar segment compound, enlarged;
with
in
compared
a
clear results.
more
Complex
from
homoplastic
a
parsimony,
of
presumption
give still
This
12).
presumption
the
of
as
italics
(cf.
state
equivalent
antenna
one,
homoplasy
hypothesis (4)
character
the
xyeloid
segmented
of
cases
in
founded. Hypothesis 3, of
of the
origin
homonomously
abundant
the
best
not
are
likely
their
assume
hypotheses
being the
less
are
to
ways of other characters.
thus such
originate with great difficulty;
with
postgenae contiguous;
n-like
fore
bent;
wing
mesosomal
tentorial
mesonotum with
with
SC
lost;
propodeum;
Rasnitsyn (1988a: 123,
item
1st
larva
28).
bridge
adlateral
lines
abdominal tergum
parasitic;
etc.,
as
22
A.P.
Fig.
that
12.
of the lower
Cladogram
in the
similarity
11, with
the
1 — Same
xyeloid
in
as
apex
fore
see
and RS far
1mcu cell
cross-vein-like
item
120,
1st abscissa
with
6 — fore wing lacking
—
items
flagellum
secondarily
m-cu
boring
wing
fern stem,
with
with
larva
with
with
eye
and antenna
mandible
suprapedal
further
with
and
lacking
distant,
incisive
lobes
subspiracular
synapomorphies
composite
2nd claw
see
Rasnitsyn,
segment;
larval
leg
ordinary;
hind
wing
cross-vein;
strongly
tibial
spurs
in
hypothesis [(4)
Otherwise
as
in
the
lost;
male
apically;
text]
10—
Figs.
rows
with
salivary
glandular cells;
tentorium
with
issuing
between
anterior
subanal
toward base of anal
with
slit;
ductus
and
ventro-
pseudosternal
with sternum 8
larva
with
hardly
prolegs
2-
larva
longitudinal
salivary gland
covered
or
almost all
appendages setiform,
length
of mesothoracic
13 — antenna pectinate,
ordinary;
entire
fore
wing
straight line,
larva
with
with
distant from
antenna
feeding
on
diminished
in
size
eye;
over
all
venter;
sections
A2+3
of RS
and M
straight, running
herbaceous
14 — basiflagellar segment
silk-
basiflagellar segment seemingly
basal
and with
by
quadrangular in section;
11 — ovipositor small, claw-like modified; larva exophytic,
with
and
shifted
appendages segmented,
larval
oral
arms
gonomacula subapical;
lacking prolegs
sterna
envelope,
of
bridge; propleurae contiguous
and with
oblique sulci,
common
2
hypostomae contiguous
gonostylus
abdominal
and
appendages lost,
with
with
tentorial
male
angiosperm plants;
subdivided
(subequal
forming
behind area
in
into
width
primary segments
though
to,
much
longer than, following segments);
15 — tibial
segment seemingly ordinary;
excised
subanal
margined
occipital foramen;
below
mesally;
and
modified;
segment seemingly
sulci lost; preapical
from
aspera;
beyond
the
12 — both wings lacking SC; pseudosternum extending
5, 6, 8);
2-clawed
9 — basiflagellar
visible,
with
lost
8 — basiflagellar
lacking
larva
one; fore
180°;
2r-rs;
for numerous
1988a: 120,
7
and
flange,
merged;
except
and
protected,
1-segmented,
antenna
molar
lost
modified;
rotated
with
synapomorphic.
are
10 — head capsule
basally
submarginal cells;
structures
5 — male genitalia
with
SC
fore branch;
RS lost between
ductus
cavity
with
2—3);
4 — single flagellomere beyond composite
caudal
of RS short,
possible synapomorphies,
fore wing
medially;
basi-
connecting pterostig-
with
segmented,
gland
segments beyond
with
similarities
inferred:
2r-rs
further
taxonomy, and nomenclature
Phytogeny,
-
constructed consistent
and the alternative
large;
6
to
with
beyond 2r-m,
short
3 — pronotum
(up
wing
large (for
1988a:
Rasnitsyn,
and very
short
2 — antennal flagellum
and with
(suborder Siricina)
10—11, and additionally, basiflagellar (3rd
Figs.
flagellar segment);
insects
symplesiomorphic,
following synapomorphies
antennal) segment compound
mal
hymenopterous
antenna is
Rasnitsyn
spurs
with
soft
tips;
inner
tooth
of
tarsal
claws
reduced; hypopygial depression large;
16 — membrane
on
of fore
Angiospermae;
wing
more
or
less
folded
apically;
larva
Contributions
texyela.
It forces
different
66
Zoology,
to
make
to
us
-
(1)
choice between the
a
that claim that the
hypotheses
structure evolved either by reduction
The
ple origin.
favour of the
The
plete,
rather
the
xyeloid
antennae.
need
them,
two
those
far
and
explicit
the
in
form,
with
SC
hy-
The
picture.
de-
hotly
antenna
a
The
(1976).
femora
lacking fore branch;
as
in
(1937) and,
Königsmann
I
man.
widened;
hypopygial depression medium-sized;
18 — vertex
claws
with
with
short
and
deep
bristles
long
two
19 — fore wing
very
grooves
with
tarsal
subparallel anteriorly;
short,
very
dark
conspicuous
with
1r-rs
with
space
still
(though
sheath
differentiated
setose
depression;
tarsal
lobe;
with
tibia
with
mesofurca
with
fore
xylophagous
hind
living
on
Rasnitsyn (1988a: 122,
26 — fore wing
with
(outer)
SC
rudimentary (or lost);
spur
fused for
distance;
some
modified
plants,
item
larva
described
as
by
in basal
third
with
cenchri
abscissa
1mcu
short, appressed
to
R, lacking
fore
area
short,
Cu
aspera
with
with
straight
lost;
appearing
and
later
wing
costal
with
wing
with
abdomen
costal
space
synapomorphy
1r-rs
narrow;
with
A2
larva
in
base
by symbiotic fungi,
1st
width
though
to,
capsule
with
is
of
general-
transformation
changes
are
22);
somewhat
diminished
cross-vein
dissociated
r-m
into
in
fore
size;
each;
primary
flagellar
seemingly ordinary (unknown
to be
dissociate
into
size
of
as
a
whole,
or
to
ordinary size);
segment
subdivided
into
reduced
size
as
in
size
in
as
diminished
segment
much
primary flagellar
whole
a
(still
subcontiguous;
mesoscutum
concealed
prepectus
under
in
head
bearing
posterolateral
edge
of
37
basiflagellar segment seemingly ordinary;
—
(subequal
longer than, following segments);
postgenae
suture;
pronotum;
postgenae contiguous;
head
capsule
mesoscutum with transscutal
fore
cephalad laterally;
38 — mesoscutum
SC lost
39 — pronotum
elongate;
in
(unless
on
dead
explained
in
with
wing
larva
with
basal
abscissa
suture
of RS
short
bent;
wing
segment
mesosomal
mesonotum
with
dorsally; propleurae
medially,
postgenae contiguous;
n-like
lines; fore wing
adlateral
with
fore branch;
neck-like
feeding on angiosperm wood;
40 — basiflagellar
fore
incipient
except crossvein-like
straight,
Siricoidea
feeding
modified
our
case
rounded;
36 — basiflagellar
and cell
boring
probably
remaining
larva
are
than
more
subvertical;
metanotum
long
with
compressed;
of all
it
ir-
irrevers-
framework of
particular
single
segment
diminished
33 — scutellar
bent
space narrow,
1m-cu,
hardly
evolutionary
item
of the
treat
Indeed,
analogy.
of
Königsmann
particular
more
a
in-
law, but exceptions
In the
one
within
basiflagellar
either
space very
costal
ordinary;
cross-vein
before
Vespina) ovipositor needle-like;
wood conditioned
and hind
a
(Königs-
partial
a
presumption
that it is
of
segments (still widened); ovipositor scarcely extending;
with
with
as a
that
Rasnitsyn (1988a: 122,
with
angiosperm plants;
29 — fore wing
of
directions
series,
mesonotum
half; ovipositor long extending;
lost; fore wing
with
fore
segment seemingly
of RS
very
large,
and with
or
just
presumption
transscutal
19);
except laterally;
28 — basiflagellar
is
for
respect
and
disproportionally large);
27 — basiflagellar segment seemingly ordinary;
narrow
presumption.
it
35 — basiflagellar
branch;
membranous
another
agreed
wing
as
Ross
shortage
segments;
large stylus;
long,
arms
even
34 — basiflagellar segment
24 — basiflagellar segment seemingly ordinary;
25 — fore
ly
or
so numerous
primarily segments
22 — basiflagellar segment seemingly ordinary;
23 — ovipositor
the
the
be
approach
32
patches;
thin
disproportionally long);
claws with acute basal
just
rule
a
to
of
The
provides only
on
31 — basiflagellar segment
of RS
20 — basiflagellar segment seemingly entire,
21 — malar
based
30 — basiflagellar segment
medially;
1st abscissa
and with
thin,
and
as
or
weak.
of evolution. We often
reversibility
known
of
opinions
of view of Ross and
point
additionally
ibility
to ana-
(Ross)
(Ras-
hypothesis evidently
last
all
at
xyeloid
mine
1988a).
the
fossil record
The
sight.
not
are
about
the
the
but
is
Vespida
1980,
here,
loss of the
multiple
a
order
1969,
preferable,
mann)
is
the
portrayed
knowledge
they have
xyeloid
be found in Ross
can
the
straw
a
As
1968,
Königsmann
com-
possessing
have been
the
considering
one
nitsyn,
mostly
attempt
an
complete
to
not
are
know,
creating
hypotheses actually
17 — fore wing
still
within
antenna
looks
groups
I
as
hypothesis proposing
in
clearly
concerning
of all
bit like
a
however,
homoplastic
more
As
multi-
relevant presump-
one
proposed before,
bated. The
is
hypotheses
monophyly
them looks
other
be
a
hypothesis.
omitted
obvious, especially
been
by
or
to
seems
enigmatic
the above considerations
on
lyse
of
have
we
pothesis
never
loss
repeated
Indeed,
tion.
option again
comparison
for
23
1996
SC
lost;
propodeum;
Rasnitsyn (1988a: 123,
seemingly ordinary;
tentorial
with
bridge
adlateral
1st abdominal
larva
item
28).
lines
well
tergum
parasitic;
head
etc.,
capsule
stripe-like,
narrow
developed;
modified
as
listed
into
in
24
A. P.
with
commonly asymmetrical,
pearing
realized
changes
cladogram supposing
site direction,
one
ought
changes
no
the least
or
unidirectional
note
one can
definitely realized,
as
their
of
prefer
to
in the oppo-
unless
number,
and until there is strong contrary evidence.
Indeed,
tried
to
pends
show
on
evidence
contrary
above,
what
of the
complex
against
the
the
permit
selected,
to
us
the
vs.
of
change
of the
when
occurred,
hope
convincing
se
the
higher
gain
of evolution. I
per
assess
a
system
of
bionomically,
we
with
normal
a
with
predict
a
it
rather
to
a
insect
The
on.
the
taxonomie classification
diversity.
created
There
cause).
are
fulfil
to
of pests
universally
very
accepted
To
meet
as
possible.
for
an
a
the
taxa
For
are
purpose
a
as
tween-group
words,
it
characters
as
must
two
once
One of them is
within
they
bio-
to
on
to
the system
for
a
a
as
considerations
mention them
are
only
as
taxa
the
be
many
must
users
a
for
the
and
third
original
518).
misms
are,
are
however,
"bad
more
"eclectic",
etc.
not
1979:
just
terms
taxonomy")
but
dently
euphe-
a
(Rasnitsyn,
name.
of the
distinct
latter,
tax-
how-
Once
name.
seems
I
is evi-
1992b), which
Phylistics
the
reveal
to
principles of traditional
equivalent
deserves
awkward
an
adequate
the
the
497,
qualify
to
explicit
full
to
proposed
"synthetic",
(Farris,
approach.
ever, and thus it
cladistics
names
and make
not a
mis-
equally
call
This version of taxonomy tries
onomy. It is
"phylo-
"phylogenetic" relevant
alternative:
(replacing
people
most
"syncretistic" systematics,
These
it from
and
the
"evolu-
name
difference between the terms
There
context?)
the
an
for what
what is
"evolutionary"
present
distinguish
to
and
cladism,
misleading
a
a
more
qualification.
must
cook, and
present its
and
possible.
of all
and
as
In other
possible
different
lesser
be-
importance.
store the information in the
Phenetics
be-
property allows the system to fulno
name
in the taxonomy of
definite notion
hunter,
majority
functions of
to
to
one
lived and
specimen
pheneticism,
-
called with
called it tradistics
homogeneous
have the
similar
more
for
a
users.
heterogeneous
tween them. This
fil
as
7
(interdiscipli-
common
all
be so, the system
to
within-group
a
meet
(e.g.,
users
of existence is
meaningful
and also
to
problem
example: "bird" is
taxonomist,
such
concentrate
above conditions,
that
artist. And
taxa
of
little
a
nary) reference system for
provide
kinds
that I shall
as
in
to the type of harm
according
can
contemporary
a
three main trends
decades
(indeed,
concerning that
functions and
particular
They present only
whose
living be-
kinds of classifications
many
so
of
diversity
activity
our
particular
of
logical taxonomy,
system,
biological
facilitates
requirements
system
the
arrange
We
etc.
clearer for further consid-
background
are
last
leading
to
function,
meiosis, mitochon-
above
I need
genetic systematics",
way that it
or
museum
commonplace.
make the
There
out-
Taxonomy
ings
genetically.
reasonable confidence where
single
feeded
stud-
insect consists of
structure and
mitosis and
equally
fossil
particular
species (in
been
ever
and
that any
eucaryote
tionary systematics"
a
have
studied
not yet
dria, DNA, particular kinds of RNA,
third,
need
very
including
Another func-
taxa
few insect
number)
knows
us
erations.
phylogenetic presumptions is
lined in Table I.
We
a
sure
chromosomes,
what
to
of
course
cytologically,
be
can
that
not,
or
Indeed,
their full
to
yet
(of
of
predict properties
known from
cladogram reconstructions.
The
detail.
And
Each
form.
sparrow
of higher taxa of birds).
in full
relation
a
say,
to
and
proposed approach
available
tion is
cells of
and
stronger
function
enigmatic
irreversibility
example
does
of
mode
one
loss
multiple
structure
than
more
I have
as
final decision de-
be
to
exist,
and
about,
knowledge
ied
I argue in favour of the
sounder arguments.
probability
that the
so
consider
we
does
much
taxonomy, and nomenclature
Phylogeny,
concise
direction ap-
one
often than alternative
more
when
Therefore,
ones.
the
be
to
-
Rasnitsyn
most
Of
the
and
means.
plicitly
way
three
to be
seems
most
approaches,
straightforward
Its claim is that the
constructed in
permits
larity
competing
the
one to
terms
avoid
of
system
phenetics
in
similarity,
subjectivity
its
must
in
goals
be
ex-
and this
the simi-
calculation. The methods of calculation
are
Contributions
Table I.
to
Zoology, 66 (1)
25
1996
-
Phylogenetic presumptions.
Scope
Term
Statement
General
"Rnowability"
of
phylogeny
until
Group analysis
the reverse
Among
Paleontological
sound
is
considered
reasonably
earlier should
contrary
evidence
related
the
groups,
be considered
is
unless and
inherited,
as
proven
apparently closely
two
fossil record
Character
should be
Any similarity
one
entering
the
unless and until
ancestral,
presented
analysis
Dissimilarity
analysis (polarizing
transformation
series)
Transformation
Paleontological
succession
unless
and until
Transformation
Biogenetic
series
of the
should be
sound
until
Of
If
analogy
there
series
should be
transformation
a
be considered
When
irreversibility
we
should
direction,
is
unless
and
otherwise
in
polarized
with the
agreement
character states,
decide
for another
the
a
the results
group,
unless
group,
one
direction
of changes as
cladogram supposing
the least their
or
in
polarized
to
record,
presented
and until
should
sound
presented
are
can see
prefer
series
are
respective
reasons
valid
as
data
contrary
Of
of the
serious
are
data
contrary
ontogenetic succession
to the
polarized according
character states in the fossil
respective
number,
no
unless
definitely realized,
to the
changes
and until
we
we
opposite
have
strong
contrary evidence
Of functional
Of two character
efficiency
reliable
Of complexity
A
a
The
vestiges
character
in
Similarity analysis (identifying homoplasies
Of
case
character
The
another
Of
weighted similarity
and
until
occurs
and until
showing
found
respect
any
as
in relation
apomorphic
have
we
of
signs
the
over
within
only
to that
and until
likely
strong
reason
modified
being
unless
latter,
a
and until
should
group
distributed
strong contrary
cladogram
unless
cladogram
conflicting
inherited
in
efficient
more
unless
to
from
strong
be
both within and
evidence
considered
outside this
appears
similarities)
most
homoplasies,
Of
a
apomorphic,
exists
state
in
unless
of conflicting
parsimony
corresponding to
should be considered
apomorphic
evidence
apomorphic
group,
evidence
one, unless
state
state is
contrary
A
one
considered
otherwise
another
Outgroup
structure
simple
more
decide
Of
be
contradicting
complex
to
states the
should
adaptation
as
implicating
the most
likely
that
similarities,
related
that
there
is that known to be
closely
is
and until
which
more
unless
groups),
the least
sound
are
should
reliable
in
and until
number
reasons
be
other
considered
cases
of
indicating
as
(especially
strong contra-arguments
appear
called
monly
proved
adequate,
the
Unfortunately,
developed
this
opinion,
has
less
though
onomy".
been
"taxometry"
collectively
to
been
proceeded
to
was
be
not
more
back
enough
methods of taxometry
level.
proclaimed
because
the
by cladistics
in
are
com-
methods have
level
unattainable but because
forced
far
the
or,
"numerical
its useful
tax-
never
In
has
my
been
phenetics
before
it
had
and
is
a
easy
objective
adherent
e.g.,
and cannot be
manner
to
any
Mickevich,
labour
to
particular
work.
Indeed,
weak
and
diverse,
before
numerous
determination of the
not
adequate strategy
made in
However,
approach (including
1978),
and
it
sufficiently
a
this is
a
problem
cladistics:
evidently
see
needs hard
evaluate what is the natural domain of
method
sides.
it
yet.
most
and
Phenetics
was
which
are
proceeded
"consumed"
by
the
its
far
a
strong and
in
more
this
way
vigorous
26
A.P.
of cladistics, but
discipline
a
conscious rather than
enough
render
to
adequate clustering method possible.
an
The above failure does
weak
important
ably
is
phenetics,
is
that the
of the characters
larity
sonably
the
shared and
the characters
cluding
to be
never
shared
not
thing,
important enough
in any
both cladistics and
kind of balance
a
that
are
in-
compared,
taxa
(and maybe
This kind of
is
the
Second,
characters
similarity
the
be
to
1979:
raises
process
genetic
process? Being unanswered,
leaves
no
phylistics try
do.
to
Cladistics
dently
cerned
cladistic
all
and
is
not
cladistics differs
depending
Platnick,
1979)
state
the
to
re-instate
tematics"
the
yet,
so
term
for
the
there
exists
cladistic
only
character
while
grouping
(Farris,
entire
It
487).
by
ones
ver-
central
classifications
to
tem
isomorphic
fective.
That
phylogenetic.
however.
to
is
and
There is
First,
apomorphies
the
an
diagram
or
system
alternative
is
nature
specifiable pattern which
branching
make
most
the
phylogeny and thus the
why
ordered
can
be
by
synapomorphy"
original
widely accepted explanation by Hennig (1966)
the taxa marked with
are
similar-
raw
recognized
are
the
latter
was
point of view,
in
a
single
by
problem
a
classification.
legitimized
solely
is
by
com-
acquired by the
not
group.
their
because
any
while
apomorphy
an
that
means
has
been
acquired
as
a
tetrapod
apomorphy
an
is
have
(Snakes
"legness"
but the fact that the
ever
of
This definition
unloosable.
considerations
with groups
such
nically
polyphyletic
fined by
is
particular
acquired (a fact of
are
still
inadequate
to
split
to
by
groups
above
split
so
as
of
to
other
Most
see note
procedure,
to
Many
they
are
homoplasy,
or
de-
respec-
subgroups marked
monophyletic
below).
This
paraphyletic
are
Queiroz
taxa
14-16)
may
&
incomplete
serve
only temporary
knowledge
resisting
cannot
be
Donoghue,
groups
at-
ancestral in respect
as
of this kind could be settled
as
meets
all demonstra-
hymenopterous
as
rather,
or,
policy
i.e., those which
Hymenoptera
ing the difficulties
our
9
(De
fossil
above (pp.
cases
or
give subgroups that
holophyletic
as
tested
(that is
in metataxa,
problem
paraphyletic
whether
on
of both sorts into
apomorphies
the
termed
are
groups
depending
general cladistic policy in that respect is
holophyletic,
a
lacking apomorphies. Tech-
symplesiomorphy
The
tively.
1990b).
termed
what the central
group).
The above
ble
sys-
most ef-
represented
hierarchical
that
con-
explain the cladistic approach, because there exists
a
splitting,
inference,
putative
is
this
Despite
"Phenetic
according
1979:
apply
a
indepen-
Synapomorphy
lost,
as
state
state
of
history
sys-
using the
by grouping according
to
to
gained general accept-
and,
phylogenetic
and
transformed
continue
field).
stable
a
"phylogenetic
its
to
not
that I shall
statement:
constructed
term
Hennig's version,
(the proposal has
ance
ity,
original
"cladistics"
term
sion
the
denote
to
character
that
De
be
synapomorphy.) This
not a
Queiroz & Donoghue (1990a) apparently proposed
can
but
phylo-
question
group and thus characteristic
treated
legs
the
proposal
character state
a
the
not
that I shall be
be raised is
however,
correct,
Hennig,
e.g.,
significantly,
so
on
to
(taxon
a
pattern into
latter.
means.
as
so
members of that
only
universally
appreciation of different students (cf.
1966;
question
claim
species of
stem
the
analyse
with the
only
The first
lost their
meaning of
to
way
if
pattern
of classic cladistics,
monly defined
to
precisely what
the
sets
This ap-
what could be the gen-
words,
the
derived
hypothesized
secondary.
what
sets
evo-
hence claimed
erating
evaluate it
This is
thus
pattern is
phylogeny
of
of
knowledge
our
pattern
The
question:
a
by sampling
internested
classifications, is
our
538).
nature? or, in other
synapomorphy)
way.
like
and
primary
proach
Third,
hierarchic
(Platnick,
be estimated
can
finding replicated,
lutionary history,
from
character
The
pattern
and
directly available but
justify attempts
to
rea-
studied
yet
not
possible indirect
be
qualifies
properties
the
by
not
investigated).
rather abstract
must
than that. On the
It is
and
studied
with simi-
dealing
more
sort.
characters
all
most
what the
as
similarity which
is of another
system
between
same
were
not
for
quest
the
however. Prob-
studied, which
but
numerous
contrary,
be
to
taxonomy, and nomenclature
Phytogeny,
of synapomorphies.
similarity
the
precisely
not
seem
not
of
needs. Pheneticists
system
the
point
important
more
by phenetics
a
far
not
chance determination of
a
-
Rasnitsyn
(this
as
is
examples.
by
a
suppos-
result of
especially
Contributions
in the
tempting
is
rance
lution,
not
as
Besides,
phyletic
group
the
to
further
in
evo-
sure
in
refuge
prohibitive
all
(De
to
from the
be
cannot
&
Queiroz
a
split
any
Donoghue,
Some
students
by claiming
here
verified
as
such
verified
finally
to
in
some
another
and
some
This
is
not
all
at
precisely equal
(minus
must
A
Moreover,
that
himself.
phylogeny.
cies is
the
not
dent
ties"
to
by
which it
into
two
by
its
more
taxa,
marker of
while
species is
appeared
after each
the
products
of
show the
as
only
one
process
must
as
event.
be
tors,
ity
that di-
that gen-
just
a
a taxon.
having
Even if
event
its
does
ancestor
considered
do
can
of
it
through
divergence
similar
position
280-281)
to
Since
with
proceed
sults in
invert the
to
sequenced
used
only
is
value,
sides
not
classical
of
by
in
special
appearing
the
of a
event)
in their full
one
phylogenetically
step
are
be
usually
(during
form, for
complex,
by
at
least
study.
of
equal
be-
example,
fully placed
in
a
hardly available just
a
most
not
ternested taxa created
a
For
being
(no
sition of
precisely
in
even
are
phylogenetic
study.
of
problems for the identification
a
re-
and deficiencies of either
not
as
This
unattainable
are
cou-
easily
can
fully sequenced
framework of
cannot
dis-
gain
divergence.
for
they
advantages
one
that
approaches
two
that
necessarily
not
a
being
the
A
wasps.
explanation of taxa".
is
event
apomorphy,
why
in
study
without modifica-
that "descent
apomorphies
They
used
of the
apparently
to say
counting
proach.
already
allowed Nelson (1989:
impossibility
sequence,
with
tool of the
of aculeate
deserve
to
the
ultimately
taxonomy
in their value
but the
a
has been
apomorphies
comparable
as
in
his
proposition
as
apomorphies,
complete knowledge
That is
theory.
system
approach
say
gain
advantages
makes
compari-
unaccompanied
a
of
array
a
limited
the claim
divergence
a
In-
correctly.
at some
to
however, is insufficient
tion,
pled
This
cladistic
(1975)
and
phylogeny
arrive
Indeed, contrary
events
The
Brothers
ances-
in
possible
seems
sequencing.
(possi-
less evident
approach
consider the
for
only
expect
above,
the
If
some
to
approximation.
and
definable
objective
as
differ from their
important.
not
cladogram topology
cladistic
to
phylistics
have any theoretical possibil-
even
with alternatives.
son
seem
reasoning
of
we
the
see
than that.
more
identify
to
to
(but
respective cladogram,
the
as
the
becomes
usually opposed
of
by
lit-
gives
It
it
Firstly,
differ-
a
meant
to me a
seems
system appears
grounded
do not
we
This
of view is that the
divergence
it
spe-
communi-
becomes
point
a
indepen-
an
of
way
slightest difference from
apomorphy acquired),
system
speciation:
reproductive
This
as
only by
duration of
origin
divergence
a
less
made
(basic, central)
to be considered
of
taxon
problems
between the
apomorphy
ancestral
clade
apomorphy
divergence and, by inference,
Another implication of this
is
species
already
processes
makes divergence itself the
of
solid
temporal
two
owes
or
total
not
is
nature
the
bly many) species
again
8
it
The last observation is
stead,
central cladistic
definition but
more
though
proposal
advantage.
as
Secondly,
well
as
of the
only by apomorphy)
1966: 66).
(Hennig,
erates
below).
himself actually
least consistent.
at
syncretistic
claimed
degree
existed
single
a
reproductive community, and the
vides it
monistic,
birth
the
theoretical
isomorphy
an
"The
determined
one
clearly
by
than
He considered
taxon
of
sur-
researches.
ever
step has been
implication of another,
be
species.
more
definable
that is
same
holophyletic
every
insuperable
by Hennig
statement
aware
because
surprise,
ancestral
necessary
a
can
from
species
do the
to
species
rigid. This
legitimizing
Popper
the number of terminal
own
(taxa
statement
are
paleontological
a
to
impose
we
a
for cladistics unless it makes the
and
system
be
identified in number in
are
ancestral to
species
to
hypothesis
because
one),
have its
seems
scientific
species
of ancestral
number
Thanks to
populations after giving
detailed
more
ignorance
can never
1973a).
and then
species,
species
preventing
of its
Such
again.
all.
at
factors
biological
viving
no
in
refuge
stem
(Nelson,
it became clear that
no
take
again
that the
double
a
and
1990a).
but it is
strange,
1984, for
Sluys,
see
Hennig
his deviation rule). This
tle
stem
species (but
view of what
ent
non-holo-
of
us
different
de-
necessary
splitting
leads
which
species
way
is
above,
obligatory
this
27
igno-
deciphered
stem
The
fossils).
inescapably
taxa
1996
of
made
cladogram being
tails.
of
-
(1)
however, because parallel
case
secure,
we
66
Zoology,
to
Brothers' ap-
hypothesized
single
speciation
apomorphies
unitary. This
of the
successive
complex apomorphy.
As
as
are
causes
system of in-
steps
in
acqui-
has been dis-
A.P.
28
cussed above
not
able
regards
as
do this
to
phenetics,
for cladistics
tive manner,
so
certainty
classification.
to
that the
Brothers' version
compared
tent as
seems
consideration both
with
distics
be
with that of
yet
as
more
other
un-
hand,
richer in information
Hennig,
amount
of
advantages, cla-
also has
versions)
considered
later
resist
to
shortcomings.
in
on
comparison
phylistics.
phenetic and cladistic approaches
take
relatedness. To reach this result, the
is
to
and
be
phenetically
be
must
monophyletic.
has been termed
marenko &
there
with
similar
bers
of
to
each
in
of them
the
same
This kind
of
taxon
close
to
long
cover
is used
or
means:
subordinate
not) of
phenetically
than to
neighbours
The
It
looks
definition
eclectic,
criteria of
independent
a
more
independent criteria
tion.
Eventually
to
is
a
bad
define
eclecticism is
cases
real
or
to
we
are
that
ways
able
to
result
use.
two
So
The
continuum
such
a
were
rodents
of them
phyletic
are
potheses
there is
deepen
and
in
uses
as
con-
should
clear whether the
We have
real
probabilistic
case
seen
phylistic
is
result,
polyphyletic.
we
is
also
When
should try to
escapable.
in conflict when
we
falsify it,
a
encounter
as
we usu-
can
taxa
practice,
possibility
to
level
similarity
fully
we
easily
are
certain of
rely
more
why
reassess
by monophyletic
usefulness.
fields
ogy,
I refer
such
as
larity,
total
ones
as
well
as
to
spite
of their
without
our
array
(see the
(monophyly)
us
to
assess
those
relatedness is used here
simi-
on
is used
similarity
including
next
evidence that
as
relies ultimately
approach
of characters
The
replaced
parasitology, mycol-
I consider this
allow
be
significant loss in
many procaryote taxa
while relatedness
known.
cannot
a
traditional
undisputable
various kinds of parataxa in
paleontology,
details).
to
when
however,
in
polyphyly, namely when they
the traditional
of
we
hy-
taxa studied
be
it
theory,
In
likely.
for the
never
In
intuitive criteria. That is
examples,
are
taxonomy retains
two
in
versions
and
to
mono-
where both
in both the
can
separate
similar
results and escape the conflict.
this eclecticism
are
1989).
cases
equally
of contradiction
section for
criteria
We
the
form the
to
phylogenetic relations of the
and
of phylistics
find
danger there,
exhaustible.
In
will be
but nevertheless all
(Wilson,
to
knowledge
our
seemingly eclectic,
alternative
considered
also
as
convergently
(Gidley, 1912),
will appear
no
established
only
Glires
taxon
orders.
of the hares and their rela-
case
once
now
quite possible
indirect clue
eclecticism
the
in the
Lagomorpha,
true
and
discarded in fa-
(monophyletic)
hypothesis of polyphyly
order
on
cladistics.
Indeed,
the
They
in
same no-
we
natural
tives.
the
it has been
eventually
as
19th centuries
elephants, tapirs, rhinos, hippos,
more
falsified,
of
and
Ephe-
Pachydermata
18th and
either aspect of the system, and thus
because it
criteria
the
mammal order
seems
similarity
and the
only apparent.
escape
in
to
make it
above that cladistics is also
that
one
shall find the
we
flicting and thus impossible
inquire into such
one
Plecoptera,
the
Pso-
Isoptera,
not
however,
under
in
Neuropteroidea
Odonata,
during the
use
cases,
There
approach
the
comprised
mem-
relatedness.
An eclectic
in
until
of
other
is
(Pono-
the traditional understanding
"taxon".
combining
continuum
Monophyly
being
other continua.
any
term
(branching
homogeneous
and
Continuum
note 9.
any of its direct
clear and
the
1971).
Rasnitsyn,
taxon
at
monophyletic
explained
as
continuous chain
taxa
a
phylistic
to
and
similarity
within-group
between-group heterogeneous
time it
both
in its attempts
into consideration both
directly
of
advantages
the
Similarly,
vour
combine
from
Mallophaga,
related. For
only remotely
still
later
meroptera, Panorpata (= Mecoptera), and Trichop-
pigs,
to
be
to
the
was
which has
which
groups
also the Embioptera,
sense,
coptera,
to
tries
comprise
to
apart
name,
was
Phylistics
used
order Neuroptera,
instance, Sharp (1895-1899)
tera.
Phylistics
the insect
been
testing. That
sophisticated
more
have been found
current
Some
encountering conflicting data.
polyphyly will show similarity by failing
with
case
taxonomy, and nomenclature
Phylogeny,
of
cases
this
undoubtable
-
do when
ally
long
con-
for it takes into
below).
see
its
both its
(in
They will
the
relatedness and the
change acquired (but
Together
On
are
fully objec-
a
adds
approach
that of
to
we
in
Rasnitsyn
as
an
in the
yet
un-
similarly
to
show how apomorphy is used in Hennigian cladistics and
divergence
events in the
Brothersian
one.
Contributions
As
a
tic,
eclectic):
timately
and
all three
result,
not
Brothersian cladistics
cession
of
the
uses
divergence
relies
morphies. Phylistics
the
(in
indicated
(as
prove results of
the
list
and
apo-
similarity
on
and
characters)
by apomorphies)
we
return
can
and
advantages
In
phylistics.
phylistic
satisfied
is,
with
ist. In this
use
any
dent ones, it is
exceedingly
cladistic
mately
the
relies
states.
the
As
ing
It
comparison
to
a
nearly
tive.
That
is
("plesions")
that
are
less
complete
One
relatives.
that the
enough
in the
why
many
to
cladistics
ulti-
solely
infer-
on
succession
of
system
hypothesiz-
series
cladistics
is,
has
are
to
groups
be considered
as
are
restric-
parataxa
despite
for
the
characterized
their
fact
well
(normal taxa)
makes the domain of
advantages
able
with
of
phylistics
seem
to
the
more
the
taxa
to
of
it
in fact the
tained
in
the
gram, and
metrical
as
the
to
bits
3
rank
a
result,
of the
cladistic
the
is
information
phenetic information,
only
is
that the
the
and vice
ing
one.
ranking
That is
is the
most
we
cession
a
in
time
practice
virtually
that is
of
was
supposed
divergence
found
bit
a
Unless
amount
can
of
introof
amount
expense
a
of any
only by
arrangement
with its
to
us-
highly divided
informative
original Hennigian system
number of ranks
is
the
sym-
versa.
system, especially
why
a
not
added.
additional
at
con-
dendro-
with 4 terminal
Additional information can be added
a
that
of information and
an
of
of the
the total infor-
instance,
dendrogram
system
or,
topology
respective
information
not
such,
as
length
se
per
correlated similarity and relatedness,
into
sys-
content.
description
system
the
on
ranking is involved, and beyond
duce
non-
phylistic
and in the
such it is limited. For
dichotomous
unless
similarity of
The above dis-
but in its
of the system
geometry
contains
taxa
cladis-
for
problem of informa-
solely
dendrogram
content
oper-
information that is
As
internodes. As
dendrogram
mation
se
background.
respective
the
why
the
to
se-
Phylistics,
richer in its information
contains information
the
to
which is connected with the
hierarchic system.
a
concerns
in its
we are
into its system the
incorporate
stored in the system per
rather,
has
of characters.
characters. That is
content
find
we
result,
add the information about
to
potentially
cussion
a
that cladistics
set
There is another side
tion
As
of information available
involved,
polarized
tem is
so
incomplete
amount
tics, and
we
polarize the whole transformation
however, is able
total
fullest
transformation
characters
new
This
information
more
yet polarized.
not
than
is.
the
because
of the
polarization
characters
phylistics)
however,
attainable. The
to
an
related-
informative
more
inference,
by
ries that is available,
such
narrow.
putative
(and,
of all
origin
thus
disputable,
available,
never
characteristics
orthotaxa
This
as
highly
require-
more
available
do so,
is
has
consider
than those
to
much
whose
groups
system.
phylistics
is
with the series
more,
is that
while cladistics
forced
respective
phylistic
Other
is
it
is
the
phylistics.
for characters that
ments
and
cladistic
lower level of
"omnivorous"
"stenophagous",
evi-
this,
to
is based
the
result,
when
more
knowable
Another side of the above result
is
the
is
respect).
data
primarily
evolutionary
a
contains inference of
in
in this
resulting system
that is
interpretation.
concerning
character
both
relatedness
from character
ences
claim is
and
information. Farris
stores more
specifies
and
phenetics
ex-
phenetics
hypothetical. Opposite
system
on
no
reliability
as
only
uses
that the
possible
when
phylogenetic
and because it
they exist,
alone
phenetics
reliable
on
such. It
may be
as
empiric
as
like
just
ultimately
reasonable
any
important
most
it relies
similarity
it will be
case,
of cladistics and
the
that
characters
on
raw
fact it will be
will
is
data of
phylogenetic
not
available
be claimed
can
similarity
this in that the cladistic system in
to
form
not
of compara-
question
opinion,
my
that
the
to
disadvantages
advantage
similarity,
It
fullest
its
ate
Now
(in
(1979) objects
about
alone.
tive
and, therefore,
ness
less evident. It
or
records both
acquire
similarity
on
important
phylistics
form is
remember phe-
ultimately
that
im-
of charac-
sets
be less
uses
to
straightforward in relying
and
proximately
suc-
the
sequence
ultimately
complete,
netics: it is monistic and
both
them.
the
on
apomorphy,
study of incomplete
To make
ters.
to
array of
complete
most
relatedness
acquiring
ul-
events,
identify
ultimately
events
(monis-
relies
divergence
mark and
to
29
cladistics
of the
relies
of
events
1996
-
consistent
equally
are
apomorphies
uses
(1)
Hennigian
succession
on
66
Zoology,
to
incredible
reflect the
events.
impractical,
suc-
However,
and
now
it
abandoned in favour of traditional Lin-
30
A.P.
nean
ranking
in
is
completely
a
transforms
tic
1974)
that
Artificial
comb-like
in
a
a
sequence
group
of
with
the
an
or
basic
crown
and
early
pair diverged
most
lately
still
group
nig). Indeed, the
by definition
paraphyletic
a
It has been claimed that
cladistics,
contrast to
by
is, paraphyletic
of
not
which is
mean
a
only
1979), that
the
absence
nor
(p. 26)
charac-
a
hypothesis concerning the his-
absence of
apomorphy
by
character
a
Absence of
taxon.
of snakes:
apomorphy does
an
Lizards
character.
a
they
comes
defined
exclusive.
or to
criteria of
holowe
problem
of
have
also be discussed here. The problem
to
be
disregarded
category
is
rived from
the
grass
As
(Poaceae),
species,
typically
to
certain
genera
be
The
first
than the entire
for
being
example,
Tribe
300 of which
de-
while
Tzvelev (1993),
the
in the tribe consist of
higher plant
not
taxa.
As
result,
a
to
rely
a
them
on
(as
a
in the
I
a
as
vote
which
taxon
taxon),
neither of the two,
in the traditional
taxa
sense
ultimately
phylistic taxon), and
are
polyphyletic
so
not
choose
cladistic
a
their
are
to
a taxon.
as
as
dif-
to
to
which
have
we
group
those which
problem
fect these
(the
of
and
hybrid
symbi-
to
hold
true
for many
taxa
do pose
does not af-
problem
only cladistics,
that
this
However,
phylistics,
not
parent species
are
paraphyletic
problem
nor,
taxa
of
concerns
phe-
course,
netics.
Other considerations relevant
seems
its
taxonomie
leading
the
Nevertheless,
own
advantages
and
between them does
my
opinion,
be inferior
the
seem
a
of
prognostic
of
system
to
consider the
points
which
phylogeny,
cladistics, which rejects
taxonomie constructions with
I believe
tion somewhat
tem.
Therefore,
this
more
I
makes
shaky
a
two
are:
fails
to
and the
too
large
produces
high hypothetical
the whole
than the
construc-
phylistic
expect that phylistics
the winner of the contest.
In
demonstrate
fraction of available characters and also
content.
has
The balance
These weak
phenetics,
power
obtained
equal, however.
and cladistics
phylistics.
shortsightedness
the
comparison
already
disadvantages.
not
phenetics
to
the
concepts certainly
result
deficiencies important enough
to
to
conclude that each
sufficient to
fastidiousness of
hybrid species.
ancestor!)
common
in
caryo-
symbiotic
chain between both
definition.
by
and
themselves but their ancestral
use
Triticeae
are
taxa
complete
and their
hybrid
for taxonomy. This
of
intergeneric hybrids, and the
A similar
pattern is claimed
other
enough
organelles
reviewed by
family
consists of 500
largest
important,
exotic.
less
no
cell
is
common
characteristic,
very
higher plants.
the
(their
by
(either
future: if we accept
Nevertheless,
a
of the three
must
taxa
originally free-living organisms),
is
latter
are
something
represented
Eucaryota
taxon
as
they refer
respectively),
paraphyletic
sense, and not
lack the
legs instead.
symbiotic and hybrid
because both kinds of taxa
or
be-
mutually
not
are
for
strictly holophyletic (as
exclude
may
thus
symbiosis.
or
of taxa
history
aspects
shall avoid
we
holophyly
availability of
favour of the
if
terminal
taxon, which
surprise,
no
their future,
between these
exist.
The
This is
mutually exclusive.
or
time,
of all
hybridization
and
ferent aspects of the
is either
more
synapomorphy(ies) gained
of the
Hence, polyphyly
in
by
same
holophyletic
a
ancestor
in that
the
by
direct result
phylistics,
and
taxa characterized
it is shown above
apomorphy is neither
of the
explic-
be included
past,
the
and all descendants of that ancestor,
easily
a
At
definition of
common
polyphyletic,
are
is crossed
ancestry.
the
nearest
a
taxa
otic taxa.
characterized
taxa
ter state but rather
tory
Hen-
Ax, 1985) is
e.g.,
taxon
phenetics
use
apomorphy. However,
that
by Hennig)
sensu
absence of characters (Platnick,
an
from
in cladistics.
itly prohibited
in
*taxon
*taxon
(see
stem group
sim-
popular concept of
of the
minus
a
a
polytomy.
deviation
striking
more
from
in form than
unresolved
an
claim is the
(taxon
derived
being
complex
(equivalent
group
stem
we
from the one, which is
by
more
cladistic
subtaxa,
hybrid
boundary
line of
satisfy
can
taxon:
most
their lower
single
a
Cracraft,
convention
and
symbiotic
Nelson's
rank listed in
same
than
taxonomy, and nomenclature
Phytogeny,
-
because
they
diverged
representing
Another and
eclec-
arbitrary order). However,
arbitrarily
cladogram either
ple comb,
a
it
should be transformed
taxon
distinguish this list
sequenced
like
as
thus
an
1973b;
of one and the
listed the last (in
cannot
into
accept
can
cladogram
taxa
the first, and the
listed
the
(Nelson,
of little help. We
and
way,
conventions
sequencing
are
a
arbitrary
post-Hennigian cladistics
concept.
phyletic
The
1979). This form of ranking
(Wiley,
contains little cladistic information, being used
it
Rasnitsyn
sys-
will appear
Contributions
66
Zoology,
to
(1)
1996
-
31
1982, 1986,
Rasnitsyn,
Nomenclature
3. Identical
Nomenclature
how
monly
be
to
simple basic
the
use
principles
of conventions
of
names
field of
a
is
the
appreciated
and
in the Codes (e.g.,
basic
only
or
basic
as
ICBN,
principles
formed
hierarchy
the
ferred
of
principles
their
to
own
not
in
con-
is
below,
distinction between the
useful,
seems
and
names
personal
considerations
so
will be
that
and
experience,
on
the
seems to
re-
taxonomy-
for various Codes differ
the follow-
rules
of
made in favour of that
5.
the
long
and
syntax
often
are
and
all,
and
plain
they
are
judgment,
regulation
is
not
which
largely
in
details,
particular
be
Ar-
standardized and rank-indicat-
a
taxon
of the
The
last
name.
It
the
on
taxon
base
a
presents
language,
for
grammar
the
including
name,
the
among
the
principles is
princi-
Articles 4-
(ICZN
rule
on
taxonomy-independent
principle of superior priority
International Commission
case
any
of the
Nomenclature which
on
the
to
contrary
Code
(ICZN:
xiv).
Taxonomy-dependent
three
are
principles
fully
correct
cases
infringing
not
upon
be made
(ICZN: xiii).
taxo-
subject
This state-
because of the existence of
principles
that
validate
only
in taxonomy. The statement should
strictions
In
clature
explicitly
zoology,
is
see
the
area
ICZN Article
up
1).
a
subject
to
nomen-
level
superfamily (for
Exclusion of
from the scope of the Code
it is
principles
of
suprapopulational
to
subspecies
of
seems
for another
higher
to me a
de-
taxa
mistake,
discussion (see
to
respect
on
The
the
it bears is
name
cle
61a).
In other
the system
type. To
assess
alternative
by
tifies
ters
are
of
system
to
as
a
matter
can
be
we
of
how the
Arti-
be introduced
referring
to
its
should consider
consider is the way
1974,
a
its
to
class
ones
1987).
doing this;
-
no
a taxon
this decision
defining
(Ghiselin,
ties
thing
taxon
a
objective
change" (ICZN
ultimately only
by referring
taxon
the Code
which the application
determined,
words,
to
the
provides
type
demon-
imposed by
possibilities.
The first
a
most
According
boundaries of the taxon may
into
and
the restrictions
nomenclature.
name-bearing
of name-bear-
principle
important
most
standard of reference
imposed by the present Code".
restricted
groups from
re-
taxonomy-dependent principles.
concept.
is the
taxonomy
"The
probably be completed with the words "beyond
though
must
(ICZN
11,25-34).
6.
ing types
need but little comment,
must
restraint"
or
taxonomy-dependent
tails
earlier
zoo-
discussed here first.
"The Code refrains from
nomie
2.
(iii)
animals and
underlying principles.
These
ment
ho-
and
homonyms,
proposed
array of rules
strative with
to
group
(ii)
taxa
to
re-
make little dif-
Taxonomy-independent principles
1.
be
among compet-
name
or
of binominal nomenclature
ple
6,
of
Principle
form of the
1. The type
at
species
genera,
belonging
taxa
Selection of the proper
There
if
(i)
(and between) higher
ones, either synonyms
ing
they
further
arbitrarily
the
different
to
not
must
plants.
4.
can
based
are
nomenclature. This
logical
discussed
is
homonymy
concern
between
homonymy
ing
a
respectively.
ones,
Because of my
ference,
is
as
In
to
belonging
monymy with
to
of
not
as
taxa
ticle 23). This is the principle of priority.
eas-
accepting
species.
taxonomy-independent
as
dependent
ing
genus and
by
different. The
essentially
deserve
fully
type principle,
kinds
principle
so
1991).
Article 52). The scope
(ICZN
names
principle
names
of bino-
conventional and
with any kind of taxonomy
ily coupled
evolution-
particular
a
the Linnean
minal nomenclature is
two
by being
valid
as
stricted
of these
some
conventions
true
inference of
by
theory. Indeed,
trast,
addition,
reflection of a particular taxonomie concept,
a
obtained
ary
not
are
1985) but
1994; ICZN,
In
principles.
mentioned
are
used
of the
com-
exactly
explicitly formulated. Some of them
not
on
details and
not
all
not
are
It is
taxa.
sophisticated
This
principles.
for the
case,
collection
a
and
to create
felt
is
1989,
of different
names
the
provided
and
to
characters.
not
There
simplest
multidimensional
that the
merely
are
is
introduce
This
diagnostic
several
a
iden-
charac-
possibili-
combinatorial
matrix
with
each
32
A.P.
compartment corresponding
version of this
A
taxon.
sometimes
used
matrix with
an
is
particular
a
to
nation of characters and thus
harbouring
by
of characters and left col-
row
sections.
This
of
sort
is
system
quite
aims of the
general system of organisms,
It is
inflexible:
too
torial
system
addition, deletion
any
will affect many other
is
tem
not
very
fit
to
not
agree
improve
cannot
we
locally
It does
a
particular
a
alternation of
or
That is
taxa.
even
common,
and
as
why
more
by
flexible
with
identification,
highly
ranked the earlier
system
acters
are
since
enough
a
&
character
a
such
a
primarily
proposal
for
meters)
other
a
popular
who
(1923,
few
in
which is able
to
the
fructification.
explicitly
called
who
characters
for
a
(para-
I
a
charge
position
of re-
from these
The
task
as
key
proposed
applied
to
characters
to
(the
Lubischew
by
and
taxonomy,
resolve
it.
However,
solution has been found, albeit in
discovered in
a
sought
for it.
convinced antiselectionist,
The field is the cladistic
claim that the characters
considered
system
of
as
with
by only
as
seen,
incan
characteristic, viz.,
the cladistic
an
a
type
cladistic
by Sundberg
Pleijel
&
practical application
is
that
cladistic
taxonomie
a
does
approach
as
about
thought
of the clade in
pomorphies
of all
synapomorphies
these
the
as
hypotheses
have been
are
acquired
collection of
believe will be
an
easy
a
but exist
attach
to
job
All
only
states
event.
So
taxonomie
a
I do
hypotheses, which
to
of
sum
inclusive clades.
divergence
told
effectively being
name to a
ances-
which character
what
at
that
result from direct ob-
type specimen does,
concerning
a
to
us
(taxon)
question plus
not
as
collection of syna-
a
more
do
synapomorphies
servation,
as
only
as
a
no-
have
we
permit
not
real group
a
name
as
could be identified and studied. Instead, the
is
This
concept.
clade. However,
ancestor
self-evi-
that
proposal
proposal
ancestor of
an
De
However,
draw the
to
(De
afraid that this will be
am
Their
refer
they
ancestor"
307).
to
of any
aware
and I
effort,
think about
not
taxonomists
persuade
do.
The
its
class,
as
members,
native.
The
group defined
a
has the
therein) is
suggestion
taxon
ring just
rity
thus
must
to
and
enough
to
by
individual
individual (Ghiselin,
to
as
the
more
taxon
the characters of
its
logical
consider the
taxon
quite natural.
As
an
name.
For
this, it
the
taxon
than
one
name.
of them
to
must
dication of its
an
member/part)
concept. Any part of
an
by
integ-
developed
be born and
a
a
have its
to
die. It
equal enough
might pretend
This makes
the ostensive definitionof
an
refer-
wholeness
spatiotemporal
permit
as
individual,
then be able to be introduced
its
alter-
1987, and bibliography
1974,
cannot be broken down into parts
that
its central
organisms and, by
one
a
false.
the location of their taxa in the
system,
be best determined
a
Ironically,
field which Lubischew,
to
tag
This
characters of
am not
menclature should attach
to
parametric system,
a
their
been taken
difficult task.
we
and the
the Mendeleev system.
other than where he
ference,
ap-
system of
most
predict all important
hopeless
meanwhile
was
this
has
if
same
should abandon the
step
of their
tor
deter-
atoms
Lubischew himself failed
it
char-
nuclear
discovering
involved
parameters).
place
the gen-
rank
system)
of
rank
highest
elements in
would result
seemed
It is
in somewhat
based his
1966),
like
characters,
taxa
more
key.
that could determine the distribution of all
mines the features of
spective
key
taxonomists
among
characters
on
iden-
taxa
as
highest
the
shape
to
has been formulated
Lubischew
search
the
next
from
the
1990:
Gauthier,
dent inference
taxonomy
sys-
from
stemming
refer to
to
into the system.
synonymous
and Gauthier failed
being
good
as
are
Queiroz
for
evident arbitrariness of
few of the
Linnaeus (1751),
plants
This
(that
has remained
proach
not
"names
been
atten-
need its
not
enough
a taxon
Queiroz
in the
occur
rankings. Nevertheless,
modified form
by
Indeed,
introduce
to
combina-
standard
characters
because of the
the character
in the
as
"typified"
draw
to
does
system
the
meaning
system have
want
because it is
clades
taxon,
with the
one
they
it is
quite handy, although
eral
is
system
ranked characters,
fact that the
the parameter
(1994).
tified
for
be
only
Here I
however.
tool.
A
the
to
becomes
thus
definition. The
of the cladistic
potentialities
taxa to
with the
identification
an
Relatedness
parameter by
discussed above.
tion
inter-
really simple
sometimes
useful.
the
at
Lubischew
and
taxonomy, and nomenclature
Phylogeny,
their relatedness.
rectangular
a
-
of taxa, and with character states
umn
combi-
separate
identification table
an
taxonomists
upper
a
-
Rasnitsyn
to
so
inherit
difference between
individual
and the
use
(direct
in-
of the type
individual may be
equally
Contributions
intended
used for the
how
not
select and
to
to
-
(1)
33
1996
and hard rules of
definition,
the taxonomie types
use
introduce individuals into
to
necessary
66
Zoology,
to
a
are
sys-
tem.
"Taxon" therefore is
it
if
vidual,
is
the
The
paradigmatic
wholeness based
This also
parts.
due
species,
to
pressed by
in
what
I
that
flow
Ghiselin
(1981)
phylogenetic
is
limited in
are
1988:
hypothesized
has
a
tegral.
and
son,
so
scendent
the
a
problem
species
distribution of
asymmetrical
share
common
a
viewers for
precise
description
system.
It
displays
and
exist,
die,
its internal
here
history:
but it
is
be,
is
rything,
a
history
the future is
is:
a
it
correct
my
give
birth
it is
to
be
to
not
history
in its
I
role
a
taxonomie
itself. The
characteristic
-
is
a
can
for
in-
clade it
meaning
this is
is
eve-
appar-
of the cladistic
view of the system.
However,
not
the
necessarily
While
ality.
time,
(see
the
is
name
the
dichotomy
existing
options.
does
In-
to
and
is
taxon
given,
able
is
as
between
of all
geographical
be
the individual appear to be
two
space
whole-
its
possessing
In
the
two,
extremes
is
taxon
a
rather it
or
because the
them,
its
after
appreciated
individual.
features of both class and individual,
gap
it
lose its individu-
Because of this
to
an
in
important,
more
not
extent in
1980).
so
retains its wholeness
a taxon
an
not
what is
to
multidimensional space
Schram,
e.g.,
ness,
the
is
of Homo
parts (populations
what is
and still
evolving,
in
class and
of
single
a
spectrum.
the
Consequently,
neither
is
to a
real,
proach
its
phylistic
is well suited
handling
a
identify
stead of
can
to
be
a
unlike
continuum,
as
a
taxa
and
If
compartment
an
individual is
in
which it is
possible
depends
an
a
overall
possibility
integral body
continuum
similarity)
its
and
matrix,
It is rather
change
extinct).
is
a
possible
by
delimiting
continua,
and
by
are
the
not
easily changeable
knowledge
gaps
in
its
to
integrity,
characters
of
it
con-
cloud for
a
dismembered
It is
class
a
shape and
in
existence of
in-
of particular charac-
character
two.
thing until it
(becomes
ap-
similarity
class,
a
(in
terize the continuum both
and
the
on
elements, but nevertheless it persists
appreciable
the
agree-
phylistic
introducing and
individual,
the
to
taxon
system ultimately by using
means
tinuum is neither of the
is
most
it is the continuum that
unlike
by
not
The
usage.
an
and
integrity,
is
solved
A
in
continuum
ters.
posing
being complete.
identified in relation
other
a
a
sense
spatiotemporal wholeness
using types:
taxa
above
the task of
to
be introduced into
type. Indeed,
to
far from
the
individual. It is
true
a
be best defined
to
in
taxon
nor
cloud whose
though
with
ment
class
a true
Both criteria
class-individual
exhaust
develop (evolve)
can
in
taxon has
species), and,
to
mem-
Alex Rasnitsyn
time, similarly
same
a
and it has
specimen [example]
a
charac-
possesses
diagnosis,
Ghiselin,
to
individual,
relation
class it
a
its
by
to
At the
sapiens).,
can
evolu-
past history
nothing
re-
segregate
single
of
not
born,
how "brilliant and
the
to
a
I
another
to
only
as
part but also
a
seems
kind
special
can
(contrary
similar
cladistic
all-depressing
matter
next
in-
and de-
father but
of the
bers
an
history:
common
what
taxon
world,
comparable
fairly
to
into subclades (subtaxa).
no
to
as
equally
ancestor
essence
can never
the
the
for all of
dichotomy]. This
clearly
determining
common
ently
this
exaggerated,
nothing in
fluential" in
could
very
cladistic
tionary invention,
as
me
of the
contents
an
are
my
The only thing available
taxon.
see
a
taxon
of my anonymous
to one
showing
of individual
with
history
[thanks
son
that
sufficient
here too,
as
fills the
difference results from
species. The only
with my
well
attested
ters,
features of both the class
possesses
notion intermediate between these
con-
301-302).
as
both its
to
only
in
related to my father and
equally
is
for
scope,
taxon
individual:
characters,
sup-
species
actually
integrity
the ancestor-descendent transitions
am
is
validity
lineage and the respective
individual. There is
I
its
has
for
true
'nondimensional situa-
(Mayr,
lineage
consider the
be
divergence
concept
populations
with each other"
tact
interaction of its
proposed
called its
where
spatiotemporal
to
the
for the
con-
individual,
an
(Mayr's biological
species
have
is,
continuous
been
model
biological
tion',
and
integrity
on
has
gene
of
case
in
comments
e.g.,
the fact that their
The
concept).
"The
(see
and the issue deserves further
has its
organism,
kind of indi-
special
very
anything
Ghiselin, 1981),
sideration.
a
the
deed,
and
com-
as
or
an
dis-
charac-
manifest
from other
its
diagnosis.
fixed, however, through being
the
course
acquisition,
so
of both evolution
that
at
any
particu-
34
A.P.
lar
they
moment
be
can
limit the taxon. That is
found insufficient
why the
identifiable through its
taxon is
attached
name
de-
to
ultimately
as
same
result
be achieved
can
differentand shorter
diversity
the
that
to
Plato's
in
cut
nature
artificial
classification,
daries instead of
this
acters.
quite
For
it is
defined
individuals,
to
in
between,
sophisticated
we
are
is
as
only legal
and
to
way to define
a
the
type
to trace
there
as
such
implication
of
by
two
are
taxa.
more
nomenclature,
ICZN.
They
are
continuum nature of
a
taxon.
2.
The
of
principle
cannot
next
more
must
be
grouped
in
fully inclusive,
that
whole,
the
Equally,
cannot
be
as
two
or
the
thus would
the definition of
As
a
obey
result the system takes
3. The
principle
member of
bear
not
a
only
single
of
a
name,
a
would
single higher
invalidation because of
not
even
more
inin
as
a
higher
overlap and
a
continuum.
fully hierarchic form.
synonymy.
extra
The
must
whole,
a
taxa, for otherwise
latter
taxa).
is, they
taxon-continuum,
member of
a
continua,
in the
way than
(higher
clude the subordinate continua
parts.
Being
other
any
continua
inclusive
be
hierarchy.
names
If
a
taxon
taxon, it
being
synonymy.
can
can
to
follow the principles of no-
be
taxonomy-independent
deviation
a
a
only
destined for
that
sense
discussed here. The
not
is
onomy-dependent principles
a
seem
conflict
it.
with the
The
tific
some
in the
The
least
sedis.
sible
as
parasitic
disturbing
its
specify
to
hierarchy.
can
be
For
assigned
moment,
to
to
any
interpreted
easily
directly
be
orthotaxon
position
at
instance,
a
principle
special
a
local
distinct
taxon was
genus
treated
with
(ICBN
as
of
is
a
one
for the
be
largely
an
and
subor-
understood
which
as
can
only
the
to a
concern
is
A
(the
rather
definition
narrow
There exist
The
as
a
in
sev-
formal
term
1986)
broader
botanical term formformal
orthotaxon,
special
principles
genus
all be attributed
cases
can
1979,
3).
a
can
existing family.
an
parataxa.
Article
of
sedis
least for the
of the
parataxonomy
of the
if it
at
family,
proposed (Rasnitsyn,
general
except
can
practical usage).
kinds
framework of
the
genus incertae
unnamed
serious
called
interpretation
impos-
particular level
Indeed,
order
an
by Melville,
comparison
a
rejection
of synonymy and
published
eral
to
new
more
field
incertae
(normal taxon)
order, but not,
an
synonym of
a
Other and
fungi).
taxon
a
ma-
particular family. This practice
as
a
is
in-
con-
complicated
characteristics make it
of hierarchy and synonymy.
dinated
the
paleontology
a
and
worms
case
an
incomplete
the
be
of groups with
It differs from
in that its
scien-
or
usually
are
into
specific imperfection of the
taxonomy
ontogeny (such
no-
classify
to
regards
reasons
terial involved. This is well known in
and
of
comes
applied
group. As
complete information, the
nected with
group
nevertheless
arises from the
importance of the
scru-
will be
of information
shortage
necessity
necessity
been
principles
particular
a
tax-
important
more
1986), they
the
following
classify
to
cases
fashion here.
cursory
menclature arise when the
necessary
these
with the
they have
(Rasnitzyn,
not
why
problems
and deserve consideration. As
Reasons for
taxonomically
no
That is
so.
principles,
necessarily subjec-
are
there
do
reason to
attributed to
taxa
latter
the
compelling
perfectly the
because
both not
of the
In
cladists.
procedure,
principles
an
no
continuum is
concept,
appreciated
an
defined in
are
meets
taxonomy-dependent
also
leave
rank border lines between it and other
Besides
to
natural
no
as to
done by
demands of the classification
the
of
aware
for
only treated in
fit
not
in
tive
are
of
case
reasons
to
character, for
a
refusing
tinized elsewhere
boun-
does
the
the
186)
of
case
and nomenclature
taxonomy,
menclature
to
with other char-
tightly
so
the taxon-continuum
contrast,
by
unless individuals
manner,
joints",
1983:
(in
Class
overlap
forces that could pack them
space
her
just impose such
we
natural
This
and to fix and
tracing them).
procedure, for
which it is
leave
at
taxonomie boundaries
as
nor
residue).
cited by Hull,
as
taxa
classification is
a
the field under classification,
rank them
a
a
each other,
aim of
("to
metaphor
by
system of
(unclassified
space
natural gaps
trace
a
accommodate all of
to
classified
implies that the main
use
is
should neither overlap
uncovered
any
by
of
means
path of reasoning. Indeed, the
aim of any classification
be
Reasons for
In
nomenclatural type.
The
Phytogeny,
its
to
tag
a
-
Rasnitsyn
taxon
but
can
only
in
system which is parallel
completely
independent
principle of homonymy.
be
the
to
of it,
These
are
Contributions
for
example
66
Zoology,
to
in
taxa
seeds,
or
Table
fos-
classify
to
beetle
or
35
1996
-
created
systems
sil detached leaves,
(1)
II.
of nomenclature.
Principles
fos-
elytra,
Taxonomically independent
silized animal traces, and other works
cases),
larval stages of
groups
with
of them
any
tainly)
should
tant
be
not
the
be found in
can
cer-
2
restricted
3
homonymy
4
priority
5
standardized
6
superior authority
cases
of this
1
claimed
the
explain (e.g.,
Others
66).
taxonomists
refer
the
of the
case
(see Rasnitsyn,
created
to
Dryinini
and
house
ICZN
be.
for
1986,
an
from
wasps
neric and tribal
position
I
Kieffer,
was
tribes
whose ge-
is unknown because of the
female-oriented taxonomy of these groups. Never-
theless, the genus Laberius has been
with
one
explained
which had stimulated him
case
synonymized
of the orthotaxa within the Gonatopodini.
As A.G. Ponomarenko
to
to
it
me,
this
was
detypified
create
parataxa for fossil beetles in Rasnitsyn (1985:
does
81). Detypification
onymy.
perhaps
Yet
a
than
gerous
it prevents any
the
of the
understanding
taxa
will be
a
illness
defence
medicine is
itself.
aims
thus
synonymy,
in which the
case
ungrounded
prevent
hope
I
and
against
a
47syn-
being
more
that
dan-
better
functions of para-
their
arbitrary
syno-
nymization.
The
taxon.
that
rank,
can
cannot
be
above
sort.
in
the
be
a
For
there in
example,
present state
group
placed
with
Article
to
of
certainty
The
parataxon is
to
a
higher
“Cercaria
taxon
known
as
col-
reasons
for not
to be followed
(not
belonging to
followed
be
to
and the
one
following it)
for taxa incertae
for
all
same
sedis)
parataxa
except
particular system)
According
reet.
higher
the above
and thus
taxon
scope
Cercaria
it
ply because
is
trematode type
is
type species
doubt that
zoology,
is
ample
higher
as
has
the
marenko,
as
taxa
been
It does
collective
family
Caraboidea that
family.
Carabilarva is
the
Thus
Carabilarva
type
is
synonym of Carabus
synonymized
as
are
of
boidea, that is Carabus
long
botany.
Another
the
of the
to
formally
yet
attribute
superfamily
junior
the
two
of
Cara1758.
objective
must
they belong
as
to
super-
granulatus Linnaeus,
a
Pono-
species
type
in
ex-
1985)
of the
impossible
this,
and
of the
that this
Carabilarva
beetles
A
sim-
I have little
(in Rasnitsyn,
of Mesozoic
Because
larva
not matter
genus
1985, proposed
such
as
a
type.
eventually typified
done in
larvae
a
to
ulti-
are
same
yet appreciated.
will be
house
to
the
to
similar enough
species.
not
to
be identified
can
the respec-
members
its
mately identifiable by referring
species of
the collec-
definition,
in its
equivalent
is
taxon
tive
to
to
not
be
separate
is
cannot
also
viewpoint
considerations
are
on
the
principles
listed in Table II.
Summary
is used
collective-
that
preceding
but
Müller,
[...],
a
The
of nomenclature
of the
be
genera..." (ICZN
taxon
but this
O.F.
worms
larvae
collective
detypified,
a
a
generic
special system
knowledge
in
of
usually
genus of
a
for Trematode
67m).
be
kind of
parataxon,
assigned
organized
name
there
poorest
It is
established for
1773,
on
systems.
last and
lective
those
tive
ar-
example
the
Gonatopodini (Dryinidae),
names
International Commission
of the
good
no
synonymy
to
some
which
details),
rank-indicating
of de-
drive of
As
application
Article
means
Laberius
genus
male
a
parataxa, however
might
synonymy
as
persistent
synonymize
to
such
to
1914
arise
to
difficult
are
of ichnotaxa,
case
known
are
parataxa from the
fending
bitrary
well. Some of them
as
of
and
hierarchy (not
3
type concept
area
type (with
involve the
to
restrictions
Taxonomically dependent
Meyen (1990).
sort are
the
beyond
Nomenclature
2
Some
freedom
Impor-
and functions of
nature
taxonomie
imposed by taxonomically dependent principles
yet they
with the latter.
synonymized
on
even
unrestricted
in
only
and
orthotaxon,
some
in-
dimorphic
female-based clas-
possibly (sometimes
are
considerations
parataxa
highly
1
or
worms,
from orthotaxa
differ
synonym of
a
in
traditionally
a
sification. Parataxa
that
living parasitic
unassociated males
even
sect
or
caddis
(e.g.,
stated
is incor-
Phylogenetic
relies
(2)
on
search for
about the
attempts
inference,
like
(1) observations
scientific
analogies, (3) creating
falsify
work,
(including experiments),
underlying patterns
to
any
these
and
of
hypotheses
mechanisms, (4)
hypotheses
(mostly
A.P.
36
through
their
and (5)
implications),
sults of the above attempts using
assessing
of
a set
re-
presump-
tions.
The
to
set
of
of
array
be
The
phylogenetic presumptions
of
analysis applies
group
and relies
relationships
into those relevant
to
groups.
the
analysis
subgroup
the
covers
character
array
which
of
of
delimit taxa,
and
This
the
biogenetic
help
us
to
discriminate
to
comprises
important
most
for
presumption
the
presumption,
a
of
pre-
the basis
of
vestiges,
presumptions
concern
the
The
of
group
possible
nomic
users,
i.e.,
myopic (purely
Of
cladistics
is
three
empirical).
and
Table
fastidious,
with
the
balance
main
rival
Another
taxo-
cept,
the
lizes
Phylistics
the
only
continuum.
synonymy,
may
sedis,
organized
in
or
a
pendent
tice
any
then
incertae
order
There
are
con-
type
taxonomically indede-
conventionally
are
make the taxonomic
to
par-
developmental
violate the
to
a
detached
particular
organisms).
of the
which
principles,
signed solely in
of
found
reasons
of
particular
a
group
violate
to
nor
of
or
living
a
neither valid
cept,
objects (e.g.,
organs,
of
which
special (independent) system of
ticular kind of
a
of
cases
taxon
a
as
taxonomi-
hierarchy and
material,
in
con-
less stabi-
or
followed in
either
are
designed
those of
not
deficient
taxonomically
be
be
3
dependent
other
Two
cally dependent principles,
should
more
of the taxon, that is
name
and
is the type
one
that
one
purely
is shown in
as
taxonomically
important
most
monophyletic
prac-
stable and uniform.
more
Acknowledgements
As
states).
present
written in
and
be
employs
result
is intermediate in
system
of
content.
a
that reflects
similarities
and
a
comprehen-
dissimilarities,
to
be
method
defines
a
to
taxon
as
approach
a
abroad.
I
but
I
expanded
an
was
have
circulated
benefitted
version
among
from the
of
of
R.
Schram
University
for
Sciences,
of
Dr.
San
for
(Institute
and
to
Michael
of a
text
colleagues
of
input
Prof.
in
many
and
Systematics
for their
James
Harvard
T. Ghiselin
Frederick
Population Biology,
extremely helpful
the
help concerning
M.
Uni-
(California
and Prof.
Francisco, CA),
Amsterdam)
advice
debt
Comparative Zoology,
versity, Cambridge, MA),
Academy
in
particularly
am
Carpenter (Museum
and
for
assistance
Criticism
and
in
importance
most
based
statements.
advice
with
linguistic
suggestions
of Prof.
Lawrence, KA),
an
which
discus-
pertinent literature,
is
The
Its ul-
sense.
and
colleagues,
to
the cladistic system
hypothetical
represents
paper
1989-90,
upon
extreme
a
The
I
making
am
Charles
Dr.
of two
also
D.
K.Yu.
and
other
stylistic
anonymous
visible
thankful
for
Michener
Eskov,
mistakes
many
the
were
and
of
weakly
discussions
(University
Dr. A.G.
problems.
reviewers
and
of Kansas,
Ponomarenko,
Dr.
imporS.A.
heuristic
the
Among
built
independent
are
employed,
concept
which is the
Russia
about the evolution-
while the relatedness is considered
tant
II.
principles,
sions,
goal is
they
are
system
a
between-
approach, which
character
phylistic approach
timate
create
such and thus appears
as
cladistic
of
overburdened
that
be
to
taxa should reveal maxi-
only apomorphies (inferences
history
presump-
pheneticism relies solely
approaches,
the
to
homogeneity
heterogeneity.
represents
of all,
for the maximum diver-
meaningful
available characters
sive
indepen-
first
principles,
sense
claimed
6 of which
similarity.
objective of taxonomy is
taxa are
The
analysis
vs.
These are,
similarity.
within-group
mum
ary
similarity
presumption of parsimony, and the
whose
too
in
the problem of the inherited
tion of
weighted
sity
presumption.
outgroup
engaged
dently acquired
the
the
of 9
taxonomic
a
stage
and
a
cor-
the system deserves re-investi-
or
conventional in the
evolution, of functional efficiency,
complexity,
as
whether the resulting taxon is
Taxonomic nomenclature is
on
fossil
of
used to
are
considered
is
monophyly
gation.
sumption of analogy, those of the irreversibility of
of
discontinuities (gaps)
and
rectly delimited,
of
apomorphic (de-
group
the
paleontological
the
are
characters,
states.
and
The former
that
and nomenclature
taxonomy,
taxonomically dependent principles,
separable
differences,
presumptions
presumptions,
I).
presumptions
further
similarity.
plesiomorphic (ancestral)
wide
The
are
study
of
(Table
the paleonto-
on
transformation series, i.e.,
polarize
rived)
where
tinuum,
possible
ancestor-descendant
to
analysis
those used in the
the
phylogeny,
analysis
primarily
for
logical presumption
used in the character
and
groups, involved either in
two
in character
or
the
covers
particular presumptions
into
analysis
of
"knowability"
more
segregated
group
Phylogeny,
means to assess
presumption
an
-
Rasnitsyn
this
monophyletic
goal.
con-
Rautian,
Dr. M.A.
tological Institute,
and
Dr.
I.Ya.
Shishkin,
Russian
Pavlinov
Dr.
Academy
(Moscow
V.V.
of
State
Zherikhin
Sciences,
(Paleon-
Moscow),
University, Moscow).
Contributions
Very
important
Florence
my
Dr. R.
of
insects,
insect
inclusions
Invertebrate
I
in
the
of Mr.
of
Andrew J.
Natural
the
37
Schram,
of the
style
Ross,
eludes
edit
1983).
Contribu-
in the
Department
is
account
and
other
of
need
London.
in
published
as
5.
for the term
Farris
has
parsi-
hypothesis
said
on
an-
there is
'phenetics',
would
'phenetics'; 'everything'
do
no
just
488).
lobe
galea (a
of the
of the
It is
post-Triassic origin.
and is not known
in
living xyelids
is
mouthparts)
for
proved only
the subfamilies
comprising
himself
is
(Farris,
of the most
of the best
synonym
everything
apomorphic similarity
divided
(Rasnitsyn, 1992a, b).]
1979:
well";
An
"If
a
(as
unnecessary
occasion,
evidence
phylogenetic
makes the notion
approach
hypothesis just
thus
of
of the
weighting
This
monious
Curator of
History Museum,
present
Drs.
to
the collection
study
to
amber, kept
Burmese
version
Prof.
by
and
opportunity
Paleontology,
[An abridged
Russian
had
1996
Sluys (Amsterdam)
English
Zoology. By courtesy
fossil
-
(I)
effort
great
and
Pieters,
to fit it the rules
text
tions to
the
was
F.J.M.
66
Zoology,
to
and
Xyelinae
for the Triassic
the subof
possibly
the
subclade
Macroxyelinae
Archexyelinae.
Notes
6.
To
be
1.
This
try
only
and
either
extinct
cladists
ignored by
collection
as a
identify
fruitful
2.
is
problem
cestor
real
a
ancestor
should
among
that
to the
possibility
(pp. 7-9).
"Scias
Characterem
Characterem"
In
I
follow
be
easily
that
169],
is
sumption,
the
subject
mistakes,
consider
dangerous
would
of
reduce
approaches
a
be able
reduced
ideal of
a
the
lowers
to
of
dares to
and
support
use
and
pitiful
the traditional
There
necessary
exists
demands
for the
scientific
a
a
why
I
am
which
abilities
trying
promise
I
of
to
maxi-
develop
and
interesting
mistakes.
to
amount
more
wide
means
to
of
following
economy,
reach
understanding
the
of
aim.
which
in-
of
this
context.
At
on
lower
their
9.
studies
have
been
literature, by Nevesskaya
both
equivalent
monophyly
follow the cladistic
be
many
classified
(one
vs.
vs.
of
Equally they
absence
et
fre-
pat-
had
in
their
both in
than
one)
ancestors)
are
impor-
are
rather
published
and
of these
in
of the
in
ex-
Russian.
with rel-
(1986).
the
a
term
latter
usage.
I
depending
discernable
cross
in
classifica-
lines
their
refuse
group,
on
of
lower
according
to
that is
cov-
being
following reasons.
classifiable
of descendants
research
fossil
the results
cladistic
strong
special meaning
holophyly,
for the
a
molluscs
economic
levels
al.
compli-
and
of intensive
a
Para-
rapid,
hydrology
system,
phylogenetically
more
try (successions
line.
usage
common
A
with
(1971) using monophyly as
and
taxo-
the
recently summarized, together
paraphyly
to
as
currently they
characters,
evant
ering
in
have
have
Ashlock
in
of southwestern
seas,
the
and
mostly
the
inhabiting
subject
They
I follow
been
into
(Neogene).
taxonomie
shell
to
not
biota. The bivalve
bivalves
Besides,
pro-
me
because
usage
things
because of the
strata,
if
classifica-
classes,
seas
of the
outside
a
the
below).
changes
have been
even
are,
how
the term
in current
Tertiary
of the
century
a
oil-bearing
known.
and
respec-
permitted
into
molluscs
history
evolution
than
well
can
of parsimony
the notion
of
Paratethys
tensive
prefer
at his
both
of the
more
the
they
restriction,
living populations. Unfortunately,
of meththe
later
has
clear-cut
text,
often reversible
on
tion relies
This
and
and
effect
tance
us-
the
as
restrict
epicontinental
environmental
the
it
all
here.
this
bivalve
of
of straits inside
tern
people
on
during
to
it
ordering
not such
main
for
into consideration
here
(individuals,
are
However,
work,
and without
how
presented
follow
systems
system
a
quent,
abilities,
to turn
to work
cated
we
albeit
and
to
experience
some
extensive
popular
and character states involved.
presented
proposes
not
of the
appropriate to explain
are
suggests (see
Eurasia
least
do
and
tethys,
to
experience
(and
of
I
researches
E.g.
for
More
variety
8.
strategy
is not mine.
broad
(1974)
theory)
belief
scientific
unapplied.
rest
source
interpretation
another,
our
work,
mistakes
correct
Ockham's razor and related
least
This
results).
permits everybody
That is
case,
data
have
taking
available
applicable only
as
nomie
differ-
(those
ideas
the conclusions
classes
mis-
either
without
of all taxa
that the
Griffiths
tion
faultless
first
In the
science
7.
It
experi-
two
of
use
reach
pre-
then
and
I
one
and could do it.
and rather
posed approach works,
the
thing.
prefer
methods
would
good, as
the approaches
findings
new
and
could
methods, while high
is free to
make
efficiency.
mum
the
in
any
no
are
of scientists.
standardized
mistakes to their
is
there
data
new
methods.
special,
slightly outdated,
said
using
diversity
the
participate
diversity
that
a
senseless
systems.
even
there
safe
incorrect
and
be
I believe
Genus
considerable
them.
most
skillfulness
which
discoveries),
I
giving
mass,
science
ods,
4.
the
dramatically
and
we
particularly gifted persons
very
and
be
here
cladograms presented
prepared using
(Rasnitsyn, 1995)
tive réévaluation
length
character
like
groups,
topics,
here:
the
ir-
the character"
straightforward
However,
to correct
or
only
and thus
knowledge,
would
would
Ghiselin).
examples
Of course,
at all.
cognition
avoid
the
and related
unavoidable.
are
would
few
and
easy
strategies possible
ent
M.T.
thorough knowledge of,
in,
method of
ing
an
genus
is
who
sed
Genus,
"Know,
for
presumption
not
is
but the
genus,
from these
seen
paleontological
takes
is
analysis
group
Linnaeus
constituere
non
(translation mine, corrected by
ence
approach
is discussed at
problem
short,
[1751:
discriminate
to
The
does not constitute the
needs
this
their characters
using
groups
analysis.
below
can
it should
organisms,
the
trace
we
and character
It
known
in this matter
be
cladistic
computerized
an-
an
(see below).
The fact
relevant
3.
consider
I do not believe
extant.
or
who
of plesiomorphies and refuse to
the
persuasive,
more
possibly
to
Taxa
how
ances-
border
presence
whether
A. P.
38
its
tral
line
is
descendant
is
called
for taxa with
descendant
Cladists
This
leaves
usage
for the
i.e.,
combination
not the
for
case
there
exists
taxa
That is
is
and taxon
is
valid
with
and
thus
single
a
term,
for
and
line.
a
This
is
considers
needs
term
a
The
narrow
holophyly,
in the
The cladistic
which
the
line
combined,
nonsense.
badly
over
any
nontaxon
ancestral
priority
other
meaning,
a
a
or
paraphyletic.
taxon.
broad
so
irre-
exclusively
two notions
taxon
follow Ashlock
I
why
of
replacement
a
termed
are
also
ancestral
single
a
phylistics (or phenetics)
as
of
an-
descendants
and without
monophyletic
of monophyly has
usage
line
line(s)
term
ances-
single
a
latter term
with
paraphyletic
taxa
all
the
taxa
of nontaxon
paraphyletic
covering
no
the
cladistically
it has
(whether
term for the
no
traditional
needs
approach
single
A taxon with
use
descendant
and existing
of de-
line(s)
a
polyphyletic irrespective
ancestral
while
ones,
by
than
more
traditionally monophyletic
single
a
crossed
line(s).
of its further fate
spective
only members).
is
with
called
universally
line
line
A taxon
not.
or
possessing
cestral
border
(side)
upper
scendants
broad
one, and
for the
latter.
of these terms.
usage
-
Rasnitsyn
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