Cladistics
3(4):305-332
Component-compatibility in historical
biogeography
1 AND M.C.
Zandee
M.
1 Institute for
2
Abstract
-The
problems
tional data for
be solved
groups
by applying
of components.
tial
Institute
multiplication.
the
verted into
reflect
to
tion
are
hoc
(ad
of the
the
additive
or
2
such
as
the
of endemism
for the
The
same
matrix
and
0 here
and
of
terms
in
over
of
can
and
largest
are
selected
and
states
sets
of
of
con-
amended
areas.
Area-
mutually
com-
and contradic-
best fit the
the best
other standard
be
can
missing
is illustrated
procedure
and several
as
the
boolean
taxa over areas
(vicariance)
support
par-
be used in historical
newly defined),
for the
by
of character
terms
can
for
areas
different matrices
Area-cladograms that
extinction).
Xiphophorus,
two
groups
(compatibility)
transcription into
distribution
wide-spread taxa
of endemism. The
areas
Heterandria
of
exclusion
of
means
derived data matrix
and contradiction
support
or
of distributions
by searching
evaluated in
are
by
taxa,
from distribu-
the members of those
the
represents
problems
dispersal
between
areas
It is derived from
binary coding.
accomodate the
among
genera
data matrix
binary representation
from the derived
the balance
fish
the
a
derived
cladogram
Area-cladograms
of relationships
poeciliid
from
thus
given phyletic data (Assumption
to
interpretations
regarding
tructions
gives
determined
components.
matrix
matrix
by
relationships for
relationships among
cladograms.
describes
form
1
be extracted
matrix
or more
to represent the
Assumptions
cladograms
patible
second
binary
biogeography
one
The first
can
The Netherlands.
of parsimony and mutual inclusion
principles
data
The
in
monophyletic groups
The Netherlands.
University of Leiden,
historical
reconstructing
of taxa and the cladistic
the two
sets.
Biology,
for Systematic Botany, University of Utrecht,
Components
monothetic
endemism;
of
Theoretical
2
Roos
possible
the
by
data
recon-
example
examples.
Introduction
Historical
biogeography
and
relationships
various
groups
which
terns
of
can
For this purpose,
each
terms
the
sent
area-cladograms
be
can
two
reconstructing
purpose,
1986:
1).
in
the delimitationof
The main
the
and
for
this
‘vicariance
framework for
a
one
case
map’
the main
method
(Wiley,
in
belongs
to
relationships
the
general
of biota
of endemism is
more
the
area
of
between
and Parenti
areas
(1978,
a
area
(described
coping
which
this category,
as
The
first
repre-
of wide-spread
reason-
aims
type
starting-point (Humphries
at
For this
and Parenti
and the
species
under consideration. The methods of Rosen
here.
The second type
scenarios of
Here, emphasis
with
in
of endemism.
(‘cladistic biogeography’).
1981) belong
between sister
1986).
endemism). Congruen-
areas
reseach.
manipulation
pat-
of biota
area-cladograms by replac-
general area-cladograms,
relationships
postulating speciation
is
elucidate
the historical
biogeographic
biogeography’).
problem
species-
with different purposes and different lines of
concern
or
in
depicted
constructing evolutionary
geographic separation
region might
transformed into
development
Nelson and Platnick
search for data for
study (e.g.
as
then
cladistic
ares
problems
on
Humphries 1982, Humphries
are
historical
the historical
absence of groups from
(1976)
are
types of analysis
recognized
certain
biogeo graphical entity,
the
between
correspondence
a
between the cladistic relations within
the terminals with its distributional
at
species represented
At present
ing
species cladograms
general patterns,
a
develop hypotheses
to
of the smallest relevant
cies between
there is
in
organisms occurring
be used
Nelson and Platnick 1981,
(Rosen 1978,
ing
that
assumes
area-relationships. Comparisons
is
particular
put
on
concerns
groups under
vicariance patterns
models. An essential element is the search
taxa
(Wiley
overlapping
also his
and
Mayden
ranges.
parsimony
1985;
Wiley’s
598-600).
‘ancestral
method outlined
In
species
recently
press).
To date, the
general
treatments
of cladistic
biogeography (Nelson
and Platnick
1981,
CLADISTICS
306
1982, Humphries and Parenti 1986) have been concerned with obtaining
Humphries
unambiguous
in which
ample,
of
further elucidation
more
into 3
aims
based
provide
to
the
on
2.30
by
Geesink
(1984),
general area-cladograms
ference between
be
into
integrated
in
emplified
the
a
two
one,
authors neither integrate
the
steps
nor
General
1.
DATA
analysis
Nelson
and
(1987).
The
both steps
will be
procedure
Rosen
(see
taxa.
ordered (additive
uniquely
respect
to
the
of
one
2.
MONOTHETIC
or
1981, Humphries and Parenti 1986).
given by Humphries (1982)
make the
because,
procedure
and
Hum-
by
these
knowledge,
to our
for the latter
step explicit.
Area-Cladograms
or
four
as
steps:
unique,
but
state
of
a
separately
or
some
blocks for
of
series of
a
of
be
may
step
1981).
be
cladogenetic
by
means
so,
a
the
combinations)
consist
may
a
an
words,
integrated
up
This
but build
1978).
a
implies
tree
series of
that
from
we
sets
group
for
struc-
a
none
set
as
as
try
of
a
set.
of which need
(Zandee,
1984).
sets
of
(sensu
taxa
character
as
states
this is tenable in the
(a non-homologous
first level
not to
cladon is
character type
homologous
long
as
a
monothetic
of monothetic
use
rejected
phyletic
When
partial
a
strict monothetic
whole
or
a
block
of transcription into monothetic
into different characters
convergence is
taxa.
a
building
a
name,
cladon represent
same
is the idea that
as
rank,
then is it called
then it is called
defining
kept
tree,
only
and
1975; Farris,
The rationale behind the
among
unit
combination of character states,
rather than broken
In other
to
states
unique
a
cladograms
similaritiesobserved
singular
additive
states
(neutral),
uniquely represented by
of terminal taxa, without
character
terms
must
parsimony,
states).
unordered
and the data matrix
in combination,
as
group
set
a
distributions
character)
context
of
i.e.
unique
Nelson and Platnick
building
(all
as
mixture.
define this
When it is defined in
Charater
ordered
exhaustively
1959; Sharrock and Felsenstein,
terms
be coded
may
transformations,
Clada are extracted from the data matrix
(Beckner,
defined in
(i.e.
or
of their
states
SETS
cladograms (cladon:
be
binary),
a
Character states, either
taxa
These character
sequence
either type
of terminal
sets
ex-
1978,
The data matrix shows the distribution pattern of the various intrinsic character
ture).
difcan
MATRIX
amongst the terminal
with
prin-
analysis,
area-cladograms
Therefore,
Poeciliid fishes
area-
Both
parsimony.
method for cladistic
a
The
1986).
selecting general
and
of kind.
conclusions
be summarized
can
not
1981, Mickevich
conflicting
two
and
The intricacies of this
identical examples
8.28 in
methodological principles.
same
degree,
to
and Parenti,
area-cladograms
reconstruction of the
and Parenti (1986) lead to
The cladistic
the
by
of
ex-
still needs
and Zandee and Geesink
1986),
to
8.25-27
Figs.
Humphries
develop
to
(1985,
general procedure.
some
from
in
cladograms
one
1978, Wiley
We will show also that
phries
steps is
for
(1981),
general area-cladograms,
constructing
(1985)
Roos
ruled
are
biogeographical
Platnick and Nelson
into
group-compatibility
Zandee
We shall argue that the steps from
to
of
examples
(area-cladograms).
statements
area
2.31
to
method for
a
principles
have been used earlier
recently applied by
4
or
area-cladograms
Figs.
of
terms
in which the groups under
or
3-species-statements (cladograms), including
unexplained jumps
the
from
in
problem
under consideration. Nelson and Platnick
several
or
the
of endemism,
areas
the transformation of
(e.g.
1981,
paper
cladograms
ciples
or
one area
on
converting
and Platnick,
This
two
wide-spread species,
one
step,
in
occur
concentrate
least
at
next
All authors discuss
area-cladograms.
species
is absent from
study
3
[VOL.
fit
integrated
a
explanation
for
minimal number
series
of steps.
HISTORICAL
307
307
CLIQUES
3.
A search
of
BIOGEOGRAPHY
Cladons
is made for the
is based
cliques
the
on
of
These inclusion and exclusion relations
the
when all clada in
case
representing
stands for
a
relations,
the
used here in its
a set
in the
sets
original unambiguous
maximal
can
be
clique
they
include
the
graph theory,
are
used
all
are
is
meaning
complete subgraph (Garey
other elements from
character
represented
mutually compatible,
are
the
the list of clada. The
group-compatibility (not
with other cladons when
compatible
are
present in
largest clique
concept
the
is
each other.
graph (network).
a
a
In the
clique.
connected.
mutually
Johnson 1979).
In
graph
Clique
concept from graph theory,
a
and
in
set
exclude
or
by
recognition
compatibility).
as
and
This concept, and
of the
implementation
method in
computer algorithms.
4.
CLADOGRAMS
The
largest cliques
and
evaluating
and
choosing
The first
their
Support
the number of
means
which
tion
(homoplasy)
of the
be
can
to
The latter
character
are
Eventually,
the
belong
to
in the
one
to
several
or
Humphries (1982: 445)
encompasses
species
with
areas
the cladistic
to
analyses
can
Assumption
nick
2
(see
but
(1981),
Assumption
0
also
Wiley,
did
they
true
delimitedand
actually
the
au
,
This
that in
be derived
Under
true
comparable
not
press).
give
of
other
a
(Fig.
Assumption
each
on
to
supplementarily
dadogram
statements
a
as
to
(Zandee,
with
states
the
determined
1984).
multiple
consequence these
states
here
can
it
However,
i.e.
two
to
as
0
or
implies
more
species,
identity
a
be noted that
proposed by
a
phylogeny
species
1
a
analogy,
way similar
biogeographic
the
might
0.
Assumption
is the best
possible
be
wrongly
to
resulting species
group.
undisputable
are
assum-
Consequently,
components.
be misconstrued but
species
and
Nelson and Plat-
appears
monophyletic
represent
of species
of
possibilities
that if
this
Following
method
a
exchanging
Assumption 0, Assumption 1,
were
the
of
pursuit
involves
be summarized in
must
assumptions,
full credit
is the
analogy
with sister groups.
The latter
two
the basis of these criteria
biogeography
cladistics. The
and that from the distributionof
(Nelson
taxon
of its range
monophyletic
Contradic-
and/or reversals
Construction
wide-spread species
Id the
distribution pat-
not
its
1 the component, A+B
1c).
1
wide-spread
parts
As
selected
are
homologies
under three
in
a
con-
support).
phylogeny.
a
area-Cladogram
represents
Fig.
phylogenetic relations,
can
of
phylogeny. Assumption
inhabited by
means
minus
transformation series.
be sister groups and should together represent
t
support
in
among those
with the idea that the reconstructed
begins
estimate of the
ed
out
for each
synapomorphies
described above.
analysis
used
criterion,
of all its 3-cladon
biogeography proposed
be carried
i.e. those
in the first instance.
of endemism and
the methodof historical
second
stated that cladistic
code
a
data matrix
used in association.
independent multiple origins
cladogram(s)
General
which
are
by interpreting
the
single origin (synapomorphies).
a
context
different
likely rcpresentation(s)
most
in
states
criteria
two
purpose
synapomorphies
elucidate
homoplasious
were
considered
The
states.
comparison
analysis might
which
origins
This is followed
character
character states,
the number of
the total number of
local outgroup
by
fitting
explained by assuming
means
remaining
first, pertains
of the
minimization of the value of contradiction minus
(or, equivalently,
tern
cladograms.
terms
criterion is the maximization of the value of
(quantitative)
tradiction
in
few of them. For this
a
one or
transcribed into
are
implications
group).
in
and
one
(compared
Platnick
1981:
421),
part of its range
to
other
areas
it is assumed that whatever is
must
also be
inhabited by
true
of the
the other
taxon
species
in
of the
308
CLADISTICS
In
1
species
to
that
implies
more
is
la the
Fig.
related
closely
regarded
The
224).
(occurring
A is
area
related
as
to
(Fig.
or
A+B is in
consensus
tree,
comprises
2
Assumption
of its
related
area
more
to
than
inhabit
they
areas
a
mono-
or
but the component
C than
area
D. This
means
one
and
the
is
more
species
newly
closely
D). Assumption
area
D and that
to
that if
species,
B)
in
(occurring
1
delimited
the
paraphyletic
1981:
and the
group,
delimited
wrongly
A+B+C is considered
defining
is
species
cladogram sequentially (Platnick
(Fig. Ic-e). Thus,
B
area
in the future
component
indisputable
the relations
(Nelson
taxon
in
and
species 1, 2,
(c-e) fully
derived under
components
one
Platnick
1981:
part of its range
of this
implications
that
implies
area
B is
true
for both
can
the
between
under Assumption 1.
and
Assumption
432)
might
and their distribution
3,
resolved
be
(A-D).
the
con-
0.
indicates
not
areas
area-cladograms implied by
that whatever
of the
true
taxon
is
of
true
a
in other parts
range.
The
more
take
(Fig. 2b).
closely
areas
one
A is
area
assumption
more
related
closely
to area
A and B. If the
of all
to
of
C than
species
1
to
compared
to area
to
C
area
relations of A
branch off in the
components A+B and A+B+C
relationships
illustrated in
are
related
possible positions (white
If the relations of B
possible positions
the
to
A
areas
3
species
branch off from the
comprise
doubt,
area-cladogram
(c)
widespread
B
C)
in
(occurring
than to
3
lb).
sensus
2
C than
holds for the
same
Fig. 1(a). Cladogram showing
(b).
area
closely
more
area
species,
species might actually
defining
in
actually representing
should be sister
1
wide-spread species
2
[VOL.
D
compared
In
D
to
C
Figs. 2b, c)
and D
are
be misconstrued.
and D
on
true, A
means
Fig.
2a, Assumption
(Fig. 2b)
but this is
area-cladogram (Fig. 2c).
may be in doubt. This
might
to
(Fig. 2c),
circles in
C
2.
Fig.
than
the
can
not
are
true, than
area-cladogram
take
one
of all
In other words,
that both the
Furthermore,
and/or that
necessarily
this
identity
wrongly
both
and
delimited
HISTORICAL BIOGEOGRAPHY
309
species might actually comprise
for
sequences
the
(b,c)
O
indicates
DATA
the relations
positions
which
the
missing
it
paper
is
between
procedures
each of the three
matrix
under
x
(clada
manner.
for taxa,
areas
the
product)
and
(B
A,
1,
and
2,
con-
and their distribution
3,
of Assumption 2 derived from the
respectively)
on
areas
cladogram.
take.
can
which of the three
for each
study
taxa
terminal
derived from
we
taxa
have
and
of taxa,
group
nodes; Farris
x
each
distributional
data matrix
for each group of
et
x
cladograms
a
larger
can
A data matrix
be obtained via
areas
al.
a
taxa.
area-cladogram
a
are
a
Platnick,
Assumption
0.
of
to
form
a
stepwise
a
cladogram).
1981).
When the
From
same
can
have
to
inner-
(boolean
gives
monophyletic
the
be
a
of
areas
group
distribu-
same
this data matrix
set
a
additive
Each column in the
or
belong
we
by
matrix
cladogram
area-
obtains for
joined (column wise)
cladograms)
loop.
During
For each
for several
separate group
this evaluation
The distributional types
one
from which
selected from the data matrix based
compound
groups
of taxa
general
of
on
to
reduced matrix suitable for
all
can
also
is selected for
of the nodes of this
Assumption
0 in order
taxa can
analysis
taxa
area-cladograms
area-cladogram
corresponding
reduced data matrix. Reduced matrices for each group of
together
matrix
combination
terminal taxon
clada from several
cladograms
data in
raw
binary
a
separate data matrices
x
of endemism
areas
areas). Second,
x
The
one
of
matrix with distributional data
each
general area-cladogram
kind of feed-back
extracted and evaluated.
group of
sets
(taxa
1970).
with
terms
derived.
derive
to
a
be
(areas
two
Identical distributions
resulting
data matrix
area-cladograms
but
under
analyses
in the
groups present
binary
a
clada from
Nelson and
be derived under
several groups of taxa, the
form
taxa.
analyzed
in
converted into
matrix
(areas
(character type:
can
made.
are
assumptions,
for
appropriate
patterns
final data matrix represents the distribution over
tional type
assumptions
full evaluation of these
a
the distribution
This data matrix is
comprising only
of
depends
corresponding monophyletic
for each
of
group
First,
biogeographical
tain
with all the
0
comprises
and all
taxa
binary coding (taxa
are
species
derive the data matrices
to
Assumption
areas).
cladogram
each
group
assumptions.
The data matrix
of the terminal
to
polyphyletic
application
concerned with
primarily
not
the
provides
Data
or
MATRIX
The derivation of a data matrix
This
para-,
for the
area-cladograms required
the two
the
mono-,
resulting area-cladograms.
Fig. 2(a). Cladogram showing
A—D.
a
309
to
to
ob-
be then joined
derive
a
general
area-cladogram.
The
derivation of
a
data matrix under
Assumption
0 is
equivalent
to
the
procedure
310
CLADISTICS
described
Brooks
by
distributions
the
matrix under
We
from
start
a
Assumption 0, but with
subsets of
with the
tions
possible
cupied by
in
areas
together
wide-spread
a
the
the
on
0. When
basis of joined
for
thus
wide-spread
Assumption
The derivation of
a
is made in the
two
extra
in the
of each
a
now
step,
we
as
their
This
same
take
we
From
all
means
To
compile
Assumption
the
a
areas
more
on
analyses
joined
as
oc-
sister
taxa
then pro-
can
Assumption
feed-back
a
loop,
the
for the separate groups
columns.
as
under
analysis
an
of
sets
In the actual
2
(see
taxon
should be
raw
data
2 is
Assumption
even
combination between the
a
0 but
Assumption
followed
now
a
by
with
matrix
elaborate
missing
subsets
groups
and all
areas
We take
amended for
as
these steps allow for many
Figs. 2, 4),
i.e.
all
any
but
462
et
seq.),
it does
must
implementation,
the basis of the data matrix
to
applies
not
(1981)
a
results
areas oc-
area-
exemplified
zero
weight.
assumption
must
whole. The matrix should
incongruencies
pre-
1.
2 the data matrix is
Assumption 0,
and
possible
this
because of
Assumption
in the second
as
as a
possible
of the
follow that distributions
implied by
analysis
to
Assumption
under
analysed
areas
be considered
prior
which also
of
over
all of the
wide-spread
other branch in the
weighted differentially including
taxa
one
of
for each of
consequently join
we
original
possible
duplicate
a
duplicate
separate
a
defined by Nelson and Platnick
p.
of their
all of the columns of
branch off from
might
as
implementation
a
possible
monophyletic
those from the subset distributions
as
data matrix
analysis,
above and
into several parts
principle
in this
with all
corresponding
with the
areas,
that all distributions of
down
0.
complete
missing
Assumption 2,
areas
within it,
compared
For
two
the separate
thought experiments (ibid.
over
Although
as
are
combine each of these with
wide-spread
a
be broken
sent
two
areas
occur
described for
as
with the distributions of all other
be included in the data matrix which
not
or
is obtained via
described above for
taxon
amended for
in the first step.
taxa
sequentially
cause
the
the data matrix.
to
implied by Assumption
cladogram.
by
Assumption
I i.e.
way
made in the
already
These combinations as well
added
are
operations.
duplicates,
of
will
proviso
2
the data matrix derived under
cupied by
same
general analysis
were
combine the distributions
we
combinations and
as
possible
combined
are
with the subset distribu-
this
for several groups of
in the
distributional types
way
wide-spread
In the second
taxa
com-
described for
as
areas)
more
or
of
principle
area-cladograms
from the
same
cladogram(s).
themselves
these
more
with all
correspond
analysis,
either branch off
might
is
steps.
In the first step,
areas
the
data matrix suitable for
complex. Starting
two
of
2
(occupying
In the actual
data matrices
taxa
selected
only
Data matrix under
more
parasite
area-dadograms.
the data matrix for
provisions
of taxa,
taxon
represent
taxon
The derivation of gener al
ceed
and
1
is made
two
the distributions that
wide-spread
Assumption
under
A combination between the
scored in the data matrix.
results which
from
suitable
as
data, i.e. the distributions for terminal
raw
of its sister
combinations
together
group. These
areas
are
just
hosts
over
parasite
relationships
areas.
analysis
an
of
sets
proviso;
one
of each
areas
fit for
two
same
and their cladogram(s).
taxa
distributions
parasite
hosts with
data matrix
the
data matrix from
a
infer host
1
Assumption
The derivation of
plex.
from
cladograms
to
that the method described here is
suggest
we
to extract
in order
parasite cladograms
by simply substituting
cladograms
Data
fact
host
general
which he used
(1981: 232-234)
hosts and
data. In
parasite
derive
to
over
3
[VOL.
expanding rapidly,
it is conservative nevertheless.
extra
derived in the first
step
we
extra
can
step,
amend for
i.e. after
missing
amending
HISTORICAL
BIOGEOGRAPHY
311
311
for
terminal
wide-spread
Parent!
examples they
Neither Nelson and Platnick
taxa.
indicated which
(1986)
deal either with
should be
implementation
missing
areas or
with
(1981)
nor
but
taxa
wide-spread
and
Humphries
because in
preferred,
not
their
with both
simultaneously.
The data matrix for
for the separate
general analysis
a
groups of
the evaluation and selection of
In that
appendix).
made in the
types
case
the
be obtained
for
taxa
wide-spread
feed-back
a
and
of each separate group of taxa, thus
analysis
the data matrices
by joining
i.e. after
loop,
for each separate group of
area-cladogram
provisions
It should be noted that the
with respect
to
missing
areas
taxa
were
(see
already
selected distributional
only
be
connected in
in
occurring
these
used in the data matrix is
coding
areas.
the clada among themselves
to
an
can
also be obtained via
can
joined together.
are
(or neutral)
are
It
taxa.
areas
is because
connected
are
in
the
necessarily
is
coding
constitute internested
already
sequence, it
a
kind of additive
Any
of terminal
sets
terminal
taxa
or
hierarchical sequence
a
unordered
because
superfluous
If areas
taxa.
groups of
taxa
the
provided by
cladograms.
Components
In this step the data matrix is transcribed into
This
areas.
is
transcription
equivalent
A component(analogous
nick,
1981).
tion.
Components
area-cladograms
serve
to
a
list of
monothetic
(partial)
the derivation of components
is characterized
cladon)
a
blocks
building
as
do clada for
(as
to
for
by
area-cladograms
a
as
particular
well
as
sets
of
and Plat-
(Nelson
distribu-
for
general
cladograms).
Cliques
Inclusion and exclusion relations
ponents and maximal
cliques
which
comprise components
AREA-CLADOGRAMS
The maximal
If only
one
cliques
group
of
several
cladograms
These
absence,
in
Synapomorphy,
states
monophyletic
do
not
share
then be
and
of
areas
groups
monophyletic
is
a
means
or
are
of the
groups of
ad-hoc
history.
or
not
fit the
not
of
com-
Cliques
other.
same
taxa
general area-cladograms.
or
one or more
is used in
taxa
(based
matrix
area-cladograms
the
involved,
area-cladograms
compound
a
areas) general
are
taxa.
and evaluated in
regard
of
resulting
will be
general
to
area-
every
areas.
General
intrinsic characters,
It
follows
group of
taxa
and thus
analysis.
does
not
really apply
area-cladograms
at
least
not
that
to
are
a
analogues
choice
for
history,
of
one
a
groups
a
few
areas
in
homology
or
such
areas
dispersal
as
as
a
and
Evaluation
by
area-cladograms.
responding
such represent support. Columns
since
to
share
consequently
group of
homoplasy.
or
may
directly
‘synapomorphy’
uniquely sharing
indicating monophyletic
as
primitive
analysis,
Indicators of non-shared
and
of vicariance,
involved in the
terms
taxon
which their descendants inherit and
criterion leads
area-cladograrn
extinction do
by
statements
parsimony
area-cladograms
used in cladistic
groups
monophyletic
Columns of the data matrix
events
as
sense,
of taxa, but
shared
consequence of
extinction,
the
to
interpreted
have members with genes
biogeography
group
extinction with
strict
a
the
‘general’.
more
can
dispersal,
among
these relations.
exclude each
the data matrix,
compile
one
or
recognized
are
expressing
AREA-CLADOGRAMS
unrelated groups of
more
will be
diagrams
character
than
to
cladograms pertaining
obtained. As
include
transcribed into
is used
more
in the network
mutually
GENERAL
are
taxa
will be obtained. If
on
AND
(compatibilities)
sought
are
to
vicariance
indicating dispersal
fit the area-dadogram and represent contradiction. As in cladistic
312
CLADISTICS
character
analysis,
timization
the
or
onwards
root
tell
might
such
After
whether the
us
taken
states are
state
have
to
and
minus support is
with
cladograms
the
evaluation of
number of
of
data. For
the
tions
that
be
can
used
tical consequences. It
results in
at
cladogram
will also
Assumptions
1
As
a
under
or
Assumption
0
There is
a
0
shown
by
a
less
blem is
analysis
already
that
be
to
analysis
an
under
1
analyses
under
yet another
areas
the
indicated
the
by
regard
are
may
that
analysing
to
degree
problem
is
an
two
when
prac-
groups
general
analysis
fish
poeciliid
area-
with
(p. 317).
analysis
an
for it
analysis
can
with
dichotomous,
at
using Assumption
once
data matrix with respect
occur
particular area-cladogram
a
of bad fit,
groups
inventory
especially
fail
dadogram.
in
in all other columns
fit
state
to
refering
an
may
not
algorithms
used
so
to
may
The pro-
monophyletic
terminal
area-cladogram.
indepen-
This
in which small
events.
refering
to
be
changes.
vicariance
of columns
Columns
always
of
area-cladograms
respond
to
in the
implemented
to
assump-
affected column. It follows that contradictions
the nested mutual interdependence
a
this
from
only
all columns
contradiction will
same
the
real
sense.
any
certain column in the
a
least
completely
not
are
real
to
without
not
on at
be unneeded if
It is
them
on
data used
general area-cladograms,
2 make
or
all counted in the
inclusive) monophyletic
by
when
regarding
data matrix with
1
Assumptions
problem
contradiction
caused
2
or
resolved
fully
with the
dealing
Assumption
results in
0
resulting
ones
means
hypotheses regarding
general area-cladogram,
example
an
based
are
of the
of view is
using Assumption
the best
among
some
includes
point
equally parsimonious explanations.
exaggerating
this
This
a
use
columns refer
extra
to
as
certainly
can
real observations rather than
This
hypotheses.
affected because these distributional types
tion
only
parsimony.
resolved
fully
to
i.e. the interal nodes of
taxa,
of
that if
one
prove
Nevertheless, they
result in
the total
to
distinction between
a
respective cladograms,
doubt with respect
evaluate
We show this in
more
include the
dent.
preference
make
to
area-
criterion for
parsimony
a
chosen in
us
Columns
of best fit for
degree
as
2 it is doubtful that the
or
opinion
terms
particular area-cladogram,
that
(i.e.
a
serves
criterion. We
parsimony
produces general area-cladogram(s)
further
0. Given
1
our
implies
least
consequence
Assumption
only produce
that
2.
to
in
events
biogeographical
taxa
it
enables
from the
expressing
It is
phylogenies.
reconstruct
the
express
as
root.
contradiction. Contradic-
represent
This criterion is
because it
columns,
Assumptions
assumptions
to
matrix,
outgroup
be examined
can
the
at
single origin
a
present from
equal length.
0 when all
Assumption
data but
to
measure
area-cladograms.
of
assuming
Until this
root.
op-
columns with
are
appropriate
an
by
for each col-
whether all of the columns in the data matrix should be used
question
empirical
to
a
evaluating area-cladograms by
all in
of
as
the data
to
changes (steps)
state
area-cladograms
We
chosen
respect
the
reversals,
All
which
states
changes. Choosing
at
estimated
are
changes
state
area-cladogram.
Character
really originated
indicating multiple parallel origins
tion
zero
support without
as
for each
computed
represent support.
considered
are
area-cladogram
an
the number of
optimization
be
can
change
state
one
internal nodes of
on
1970).
of the data matrix
umn
zero
states
(Farris,
3
[VOL.
As
taxa
yet,
are
no
not
solu-
far.
Results
In the
tionships
analyses given here,
cladograms
or
tradiction.
in
terms
Monophyletic
area-cladogram;
or
elements
to
general area-cladograms.
area-cladograms
sal
two
of components with respect
extinction
ad hoc
imply
of
i.e.
responding
hypotheses
we
the
to
necessary
contradiction.
emphasized. First,
compatibilities
Second,
parsimony,
taxa
are
their
to
balance of
explain
implied
events
the rela-
compile
interpret area-cladograms
vicariance
to
explore
we
in order
or
support and
imply
distribution in
area-
general
con-
support for the
terms
of
disper-
HISTORICAL BIOGEOGRAPHY
313
313
As
the
a
corollary,
note
we
in
area-cladograms
with the
area-cladograms
mon
i.e.
cause,
it
However,
that
analogous
chosen
a
of
terms
synapomorphy.
to
ing biogeographic patterns.
a
and the
by dispersal
i.e.
with the
be evidence of extinctions
in
general area-cladograms
interpreted
is
to
due
as
to
to
that
absence of
in
taxa
a
of
explainin-
‘positive’
be caused
particular
absence. We
com-
homoplasy.
mechanism for
analogy implies
primitive
or
interpret
Congruency
analogous
only
from the literature, where
cases
be used
general area-cladogram) might
‘negative’ incongruencies (the
‘reversals’) might
tion for
be
the
not
the
consequence,
congruencies (occurrences incongruent
can
Incongruency
that vicariance is
As
can
shown between them.
general area-cladogram
be stressed
must
general area-cladogram
incongruencies
areas
this evalua-
use
original
data matrices
illustrated
by comparing
unavailable.
are
CRITICAL
The
some
EXAMPLES
problems
UNDER
ASSUMTIONS
encountered
we
1 and 2
using Assumption
discussed
hypothetical examples
1
are
and
by Humphries (1982)
Humphries
and Parent!
(1986).
Figure
3
Figures 3a, b,
of
A,
=
is
NG
present
Figs.
13.iii,
the
find character
conclusions
Humphries
states
and another for
resolved
using
yields
defining
cladogram (Fig.
Taking Assumption
ed
among
1
are
A+B+D
In
cladogram.
a
can
be
similar
Fig.
Figs.
and AF
13.v
2.28e
2.28.C and 2.28.d
follows: AUS
=
=
(Australia)
(Africa)
our
our
Systematists
that defines A+B
one
their
(cf.
Fig.
=
D.
It
differs
3d)
Fig 3e). Moreover,
would be
(cf.
pleased
to
their component
This would yield
component 0).
sense
for
that
by
area-cladogram
(A+B)
a
fully
is
be
can
1
analys-
(Fig. 3a)
The
undisputable.
components from different
situation harmonizing the
from these
(Fig. 3b) generates
three
remaining
in the final
area-cladogram
2
example
Area-cladogram
consensus
Consequently,
this
this
granted,
they represent
3a represents the
supported
way,
1981)
previous paragraph.
contradictions among these components.
one
Fig.
C,
in Table 1. One component
in the
(alternate) area-cladograms. Fig.
ponents only
their
(1986,
and Platnick,
given
disputable
are
his
=
as
3.e).
(Nelson
the components
(1982,
and D,
the method outlined in the
other three
is modified
counter-intuitive.
C
A, B,
and
Humphries (1982
coding
(South America)
and Parenti
Humphries
cases
SA
B,
=
clear that the conclusion of
in both
iv of
1986. The
Parenti,
(New Guinea)
from that of
3)
e
and
Humphries
general
the components
com-
area-
given
in Table 2.
Taken
Table 3 lists the components,
together,
supporting area-cladograms (u
given
support
Fig. 3c)
possible
the
From this
sequence
list,
the
number of their
cliques present
cliques (general area-cladograms)
to
the
and differs from the conclusions
are
and Parent!
(1986;
our
choice for
clique
2
given by Humphries (1982;
Fig.
with
regard
to
(general area-cladogram
our
Fig.
3d
as
well
3e).
4
Figures
4a-c represent
and Parenti
both
four
minus contradiction leads
Humphries
Figure
including
undisputable).
in Table 4.
Evaluation of these
as
=
1986;
publications
see
Fig.
coding
differ
14,iv-vi of
in
(his Fig.
Fig.
3).
14vii
=
Humphries (1982;
In
this
our
case
Fig. 4g
=
2.29.d-f of
Humphries
also, the conclusions reached by
and their
Fig.
2.29g
our
=
Fig.
4h, respectively).
The three
(in
this
case
area-cladograms (Figs. 4a-c) yield
also,
one
the
component is undisputable,
components presented
in
whereas of the other three
Table 5
only
one
314
CLADISTICS
can
be
ponents
is
Fig. 3(a).
for
1.
a
the
supported by
given
group,
based
area-cladogram
(c).
and
Humphries
Parenti
on
a
derived from
A
Humphries (1982) for
(1986)
for
the
same
A +B
A
derived from
the
two
same
available
consensus
area-cladograms
data.
com-
(Table 7).
area-cladogram
under
Assumption
(e). generalarea-cladogram
data.
3a
under
Assumption
1.
undisputable
+C
B+D
+
are
3
1.
Fig.
B
+
these
the list of the
together,
cliques
particular monophyletic group, (b).
Table
Components
maximal
list, eight
general area-cladogram
(d). generalarea-cladogram presented by
presented by
Taken
respective area-cladograms).
in Table 6. From this
Consensus
different
[VOL.
only
one
C +D
Table
Components
derived
from
2.
Fig.
A+B+D
General
minus
Assumption
1
undispu table
B
A
+
A
+
D
+
D
B
3b under
area-cladograms
1 and 2
contradition. General
(Fig.
only
one
4d and
4e)
area-cladogram
3
is
are
alike with
second best
regard
(Fig. 4f).
to
support
These
three
HISTORICAL
BIOGEOGRAPHY
315
315
possibilities
Fig.
14vii)
do
not
15
the
Using
Humphries
possible general
analysis
present
Table
Components
B
A
+
A
+D
derived
from
+
found in
one
derived from
components
of the
fail
b
original area-cladograms. “ —
=
Humphries (1982:
these four
areas)
confirm this conclusion.
to
under
Assumption
A
D 2
1
+
B
A +B
1.
+C
I
+D
1, 2(u)
4.
area-cladograms in Fig.
”
(1986, Fig. 2.29g).
3.
C +D
2
Table
from the
Cliques
Parenti
area-cladograms (of
we
Fig. 3a,
B
2
l(u),
and
the difference in likelihood. Moreover,
recognize
concludes that all
equally likely.
are
in the conclusion of
implied
are
However, they
3a and 3b.
“
”
+
contradiction: component
=
support: component
absent in
one
of the
original
area-cladograms.
Component
1
A
+
B
1
x
CM
u;
A +B +C
+
B
1
A
+
B +D
1
x
u;
x
u;
3 A +D
A+B
4 B
D
+
X
lx
2 A
1
x
u;
+D
A +B +D
1
Contradiction
Support
x
u;
Summary
+
1
+,
CM
X
2
x
+
1
x
+,
2
x
+
I
x
+,
2
x
+
X
-
1
X
-
1
X
3
x
+
,
1
X
1
X
1
X
-
+
4
x
+
3
x
+
3
x
-
,
-
+
,
-
Table 5.
Components
5.1
A
+
B
derived
5.2A
u
A
A
+
C
+D
B
+
next
together,
the
Humphries
14.i—iii).
are
are
The distribution of
given
in Table 10. The
it lacks
value of
j
A
+
B
A
+
D
I only
B
+
D
j
j
u
one
are
A+B+D
Assumption
2 is also taken from Hum-
in Table 9, Within this list all
support
of them
of which is
being undisputable).
not
and contradiction for the
contradition. There
in
15
com-
Taken all
cliques possi-
presented by Flumphries (1982)
these authors do
area-cladogram
minus contradiction.
the
given area-cladograms (Fig. 5a-c),
presented
one
u
2
(none
general
(unambiguous)
support
1.
in Table 8
(1986). Moreover,
tions.
as
one
determined under
found, only
and Parent!
Assumption
5.3
I only
For each of the
presented
components
areas
under
\
ASSUMPTION
example (Fig. 5),
found
ble for four
C
4a-c
one
UNDER
phries (1982: Fig.
ponents
B
+
Fig.
A+B+C
EXAMPLES
The
I only
D
+
B+C
\
A+B+C
from
Fig.
are
general area-cladogram
5d is the
seven
and
mention alternative solu-
most
is
likely hypothesis
possibilities showing
4
as
the
316
CLADISTICS
Three
Fig. 4(a-c).
area-cladograms
under
cladogram
and
and
consensus
derived
area-cladograms
from
Assumption
1.
these
of three
different
area-cladograms
under
assumption, (h). general area-cladogram presented by Humphries
1.
(d,e).
groups;
(f).
the
the
two
second best
and Parenti
best
for the
(1986)
general
general
for the
3
area-
same
same
data
data
assumption.
Table
A
and
6
4a-c
2
B +D
3
A +D
3
C +D
(u), 2,
(or
3
have shown here is that
we
or
on
judging
at one
a
their compatibilities
especially
one
in those
(or
A
from
a
few) general
in which
a
list of
B
+
C
1, 2(u)
+
B
+
D
1, 3(u)
possible components
derived from each
cannot
on
the basis of
a
and
Humphries
we
can
arrive
criterion.
explanations given
and Parenti
alternate, conflicting possibilities
area-cladogram(s).
pre-
area-cladogram,
parsimony
be derived from the
(1981), Humphries (1982),
examples
+
building general area-cladograms,
few) likely general area-cladograms
Nelson and Platnick
Assumption 1.
1
by building
and
under
A
the basis of additional assumptions
comparable, equally explicit procedure
into
Fig.
B +C
in,
1
from
2
+
B
derived
A +C
What
sent
by
monophyletic
Assumption
(g). general area-cladogram presented by Humphries (1982)
Components
A
[VOL.
are
(1986),
comprised
HISTORICAL
BIOGEOGRAPHY
317
317
Table
from
Cliques
the
Fig.
1
A+B
lx
A+B+C
lx
u;
u;
lx
2 A+B
A+B+D
lx
lx
3 A+B
3x
+
2x
+,
u;
3x
u;
2x
u;
3x
C+D
lx
4 A+C
lx
lx
A+B+C
5
lx
A+B+D
6
lx
8
2x
u;
B+C
lx
lx
A+B+C
7
2x
u;
A+D
2x
u;
lx
A+C
B+D
lx
A+D
lx
B+C
lx
7.
derived
components
from
area-cladograms given
in
4a-c.
lx
5x
-
lx
+,
-
+
lx
+,
5x
-
lx
+,
-
+
+,
2x
-
+,
2x
-
CM
+
fO
+
X
X
lx
+,
2x
+,
lx
+,
+,
2x
2x
+,
CM X
+
2x
+,
4x
3x
1
2x
+,
CO
+
-
X
1
1
3x
-
3x
+,
-
-
3x
-
3x
+,
-
-
2x
-
X
+■
1
1
2x
-
4x
+,
-
Table 8
List
of components
derived
from
Fig.
area-cladograms given
5(a)
5(b)
A+B
A+B
A+C
A+C
A+C
A+D
A+D
A+D
B+C
B+C
B+D
B+D
B+D
C+D
C+D
A+B+C
A+B+C
A+B+C
A+C+D
A+B+D
A+B+D
B+C+D
B+C+D
nor
A+C
A+D
occur
B+D
B+C
can
together)
Table
Components
derived
from
Fig.
A+B
1, 2,
3
B+C
1,
2
A+C
1, 2,
3
B+D
1,
2,
A+D
1, 2,
3
C+D
1,
3
ROSEN’S POECILIID FISHES
Assumption
5(c)
A+B
(Neither
in
5a-c.
AS A
9
5a-c
3
under
Assumption
A+B+C
1, 2,
A+B+D
1,
2.
3
3
A+C+D
2
B+C+D
2,
3
BENCHMARK
0
The theoretical and
empirical implications
ofAssumption
0,
in
terms
of the
complexity
318
CLADISTICS
of the data matrix and of the number of
than those of the other
area
11
(for coding
see
two
area-cladograms
assumptions.
Rosen,
1976)
[VOL.
The
be
to
has
analysis
because it is assumed
to
evaluated,
been carried
be of
are
out
3
simpler
without
hybrid origin (Wiley
1981).
Fig. 5(a-c).
derived
Three
area-cladograms of three
from these
area-cladograms
different
under
monophyletic
Assumption
Table
Cliques
based
on
the
1
A
A
2
A
+
B
3
x
+
B +C
3
x
+C
2
x
(d).
the best
10.
components
cladograms
groups;
2.
in
derived
from
the
area
5a-c.
Fig.
+
>
6
+
1
+
X
x
+
x
+
-
,
A
B +C
3
x
+
3
x
+
+D
2
x
+
A+B
3
x
+
+
5
1
X
-
,
3
A+B
C
1
X
-
5
x
A
5
2
x
+
A
+C
2
x
+
A
+C +D
+
B+D
2
x
+
A+C
3
x
+
B
2
x
+
2
x
+
3
x
+
3
x
+
1
X
-
1
X
-
1
X
-
5
x
+
4
x
+
,
6
+
D
X
-
X
-
,
,
,
1
+
,
4
1
,
2
X
-
1
X
-
,
1
X
1
X
-
,
or
7
8
9
10
A +D
,
5
2
x
+
A
D
3
x
+
A+C+D
I
X
+
,
2
x
1
X
+
,
2
x
A+D
3
x
+
or
2
x
+
1
X
2
x
+
,
1
X
B +C
2
x
+
,
A+B+C
3
x
B +C
2
x
B +C
x
+
,
A+B+D
+
-
1
,
X
-
5
x
1
+
,
-
4
x
+
X
2
x
2
x
1
X
-
-
,
-
-
,
-
4
x
+
,
1
X
-
-
5
+
x
+
-
,
11
B+C +D
+
1
X
3
x
A+B+D
2
x
+
A+D
3
x
+
1
X
+
2
x
+
,
1
2
+
X
X
-
-
3
x
+
,
12
B
+
D
1
X
5
-
x
+
,
13
A+C +D
C +D
x
1
X
2
X
3
x
2
x
-
-
,
2
X
1
X
CM
X
1
X
-
1
X
-
-
4
x
+
,
14
3
,
+
-
,
-
,
A+C+D
1
X
+
C +D
2
x
+
B +C +D
2
x
+
-
3
x
+
-
,
15
,
,
4
x
+
,
-
general area-cladogram
HISTORICAL
319
BIOGEOGRAPHY
319
Fig. 6(a).
of Heterandria.
Cladogram
cladogram of Xiphophorus; adapted
(b).
Table
Species
dria and
text
for Heteran-
Xiphophorus.
Heterandria
Heterandria
1
(1976, 1978).
11.
and numbers used in
names
from Rosen
Xiphophorus
pigmaeus
pigmaeus
1
attenuata
attenuata
2
2
jonesi
2
2
3
3
litoperas
3 montezumae
3
montezumae
4
obliqua
4
4 cortezi
5
anzuetoi
anzuetoi
6
6
cataractae
cataractae
nigrensis
5
clemenciae
5 clemenciae
7
7
dirempta
8
8
bimaculata
6 alverezi
6
alverezi
7
“PMH”
8
8
signum
9
9 helleri
helleri
The distribution over
see
Table
11).
The
terminal taxa and 7
these
two
binary
areas
phylogeny
(internal) clada,
matrices
each cladon in the
is coded
a
as
a
areas
(Table 14)
From this data
17).
matrix,
A search for the
clique
and
one
8 represents
areas’. The
authors
The
a
true
It is
taxa
of mutual
compiled
among
represents
over
areas
the boolean
combining
(components)
can
be extracted
compatible components reveals
The
areas are
similar
to
those
for
pro-
distributions
indicated in each column of Table
sets
From
(Table 12).
distribution
from
names
shows 8
6a),
one
12.
(Table
maximal
wide-spread species
areas
2 and 3
are
‘sister
postulated by previous
1981).
of Xiphophorus
represented
of Xiphophorus
combinationof these
Fig.
form
binary
area-cladogram (Fig. 7d).
the other
(for species
(1975:
component under Assumption 0 and, therefore,
(e.g. Wiley,
phylogeny
set
Rosen
derived, giving
monothetic
resolved
by
The data matrix
16a).
terminal
partial
largest
relations
species
19
completely
and 8 clada, and is
for the
for all
given
and is also coded in
duct of the matrices in Tables 12 and 14.
over
as
data matrix is
cladogram (Table
matrix in Table 14
binary
of Heterandria
two
is
as
given by
in
binary
given
matrices
Rosen
in Table 15
gives
(1976: Fig. 6b),
shows 9 terminal taxa
form in Table 13. The distribution over
(for species
another matrix
names
indicating
see
Table
areas
11).
The
the distribution over
CLADISTICS
320
areas
for each cladon in the
monothetic
of
sets
areas
cladogram (Table
can
be derived
16b).
(Table
Table
From this data matrix,
18
3
partially
18).
12.
matrix for Heterandria
Binary
[VOL.
cladogram (Fig. 6a).
clada
1
2
3
4
1
1
0
0
0
0
0
0
0
0
2
0
]
0
0
0
0
0
0
0
3
0
0
1
0
0
0
0
0
0
0
0
�
0
0
0
1
0
0
0
0
0
0
5
0
0
0
0
1
0
0
0
0
6
0
0
0
0
0
1
0
0
7
0
0
0
0
0
0
1
8
0
0
0
0
0
0
0
SPECIES
6
5
7
Table
Binary
matrix for
8
10
9
11
12
13
14
15
0
0
0
0
0
1
0
0
0
0
I
1
0
1
1
1
0
1
1
I
1
0
1
I
1
1
1
0
1
1
1
1
I
1
0
1
1
1
1
1
1
1
1
1
I
1
1
1
1
1
13.
Xiphophorus cladogram (Fig. 6b).
Clada
SPECIES
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
1
1
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
2
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
3
0
0
1
0
0
0
0
0
0
0
I
0
0
0
0
1
I
4
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
1
5
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
6
0
0
0
0
0
1
0
0
0
0
0
0
0
1
1
1
1
7
0
0
0
0
0
0
1
0
0
0
0
0
1
1
1
1
1
8
0
0
0
0
0
0
0
1
0
0
0
1
1
1
1
1
1
9
0
0
0
0
0
0
0
1
0
0
1
1
1
1
1
1
Table
14.
Distributions
of Heterandria
0
species (after Rosen,
1976; 1978)
but
excluding
area
1
1
11.
areas
SPECIES
1
2
3
4
5
6
7
8
9
10
1
0
0
0
0
0
1
0
0
0
0
2
1
0
0
0
0
0
0
0
0
0
3
0
0
0
0
0
0
0
0
1
0
4
0
0
0
1
1
0
0
0
0
0
5
0
0
0
0
0
0
0
0
0
1
6
0
0
0
0
0
0
1
0
0
0
7
0
0
0
0
0
0
0
1
0
0
8
0
1
1
0
0
0
0
0
0
0
HISTORICAL
BIOGEOGRAPHY
321
321
Table
Distributions
of
15.
Xiphophorus species (after Rosen, 1976, 1978)
but
area
excluding
11.
areas
SPECIES
1
2
3
4
5
6
7
8
9
10
1
1
0
0
0
0
0
0
0
0
0
2
1
0
0
0
0
0
0
0
0
0
3
1
0
0
0
0
0
0
0
0
0
4
I
0
0
0
0
0
0
0
0
0
5
0
0
I
0
0
0
0
0
0
0
6
0
0
0
1
1
I
0
0
0
0
7
0
0
0
0
0
0
0
0
1
1
8
0
0
0
0
0
0
0
1
0
0
9
0
1
0
0
0
0
0
0
0
Fig.
7(a). Area-Cladogram
under Assumttion
based
It differs in
and 9.
(e.g.
two
components
imum size
A
two
were
one
appears
is
Fig.
types.
As
8. Three
The
the
at
are
found
of
a
two
or
more
both
derived
area-cladogram (Fig.
positions
of
3, 6,
areas
is absent from this
Xiphophorus
presented by previous
meet a
in
based
Fig.
on
in
and
general area-cladograms
15
in
Heterandria
Fig.
for
in
authors
b and
analyzed together,
c
start
no
also from
need for
analysing
and
the
Fig.
a
ofmax-
combination of
analyses
a
of each
not
be
reduced but
Assumption
under
Assumption
area-cladogram
1
areas
or
2.
0.
for Heteran-
8b is almost similar
8c differs from the
genus
selection of distribu-
under
Xiphophorus
to
in
cliques
Fig. 8).
for the subclade of
areas
the
to
4-6 which is
fully
general area-cladogram
2, 3, and 8.
general
contradictory
figs,
and
data matrices need
8a is identical
evaluation of the three
supportive
can
general area-cladogram
of the subclade
b)
where separate
case,
respective
terms
quantitative
taxa
different situation when
general area-cladograms
whereas the
of
particular
general area-cladogram
in
16a and
for each genus, there is
in 8a
26
in the
From these components, three
groups
consequence, the
We
19).
resolved. The
A
those
combined (Tables
Xiphophorus area-cladogram (Fig. 6b) except
implies
to
(the general area-cladograms
general area-cladogram
(Fig. 6a),
only
becasue
root
similar
basically
be defined(Table
area-cladogram
joined directly.
dria
7
data matrices
can
joint analysis
tional
Xiphophorus;
on
resolved
partially
one,
from that of Heterandria,
reduced data matrices. In this
yield only
based
1981).
When the
26
area
area-cladogram
Wiley,
results in
analysis
several respects
Furthermore,
This
area.
(b) area-cladogram
0.
As with Heterandria, the
7b).
Heterandria.
on
0
area-cladograms
events,
imply
to
give
an
shows that
Fig.
overall value of 11.
overall values of 9, and
12. Thus
8a
The
Fig.
CLADISTICS
322
[VOL.
3
8c has the best value of support minus contradition and also has the minimum number
of 39
state
changes.
The
other
two
both have
Table
Transposed
9-17
species,
N.B.
data matrix under
clada;
For technical
the
reasons
on
columns
the
are
right
state
refer
horizontal
to
changes.
16,
1-8
Heterandria:
Assumption 0;
numbers
40
the
and
species,
9-15
constituting species
clada; Xiphophorus:
of the
1-9
clada.
respective
vertical.
rows
(a) species
/clada
column
areas
1
nos.
2
3
4
5
6
7
8
9
10
i
0
0
0
0
0
1
0
0
0
0
2
1
0
0
0
0
0
0
0
0
0
3
0
0
0
0
0
0
0
0
1
0
4
0
0
0
1
1
0
0
0
0
0
5
0
0
0
0
0
0
0
0
0
1
6
0
0
0
0
0
0
1
0
0
0
clades
7
0
0
0
0
0
0
0
1
0
0
8
0
I
1
0
0
0
0
0
0
0
9
0
1
1
0
0
0
0
1
0
0
7+8
10
0
1
I
0
0
0
1
1
0
0
6
11
0
1
1
0
0
0
1
1
0
1
5 +6 +7 +8
12
0
1
1
1
1
0
1
1
0
1
4+5+6
13
0
1
1
1
1
0
1
1
1
3+4+5+6+7+8
14
1
1
1
1
0
1
1
1
2+3+4+5+6+7+S
15
I
1
1
1
1
1
1
1
1
1
1
+2 +3 +4
1
I
0
0
0
0
0
0
0
0
0
2
1
0
0
0
0
0
0
0
0
0
3
1
0
0
0
0
0
0
0
0
0
4
1
0
0
0
0
0
0
0
0
0
5
0
0
I
0
0
0
0
0
0
0
6
0
0
0
1
1
1
0
0
0
0
7
0
0
0
0
0
0
0
0
1
1
8
0
0
0
0
0
0
0
1
0
0
+2
1
1
1
+
7+8
+
7 +8
+5 +6
+
7 +8
(b)
9
0
1
0
0
0
0
0
0
0
0
10
1
0
0
0
0
0
0
0
0
0
1
11
1
0
0
0
0
0
0
0
0
0
3+4
12
0
1
0
0
0
0
0
1
0
0
13
0
1
0
0
0
0
0
1
1
I
7 +8+9
14
0
1
0
1
1
1
0
1
1
1
6
15
0
1
1
1
1
1
0
1
1
1
5 +6
16
I
1
1
1
1
1
0
1
I
1
3 +4 +5 +6
17
1
1
1
1
1
1
0
1
1
1
1+2+3+4+S+6+7+8+9
Choosing Fig.
by assuming
only
the
more
in
or
8c
(also Fig. 8a) implies
sequence
species
ad hoc
areas
a
in
area
hypotheses.
of vicariance
8 has
Apart
1-4, the absence of
extinction. We may
not
tip
an
that the
events
responded.
from three
endemic
the balance
phylogeny
and
8+9
+
7 +8 +9
7 +8
Xiphophorus
the
sympatric speciation
in
area
by assuming
the
+9
+
7 +8 +9
of Heterandria can be
allopatric speciation
For
species
+
7
might
first
explained
events
in which
implications require
events
be
for the
primitive
hypothesis
and
species
absence
deleting
HISTORICAL
1987]
the latter
as
it
spread species
area
9,
or
its
supposes
in
area
present
implying primitive
implying
primitive
one
7
distribution
absense
have
not
absence in this
in
323
step and is therefore less parsimonious.
extra
might
BIOGEOGRAPHY
be the
might
Species
area.
the
to
responded
latter
or
it
the vicariance
result of
a
later
5 has either
might
dispersal
into
dispersed
have
become
The wide-
event
separating
into
area
extinct
area
9
3 thus
in
areas
2+4-6+8+10.
Table
Partial monothetic
under
17.
of
sets
areas
Assumption
distributional
for Heterandria
0.
types
components
i
2
6
I
7
6
8
7
9
3
10
5
2,
3
4,
5
8
4
8
2, 3,
9
8
2, 3, 7,
2, 3, 7, 8,
10
10
11
2, 3, 4,
5, 7. 8,
10
2, 3, 4,
5, 7, 8,
9,
1, 2, 3, 4, 5,
12
10
13
14
10
7, 8, 9,
10
1, 2, 3, 4. 5, 6, 7, 8, 9,
Table
Partial
monothetic
sets
of
15
18.
areas
Assumption
distributional
for
types
1, 2, 3, 4,
2
9
3
5
8
2,
7
6
6
10
9,
4,
8,
5, 6,
1, 2, 3,
the
13
2, 4, 5, 6, 8,
2. 3,
be
11
12
10
2, 8, 9,
Choosing
10,
8
8
4, 5,
can
under
components
i
9,
Xiphophorus
0.
10
9,
10
4, 5, 6, 8, 9,
general area-cladogram
explained by assuming
14
a
15
10
in
sequence
Fig.
16,
8b
17
implies
of vicariance
that the
events
Xiphophorus phylogeny
in which
species
in
areas
CLADISTICS
324
6 and 7 have
not
For Heterandria
,
distribution of the
in
species
area
area
1
absence in
in this
areas
in
species
might
area
a
area
8
primitive
absence
3
assumed, implying
be the
must
absence in this
2-5+8+10,
it has
or
into
Table
sets
of
distributional
2
2
9(X)
3
5(X)
1
3(H)
have become
8(H)
5
4(H)
8
12
10
4,
5, 6
8
2, 3, 7,
2, 8, 9,
8
10
10
2, 4, 5, 6, 8, 9,
10
Assumption
Platnick
(X)
6
(X)
13
(X)
11
(H)
14
(X)
15
(X)
2, 3, 4, 5,
10
13(H)
7, 8, 9,
1, 2, 3, 4, 5, 6, 8, 9,
10
16,
1, 2, 3, 4,
10
14(H)
5, 7, 8, 9,
2, 3, 4, 5, 6, 7, 8, 9,
difference with
parts of the
determines the
of the
3
for the
(X)
(X)
7
10
implies
and 7
8
2, 3, 4, 5, 6, 8, 9,
general area-cladogram
8b
(X)
12(H)
in
species
species
8c
Fig.
from
15(H)
a
of the
choice here for
events
to
be
implies only
area
events
from
(X)
is that he infers
Wiley’s (1981) analysis
unique
unique
10
17
area-cladograms
general area-cladogram. However,
occurrence
11
10(H)
10
2, 3, 4, 5, 7, 8,
1,
Fig.
and
9(H)
2, 3, 7, 8,
cladograms
(H)
0.
5(H)
3
9,
incongruent
Heterandria
1, 2, 3, 4, 10,
(H),
7
2, 3,
to
into
might
(H)
9
4,
important
on
(H),
8
2,
9
The
area
6(H)
10
1,
3.
dispersed
components
i
2,
in
into
it has
3
area
that the present
9, again implying primitive absence
Assumption
types
7
for all
based
under
6
the
or
be assumed.
can
19.
areas
Xiphophorus (X)
a
7
dispersion
in
species
area
area
T5+8-10
areas
The
area.
dispersed
result of
in
3
area.
Partial monothetic
An
be
can
have become extinct in
6, implying primitive
extinct in
and thus
responded,
primitive
[VOL.
5 in
with
areas
1,
area
3
3.
and
one
unique
Choosing
to
genera
of the
explained.
one
regard
two
area-
choosing
the
affecting Xiphophorus,
event
the
events
producing
possible general
For instance,
Heterandria,
unique
without
general area-cladogram
i.e.
occurrences
in
areas
6, respectively.
1
(1981)
analysis
cladograms
considered the reduced
under
is
not
Assumption
1.
part of this
However,
area-cladogram
removal of
Assumption
but
a
obtained
incongruent
form of
by
Rosen
equivalent
subtrees from
consensus tree
analysis.
area-
It is
HISTORICAL
BIOGEOGRAPHY
325
325
a
misconception
tially
resolved
formative
in the
suggest
required
Assumption
and
extract
must
parsimony
are
nos
be
lead
to
do
terminal
corresponding
(table
taxa
to
shown in the
the
same
or
only
parin-
fully
compatibili-
next
example.
columns as
given
in Table 20a. The
given
following
manner.
From
column numbers 4 and
16a,
areas
i.e. the
column numbers 7 and
20a column numbers 1, 2, 5, and
given
Assumption 0),
in Table 21. In
four
new
comparison
8)
singletons (Table
distributional
to
An
6).
Table 17
components
are
of the
11)
analysis
wide-spread
of Heterandria
(the components
present
(i.e.
areas
yields
derived
2+8, 3+8,
2+3+5+7+8+10).
Table
Additional columns
the component
those
all subsets of
incomplete
be resolved into
3, 4, 7, and 8) and combine them with the distributions of prospective
(Table
2+3+4+7+8+10,
as
(Table 16a) plus
16a, species/cladon
under
to
comprises
terminal
the components
this,
applied
sistergroups (Table
taxa
only
cannot
they
derived from those in Table 16a in the
wide-spread
the distributions
column
can
and that
To be able
data matrix under Assumption 0
columns in Table 20a
20a,
incongruencies
1 the data matrix for Heterandria
those columns with
we
that
area-cladograms
(dichotomous) hypotheses.
ty method is
Under
to
general
(transposed horizontally)
20.
for Table
16 for
analysis
under
Assumption
1,
Heterandria,
Xiphophorus.
species
species
areas
areas
1
1
4
6
7
8
23456789
2
3
5
9
10
10
Heterandria
Heterandria
(a)
11
1
1
1
1
0110101101
0
1
0
0
0
1
2
0
1
1
1
1
0
0
0
1
0
1
3
1
0001000000
0
0
0
0
0
0
0
0
0
4
0
0
0
0
0000100000
1
0
0
0
0
0
5
1
0010000100
0
0
0
0
0
0
0
1
0
6
6
0
0
0100000100
0
I
0
0
0
1
0
0
7
7
0100000000
0
1
0
0
0
0
0
0
0
0
8
8
0
0
I
0
0
0
0
0
0
0010000000
0
(b)
based
on
species
4
based
on
on
species
8
8
on
on
species
6
on
on
species
7
7
Xiphophorus
1
0
0000010000
0
0
0
0
1
0
0
0
0
2
1
0000100000
0
0
0
0
0
0
0
0
0
3
0
1
1
0
0
0
0000110000
0
0
0
0
4
0
0
0
1
0
0
0
0
0
0001000000
0
5
0
0
0
0001010000
0
0
0
1
1
0
6
0
0001100000
0
0
0
0
1
1
0
0
0
7
7
0
0
1
1
0
0
0
0
0
0
1
0
1
11
1
8
0
0
1
1
0
0
0
1
0
0
0
1
11
1
1
9
0
0
1
1
0
0
0
0
1
1
0
0
1
11
1
10
0
1
0
0
1
0
0
0
0
0
0
1
1
11
1
11
0
1
10
0
1
10
0
1
10
0
111
1
1
1
12
0
0
1
1
0
0
1
0
0
0
1
0
11
1
13
0
0
0
0
0
0000000001
0
0
0
0
1
14
0
0
0
0000000010
0
0
0
1
0
15
1
0100000101
0
0
0
0
0
0
0
1
1
16
0
0
0
0
0100000110
1
0
0
1
1
0
0
From this component array, 9 maximal
cliques
values of support minus contradiction for the
0
can
be
complete
based
extracted,
based
each
data matrix
showing equal
(Fig. 9). Figure
9d
CLADISTICS
326
is the
derived under
area-cladogram
5, 6, 8, and 9, because they separate
to
be
one
area
(Figs. 9a, 9d,
for
areas
307 for
a
species
endemism),
and
2, 3,
in
one
and
8)
one
9g
the
each show
same
support
one
as
tion
monothetic
1;
(16)
sets of
refers
distributional
to
areas
Table
a
best fit
on
(20)
Heterandria under
to
Table
types
7(16)
9
3(16)
10
5(16)
2,
3
8(16)
4,
5
4(16)
2,
8
4
(20)
3,
8
3
(20)
9(16)
8
8
10
2, 3, 4, 7, 8,
10
2, 3, 5, 7, 8,
10
4, 5, 7. 8,
10
2, 3, 4, 5, 6,
Fig.
under
10
9.
The
nine
the data matrix
16b), together
(16)
2
(20)
1
(20)
10
area-cladograms
14
(16)
15
(16)
of Heterandria
derived
1.
comprises
with those in
monothetic
(16)
13(16)
10
7, 8, 9,
Assumption
10
11
12(16)
1, 2, 3, 4, 5, 7, 8, 9,
partial
extinction for the
(16)
8
2, 3, 4, 5, 7, 8, 9,
new
p.
components
I
2, 3,
define
(see
Assump-
6(16)
2, 3, 7, 8,
Xiphophorus
=
0
homoplasies
20a.
7
2, 3, 7,
In
It lacks
2(16)
2, 3,
(Table
(Fig. 9a).
alternatives
Assumption
21.
i
1.
area-cladograms
possible
9a.
6
0
three
homoplasy (reversal
Fig.
based
16a
authors considered
possible general
by having
3
area-cladograms 2, 3,
by ignoring
(which previous
the data matrix for
shows
Table
Partial
other
against
area-cladogram
9d and
Figs.
4 from 5
end up with three
When evaluated
and 8.
area
we
These differ from
9g).
justification)
while
entirely
of
0,
Assumption
areas
[VOL.
sets
the
Table 20b.
(components)
same
The
columns
given
under
newly incorporated
derived under
Assumption
Assumption
columns all
1
(Table 22,
HISTORICAL
BIOGEOGRAPHY
327
327
2+8+9,
areas
2+8+10,
2+4+8+9+10,
2+5+8+9+10,
2+6+8+9+10,
2+4+5+8+9+10,
2+4+6+8+9+10, 2+5+6+8+9+1+0
Table
Partial
tion
monothetic
1; (16)
refers
distributional
of
sets
to
areas
Table
22.
based
16b and
Xiphophorus under Assump-
on
(20)
to
types
Table
components
i
1, 2, 3, 4,
2
9(16)
3
5(16)
4
4(20)
5
2
(20)
6
1
(20)
8
8(16)
9
10
4,
6
5(20)
5.
6
3
2,
8
12
10
6
6(16)
9
16
10
10
Fig.
8
2, 6, 8, 9,
10
7(20)
10
12
(20)
2, 4, 6, 8, 9,
10
11
(20)
2, 5, 6, 8, 9,
10
9
10
10
45 maximal
cliques
the
Assumption
the
complete
for
14
(16)
15
(16)
17
16,
can
0 data matrix
separated (as
are
(20)
be determined, 36 of which show
(Fig.
Heterandria),
data matrix and
(16)
using
the
complete
those
10). By ignoring
we
one
end up with 9
on
the
equally
data matrix,
cliques
area-cladograms
Assumption
0
and
in which
when
data matrix
(see
8b).
10.
for
The
on
(20)
(20)
2, 4, 5, 8, 9,
values of support minus contradiction when
Fig.
used
(16)
10
1, 2, 3, 4, 5, 6, 8, 9,10
on
13
9,
2, 3, 4, 5, 6, 8, 9,
4 and 5
(20)
10
8,
(20)
15
10
2, 4, 5, 6, 8, 9,
area
(16)
4, 5,
Within these sets,
(16)
(20)
2, 8,
2, 5,
11
10,
7(16)
2, 4, 8, 9,
evaluated
(20)
6(20)
2, 8, 9,
using
(20)
5
2, 8,
3 when
14
13
4,
9,
good
20b.
The three
Assumption
analysis
area-cladograms
of Xiphophorus derived under
Assumption
1, evaluated
on
the data matrix
0.
of Heterandria
and
Tables 16a, b and 20a, b. The
Xiphophorus together comprises
complete
range
of components
a
are
data matrix based
given
in Table 23.
328
CLADISTICS
Table
Partial
monothetic
Xiphophorus (X)
(20)
refers
to
sets
under
of
areas
Assumption
The
Heterandria
refers
to
(H16),
9
(XI6)
3
5
(X16)
4
4
(X20)
5
2
(X20)
6
1
(H16),
7
6
(H16)
8
7
(H16),
9
3
(H16),
14
(X20)
5
(HI6),
13
(X20)
2,
3
8
(H16)
4,
5
4
(HI6),
4, 6
5
(X20)
5,
6
3
(X20)
2,
8
4
(H20),
3,
8
3
(H20)
7
(XI6)
4, 5,
10
6
6
(XI6)
2, 3,
8
9
(H16)
2, 8,
9
16
(X20)
15
(X20)
2, 8,
10
2, 3,
7,
8
10
10
(HI6)
13
(XI6)
(HI 6)
10
11
2, 4, 8,
9,
10
10
2, 5, 8,
9,
10
8
(X20)
2, 6, 8,
9,
10
7
(X20)
2, 3, 4,
7, 8,
10
2
(H20)
2, 3, 5,
7, 8,
10
1
(H20)
2, 4, 5,
8, 9,
10
12
2, 4, 6,
8, 9,
10
11
2, 5, 6,
8, 9,
10
9
2, 3, 4,
5,
2, 4, 5,
6, 8,
2, 3, 4,
5, 6, 8, 9,
10
2, 3, 4,
5,
10
1, 2, 3,
4, 5.
6, 8, 9,
1,
2, 3,
4, 5,
7,
1, 2, 3,
4, 5,
6, 7, 8, 9,
7, 8,
four best
complete
(H)
Table
and
16 and
components
2
2, 3,
a
on
(16)
2
2, 8, 9,
11.
1;
types
10
Fig.
23.
based
i
9,
3
Table 20.
distributional
1 from
[VOL.
7, 8,
9,
(XI6)
15
(XI6)
13
(HI6)
16,
10
general area-cladograms
data matrix.
(X16)
6
(X20)
12
(XI6)
(X20)
12
10
8
11, (XI6)
(X20)
14
8, 9,
(X20)
10,
(X20)
10
10
1
4,
(X20)
10
7, 8, 9,
1, 2, 3,
(HI6)
17. (XI6)
14
(HI6)
15
(H16)
of Heterandria and
Xiphophorus
derived
under
Assumption
HISTORICAL
BIOGEOGRAPHY
329
329
From this list 54 maximal
data matrix,
complete
best value of
tional
support
of
pattern
i.e.
shows
areas
derived from
9 and
one
second with
Figs.
8d and 9d.
is among the best
When
patibility
d.
Fig.
nick
we
can
reveals 3
9a is
they
to
be due
doubt
specific
seems
must
to
to
an
lead
matrices
far
are
column from
(1986)
general
area-
8c and 9d,
Figs.
the
0
(Fig. 8c)
to
which
when the evaluation
b
Figs. 8a,
two
reduced
a
incompletely
Plat-
resolved
several
analysis yields
our
com-
Figs. 9a,
Humphries (1982),
one,
In contrast,
build
identical with
are
give only
to
their mutual
Exploring
to
ad
our
that their method
hoc
to
procedure
hypotheses.
number of
the
too
be
possibilities,
wide-spread
(Table
For
wide-spread
a
16b)
columns
taxa
all
we extract
1-6)
possible
those derived under the former
Assumption 0,
1, and
vs
39
vs
vs
67
It is clear that
109).
As
a
subsets
23
53;
vs
9
vs
but
for
simple
briefly
the
it
16)
are
data
describe
corresponding
column 6 in
species
all its
extract
we
as
results.
These
areas.
2
cases,
following
will
we
possible
with the distributions of
Assumption
2.
areas
the
This
(e.g.
itself)
area
procedure
area
is
7 in Table
and combine
16b).
under this
79 columns under
vs
aspects
be resolved
Xiphophorus (Table
(e.g.
Heterandria 15
Assumption 2).
Therefore,
some
can
first take the
missing
(Table
the
by
Then
(here only
significantly
(for
consequence,
Heterandria 1
vs
vs
0
6).
in
even
and combine these
33 columns under
Heterandria 19
exponentially (for
bination 3
two
and 257 columns under
accordingly (For
bination 26
area
assumption compared
columns under
Assumption 2;
Assumption 1,
for their combination, 32 columns under
Assumption 2;
Assumption 1,
Assumption
in
conflicts
missing
entirety
we
trees.
doubted
Assumption
criticism, Assumption
and
taxon
for
they regard
illustrated
taxa
in both genera. For
The size of the data matrices increases
columns under
this
Assumption
these with the distributions of all other clada
23 columns under
consensus
i.e.
because
obtain the columns for
to
wide-spread
Xiphophorus)
as
be shown here in their
compiled.
(Table 20b,
for all
from
areas are
preference
a
is true;
reverse
practice
subsets of
all other clada
uses
advance,
Apart
the data matrix derived under
areas
missing
data matrices for Helerandria and
to
large
the
in actual
apply
and
taxa
and others have
(1981)
wide-spread Xiphophorus species
to
two
are
even
homoplasy.
less
one
Table 16b for the
15 for
with
The best
assumption.
columns added for
they might
repeated
of
the data matrix.
aspects of the data in
unmanageable
extended with
how
one
area-cladograms.
the fact
to
here is hard
Assumption 2,
Under
distribu-
other differs from the first
Assumption 1,
concerning wide-spread
be ad hoc. In
they might
implemented
leads
a
8b i.e.
support but differ with respect
corresponding
Nelson and Platnick
beginning.
of the data
or
2 all data
Assumption
2 which
general
show
Fig.
analysis using Assumption
an
under
Parenti
and
under this
dichotomous
same
columns in
better because it has
two
2
Assumption
In
only
there
matrix,
derive 26
components.
subsequently
general area-cladogram
from the
from
analysis
an
assumptional
Humphries
and
completely
comes
the
general area-cladograms.
slightly
(1981),
0 but the
the basis of the
sequentially.
0 data
have
They
the distributional types
use
we
from
ones
excludes
data matrix
branching
but
lla-d),
first is identical with
Assumption
Assumption
The best solution
homoplasy.
or
10
the
Fig.
The
11a, b).
on
four of them have the
assumptional columns,
better than the other. The first is identical with
cladograms,
includes
(Fig.
4+5
areas
When evaluated with
1
including
When evaluated
present.
are
the
minus contradiction (=10:
general area-cladograms
in that it
cliques
and
for
178
Assumption
Xiphophorus
0,
17
columns under
56 columns under
Assumption 0,
The number of components increases
18
Xiphophorus
vs
29
vs
the number of maximal
123; for Xiphophorus
1
vs
45
vs
85;
for their
cliques
2613;
com-
rises almost
for their
com-
8431)!
complex problems analysed
under
Assumption 2,
i.e. with several
species
CLADISTICS
330
having
a
range
of 2
or
more
When these
with those in
because
evaluated
are
Figs 9b,
4
areas
second best
two
of all
and 5
the
as
123
sister
areas
It is
same
of
the
Xiphophorus
topology
results in 2613
an
data matrix
cladograms, corresponding
with those in
sister
as
in
Fig.
against
end up with the
we
areas,
an
the
give
the selected distributionalpatterns
An
of Heterandria and
analysis
(257 columns)
tion
and 5. For the
shortest
to
Fig.
for
and
Assumption
in
15
state
changes
that supports
4
against
and
states
6
no
area-cladograms.
equally good
second best
the
to
0 revealed
of
3
those without
area-
in
Fig.
the
best
cladogram
10, which is
Fig.
area-
(223-231
the three shortest
are
topology
By ignoring
10.
showing
they
Xiphophorus
4
and 5
(451
they
are
let
part of the
Xiphophorus species
different
for
positions
It
cladogram
our
in
areas
area
4
areas
identical
to
and
that
our
only
figure
to
8a
as
a
Humphries
and
results
(1981),
possible
that
the
Parenti
are
(1986)
four
for
areas
general
4
area-
is the
(value 250)
most
or
surprising,
with the
might
positions
Humphries
but
Heterandria
assumption
be due
is
the fact
area
not
strictly applied,
mention the
hypothesis,
species
in
general
because under
area
8, might
2
8 and
10 and 3 should also be taken
these authors do
re-
our
Assumption
in
species
to
it
of
provisions
under
differ
they
as
and
general area-cladograms,
wide-spread
wide-spread
Evalua-
area-cladograms
Humphries (1982),
general area-cladograms
9 and 2,
also
One of them
Compared
valid alternative
on
general
are
8b and
13).
7. However, when this
us
there
3 and 9 take different
Discussion
biogeographic analysis,
3
because
combined data matrix
sequential branching
areas.
Fig.
1
general area-cladograms.
‘best’
two
and based
together
Assumption
area-cladograms (Figs
complete
a
show
sister
as
steps;
10 and 2,
component 2+3, based
Platnick
Both
range of the
areas
strange
seems
in
only
same
dichotomous
gives
taken
obtained under
with
obtained their results.
their small number of
they apparently
Xiphophorus
as
from the
matrix
mention
(1986). They
data matrix,
account.
come
from those obtained by Platnick
mains unclear how
because
areas
result
same
the number of area-cladograms
and Parenti
2 the
resolved
gives
the latter
(242, 245, 246, 250)
general area-cladogram
remarkably
the
used
241, 478 steps).
=
four values
next
all
cladograms,
data
complete
(homoplasy-support
that
141)
identical
cladogram
completely
generates 8431
the
against
As
11
area-
from 64-72.
homoplasies
supporting
completely
of Heterandria and
analysis
reduced data matrices will
9a).
11 third best
are
are
events,
8b.
It is obvious that
on
15
(178 columns)
142). Among
All three have
accepted
There
9.1. When evaluated
Fig.
has
be
contradictory
of the
out
emerge
identical
9a.
Fig.
the
5
Evaluation
that in
as
area-cladogram
81-90 support, balance
homoplasies,
one
90 support, balance
homoplasies,
cladograms (302 steps).
only
ones
are
areas.
that has the least total number of
data matrix
complete
There
areas.
from 49-54 and their
possibilities.
cannot
sister
and 68
states
sister
as
as
3
area-cladograms.
better
two
Assumption 1, they
rather than
also the
with
corresponds
analysis
against
data matrix
supporting
4 and 5
range
0 data matrix the best
cladograms (231
10.
with 54
values
It has the
resolved
completely
complete
sequentially
area-cladogram
one
which
A separate
Evaluation
off
area-cladograms.
areas.
Assumption
homoplasies
support
number of
unmanageable
noted earlier under
as
branch
support
them is the
Among
(108)
not
an
contradiction (65). These
support (52) and
area-cladograms
Their
cladograms.
but
c
and 5
but these also do
to
the basis of the
on
the balance between
regarding
leads
areas,
of Heterandria results in 123
Independent analysis
[VOL.
the
into
area-
Assumption
be
true.
and Conclusions
distinguish
i.e. those of Rosen
three
(1976),
different
Wiley (1980,
methods for
1981),
and
historical
Nelson
and
(1978).
Rosen’s
(1976, 1978)
in which the
method is the construction of reduced
incongruent
parts of the
consensus
compared area-cladograms
area-cladograms,
are
deleted. These
HISTORICAL
BIOGEOGRAPHY
331
331
reduced
fect all
area-cladograms
study
that observed
imply
(Wiley, 1981,
groups
p.
lose information. The first aim of historical
of the
the
character
aims
at
reconstructing
area-cladograms
in
might happen
of
from
of
Furthermore, speciation
general
a
more
by
general
more
should be
events
Brooks’
uses
missing
(Swofford 1985)
questions posed
(2)
new
the
what
are
the
far
objections
number of
that
can
simonious
of
groups
much the
arising
current
judged
factors?
well resolved
the
are
taxa
from the
as
as
solely
amply
in
practice,
aims
at
conclusion,
main
two
areas,
partly
far
so
consequence
estimating
com-
Wiley’s
removes
minimum
a
events.
parsimony
to
our
biogeographic
biogeographic
in the initial data and
gives
the
are
consensus
being
con-
encountered
general
notoriously
area-
unpar-
We have demonstrated that contradiccan
be resolved
in
perfectly
a
general
criterion.
of Nelson and Platnick’s
it leads
as
constructing general
demonstratedthat
area-components
the
fully implementation
undertake historical
to
a
at
aimed
are
the
common or
for
Program
between
relationships
and
ours,
the PAUP
results obtained
ours
codes for
binary
answering
with
method
parsimony
to
different from
using
many
Assumption
2
general area-cladograms.
because the method described here resolves conflicts present in
sent
derived
the basis
unique
coding strategies
loss of information besides
a
incompatible
strictly applied
In
what
analysis
an
on
on a
conflicting biogeographic patterns
of the considered
by applying
a
what
methods aimed
against
trees
methods allow in
spirit
same
It has been
with
coupled
from
We show also
cient
the
based
formal
strategies,
(1981) component analysis
taxa.
explanations
area-cladogram
coding
a
taxon
wide-spread species,
patterns between
we
when several
area-cladograms
cladograms
be
only
to
events.
unique
are
can
be raised
Nelson and Platnick’s
in different
as
outline of
converting
biogeography; (1)
co-evolutionary
method operates in
tions
than
tool. His method and its
resolution than
in historical
general
a
and/or with
analytical
an
as
distributions. As
sensus
be if
might
as
great degree
a
and selected
posteriori
a
factors.
by unique
It is unclear
general area-cladogram
explanation
different
explores
taxa
obtaining improved
mon
caused
met.
compatibility
interpreted
method for
(1981)
In this method he
groups with
and
Would a
and Parenti
area-cladogram.
in which he
at
are
are
with
against
For this purpose, reduced
in which there exists
groups
component
raised
obtains here.
31)
wonders what the conclusions
Recently, Wiley (in press) presented
areas.
incongruencies
relationships
(see Humphries
events.
incongruencies
incongruence.
data matrix
constitute
not
parsimony
one
revealed total
groups
that
minor
example including
At the extreme,
compound
a
implying
example, only
an
incongruence.
for several
used
are
In the sole worked
sequence of speciation
a
area-cladograms
points
to
1983 p.
method
species maps’
factors that af-
to
their
areas,
analogous
(especially by Farris,
‘ancestral
1981)
due
are
consensus
should be the determination
By omitting
established. Criticism
methods
compatibility
Wiley’s (1980,
1986)
be
cannot
areas
reduced
biogeography
between delineated areas.
relationships
other
congruencies
293). However,
under
analysis
a
can
hardly
Therefore,
data matrix, it is suffi-
Assumption
0.
method renders full account of all information
most
resolved
pre-
parsimonious general area-cladograms
possible.
Acknowledgments
Many
thanks
discussions
contributed to
our
CJ. Humphries
tions
to
improve
Haulier.
are
during
due
his
to
Dr. E.O.
tenure
as a
understanding
and
the
two
Wiley,
visiting
who
was
stimulating participant
Netherlands,
of historical biogeography.
anonymous reviewers for
manuscript.
a
scientist in the
We also
We also wish
to
their critical remarks
acknowledge
the
comments
in
and who
made
many
greatly
thank Dr.
and sugges-
by
Dr. C.H.
332
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3
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