Contributions
Zoology,
to
SPB Academic
Key words: phylogeny,
Russian
Street
Academy of Sciences, Profsoyuznaya
cladogram, ghost
fossil record,
range,
Abstract
to
of the
of
calculation
range
between
a
on
cladogram
hymenopterous
is
proposed
to
assess
and the fossil record and
this basis. The method is tested
insects
to
reveal
the
(Order Vespida),
as
and the results
but
perfect,
is
is
Contents
so are
all
and
both
why
of
251
thy of comparison
252
discussion
Conclusion
257
Acknowledgements
258
References
258
There
are
the
Introduction
Benton
and
metrics
do
extent
ward
research that
ordinary condition in cladistic
cladograms developed from different data for the
set of taxa often differ
significantly
other. This is crucial since the
one
number of indices that have been
statistical
cladogram
seems
the
1996: 3.2.5).
It
is
reasonable to seek
on
on
principle
our
it may
im-
of evidence. This
sources
well, for the
character set
That
always hypothetical.
informa-
evolutionary
cladograms,
but
‘perfection’,
in
com-
even
so
possess
are
wor-
testing phylogenies.
proposed
a
record based
of
respective
Hitchin
not
to
quality
assess
fit the
cladograms
end,
1
am
reverse
of
a
taxa
(reviewed
agrees with
task, that is,
the
taxon
In its
to
test
for their
record.
fossil
proposing here
by
these
a
To-
similar but
present form the
rough, but whenever found
it may be
propriate,
cise
rather
it
particular
slightly different approach.
method is
how
1997). Unfortunately,
of congruence with
this
on
easily improved for
more
appre-
calculation.
There
developed
devia-
Material
parsimonious
not
process
parsimony principle.
less dependent
one.
features that reflect
from the most
however, that the evolutionary
observe
from each
cladogram compari-
is aimed at selecting the most correct
(Quicke,
in
the fossil record is
of the
methods
cladogram
different
tion of the
evolutionary
254
of the fossil
to estimate
so
record and the
Material
Methods
are
sources
only degrees
are a
and
our
as
polarity
251
son
of the
strictly dependent
and transformation series
exhaustive,
never
Introduction
same
source
course,
for cladistics
true
tion, the fossil
an
Russia
Moscow,
external standard in the comparison
an
cladograms. Of
position
It is
117647
on a
is
and
Hague
phylogeny
discussed.
Results
a
not
evolutionary hypotheses,
be used
of
cladograms developed recently
of the
are
ghost
cladograms
compare
set
123,
The fossil record is
congruence
(2000)
The
Vespida, Hymenoptera
information that is
the
251-258
Rasnitsyn
Paleontological Institute,
A method
(4)
test
fossil record: the ghost
range
Testing cladograms by
Alexandr+P.
69
Publishing bv,
so
evident,
does
That
is
strictly
why
1
it
method of assessment
particular evolutionary concepts.
Fossil record of the
Vespida
=
Hymenoptera)
method for the
any fossil
enough
to
hymenopterous insects (Order
following
is
selected
reasons.
to
test
the
Incomplete like
history, the hymenopteran record is rich
represent
as
many
as
51 out of 54
ex-
252
A.P.
tant families
is
(e.g.. Fig. 1; superfamily Chalcidoidea
used here
as
for it
cladogram available
these families
sozoic,
netic
A
be
to
to
The
test.
1
Fig.
been
have
examples
is based
on
to select
Methods
Mesozoic).
When
re-
tree,
and modifications
the
2 shows the results
caption. Fig.
clade
Rasnitsyn (1988), with several
minor additions
Roy,
and
Ras-
Lindgren
(1999).
cryptic
in
as
the
fossil
the
molecular
apocritan
phylogeny
performed by Dowton, Austin, Dillon and
sky (1997:
fig. 2),
Orussidae and
of Dowton
of taxa
by
and Austin
line
I.
Fig.
lines
Relation
(more than
ranges
N -
Paleogene,
the Cenozoic,
P
thick
-
duration
of the
by
amber
Cretaceous
1986:
fig.
66
F;
Platygastridae
my
in
-
Lower
Eocene,
amber
in
and
in
the
New
Jersey
East
Siberia
Guo
et
-
P
Lower
of
Ji
as
al., 2000),
amber
Cretaceous
Lower
a
of
genus
in
the
cladogram
to
New
of
et
ah,
of Brazil
it
Miocene, N,
and
are
in
the
(Darling
outlined
visually.
These
expressed
lines
the
on
-
in
T -
are
Lower
it
Cretaceous
New
,
Known
in
the
in
of
lowermost
from
J
their aver-
or
Rasnitsyn,
amber
-
Upper
amber of
Australia
Dominican
amber
amber
in the
in
the
amber (old
1995;
my
disputably Upper
(Jell
by
and
Duncan
identification),
D.A.
Grimaldi),
Bethylonymidae
in
(my identification),
Upper
Eocene
Upper Cretaceous
Formicidae in
Diapriidae
“orphan” families
by
a
and
for some taxa
ah,
et
P
Upper (in
-
Baltic
(my identification), Rhopalosomatidae
that sometimes
marked
Cretaceous,
-
Baltic
(Ren
communication
and the rest of Platygastroidea,
are
K
Thick
long ghost
-
Middle,
Eocene
of China
1988).
lines
modified
are
ranges
Koonwarra,
(my identification),
by boundary lines, except
thin
Cretaceous of Mongolia (my
Miocene
Jersey
How-
geochronologi-
Jurassic,
-
the Burmese
Masner, personal
the
ghost
their total dura-
as
Lower,
-
the
Sharkey 1990), Sierolomorphidae
Lebanese
use
and evalu-
calculating
Spain (my identification), Diapriidae
in
in
is
to try and
comparison
double
Triassic,
Cretaceous
Cretaceous
L.
by
of inference for
a reason
Mongolia (Rasnitsyn 1995b), Scolebythidae
and
displayed
cladogram,
ranges,
indices:
found
Evaniidae
in the
well
cladogram fits the
duration of the
ghost
-
Pliocene).
the
one
extensively debatable issue, full
an
Periods:
Sapygidae
Lower
of Alava,
Upper
as
parsimonious
degree
This is
might suggest
by subscript
Megalodontesidae),
and
lower
Chalcidoidea) (modified
record).
shown
1998), Sclerogibbidae
in the
than
set of taxa,
competing cladograms.
we
thin
Jersey (identified by
in the Lower Cretaceous
simplify
-
are
Chalcidoidea(new family)
superfamily (Karatavitidae and Ephialtitidae, Serphitidae
separated
fossil
Argidae
Monomachidae
of Transbaicalia
1999). Superfamilies
(
were
more
ranges
more
for
range concept
the absolute
for
vertical
1988), Megalodontesidae
Plumariidae
Cretaceous
and Falsiformicidae
N
taxa.
cal units is still
(superfamily
(Aptian) amber
England (Rasnitsyn
Janzen, 1999),
(Dlussky
et
sources,
as
ever,
Ipntering
Oligocene,
a
These
termed
age duration per node per terminal branch.
relationship,
before
following
Scelionidae and
the Lower Cretaceous
(Brothers
and Tiphiidae
on
Upper Cretaceous
Cretaceous
Embolemidae
Ren,
(Bashibuyuk
the
tion
noth-
families
lines -
thin
the
a
apparent that the smaller
the better the
e.,
that
1-6).
configuration
in million years, either
ranges
contemporary (Holocene). Epochs
-
identification),
Mymarommatidae in the
the
hymenopteran
1886, ignored by Rasnitsyn
Jibaissodes
identification of
Thus
ranges
means
geochronological epochs
Paleocene, P,
Brischke
ones
horizontal
longevity,
comparing Rasnitsyn (1988) basing
data
dendrograms
implied ghost
ghost
ation of
lines, relationships
two horizontal
two
the
lines; the vertical thin
thin
Neogene, R
:
All
(1994).
vertical
and
show known
the duration of
style, with the known duration
being indicated by
connecting
Bartow-
families,
symphytan
i.
record,
that respect due to
added after the work
horizontal thin lines, and the
(see below) by
-
the
Stephanidae
unified in their
are
with
Figs.
particular
ghost
trees. It is
respective
particular
a
infer
to
fossil record would
in
a
their
the extension of
these extensions,
represents
have
we
thin lines
differ in
they usually
3-5
Fig. 6
11), respectively.
paleontologi-
intervals of existence
and
fig.
cases
earlier than the
(1997: fig. 2), Whitfield (1992: fig. 5), and Brothers
Carpenter (1993:
phylogenetic
a
to the
length
cladogram is developed for
in Vilhelmsen
cladograms
re-drawn from the
are
Figs.
cladogram into
‘ghost ranges’ (Norell, 1992). When
nitsyn (1988) performed by Ronquist, Rasnitsyn,
Eriksson
some
(vertical
inferred
recent
parsimony re-consideration of the concept by
In
appears
suggest
in
explained
of the
a
fit the intemode
we
subclade.
1-6.
Figs.
we convert
cally confirmed minimum duration of
enough of them
in
hori-
a
are
proposed
displayed
are
connection of two parts of
a
Me-
phyloge-
family level phylogeny of the order,
us
than
more
zontal line.
no
families
more
since the
(mostly
variety allows
and this
from the
high-level
ing
Testing cladograms by fossil record
-
38 of
as
many
starting
and 20
cladograms
the
cently for
recorded
extinct
wealth of
As
yet).
wherein many
occurred,
events
known
are
time
a
taxon because there is
single
a
Rasnitsyn
broken
members
and other
arrow.
amber
of
the Lower Cretaceous of
of
one
and the
same
Proctotrupoidea) appear
Contributions
to
Zoology,
69
(4)
-
2000
253
A.P.
254
of gaps and traps for
a
special efforts
not
the matter of issue is
fine
than
seem
tuning
sonable
here is
the
the
when
cases
inferred to
for
appearance
unit. For
the
originate
to
than
interval
all
a
first
should
grid
ghost ranges
displayed
as
ranges
ranges absent in
figs
2-6 indicate the
Fig.
in
Pompilidae and Sapygidae
in
Fig. 1, while in the
case
may be
opposite
molecular
(1993)
are
ranges,
more
these two
when
tree based
ranges,
congruent
the
cladograms in regards
long ghost
terminal
concern
Fig.
ranges.
and
taxa
to
1
the
seven
taxa and
of which
cladograms
with
the fossil
three families
lacking
ghost
that
ranges
fossil record
others (Blasticotomidae,
Argidae
and
+
Pergidae,
Sapygidae)
Late Eocene
or
later.
(e.
as
tested,
Furthermore,
taxa.
inclusions in the Baltic
cases
whenever
I shall
duces
a
tional
tree with
only
long
Relation
Dashed
double
75 terminal
be
et al.
of the
hymenopteran
have
taxa and
taxa
ranges
as
eight.
to
that ancestors
cladogram is
can
we
a
age.
infer
in
et
So when
it
to
the
we
have
longer ghost
In
fact,
al.
Fig. 1.
in
never
be iden-
group rela-
cladograms. Unlike
take
an
groups always
a
daughter for
earlier
origin and
range. I have discussed at
reasons
why
it
is
ancestors
necessary
(Rasnitsyn,
Ronquist’s cladogram (Fig. 2),
long ghost ranges (cases of Cimbicidae, Mega-
lodontesidae,
by Ronquist
not
presence of
sister
Therefore, only
available for these
acknowledge phylogenetic
four
Argidae,
seen
equal
1996).
pro-
to those
are
length elsewhere
“long”
calculated
my
(those lacking autapomorphic char-
such.
therefore
eight addi-
additional
as
sister,
a
some
however.
mother-and-daughter taxa, the sister
the
amber)
(Fig. 2)
of the considered
The results deserve
gained by
score
own
ranges
states). The other cladograms follow the strict
tionships
found in
discuss the
ghost ranges (Cimbicidae,
line - long ghost
tified
+
they consider these
cladogram from Ronquist
2.
may
in
sections
compared.
cladistic concept
Pompilidae,
record
ranges that appear in addition to these
ghost
Fig.
g.,
the fossil
acter
five
Electrotomidae
Sclerogibbidae,
enter
These individual
all other trees
The
that
clade
high
ancestral taxa
(Pergi-
dae, Xiphydriidae and Bradynobaenidae),
are
record
cladograms (each within its
peculiar. My cladogram acknowledges
the tree with 76
depicts
eight long
The
of the
occurrence
the
follows from the observation
additional discussion,
1-6 show considerable differences between
Figs.
on
1.
indicate that the
corresponding
cladograms
ranges
from Dowton with
scope) infer the minimum number of ghost
Results and discussion
a tree
(1988) and Brothers and Carpenter
than the others. This
that
1.
Bradynobaenidae
cladograms
Fig.
long
longer here than
are
of the
tree
Fig.
produces
(Fig. 6) considers 24 terminal
by Rasnitsyn
long ghost
in
seen
long ghost
true). The
The above results
are
Whitfield’s
(the long ghost
1
of
do not repeat those in
1.
Fig.
same as
ter-
Carpenter (Fig.
It
wasps.
and four
taxa
produces eight long ghost
double thin lines. The dashed double
lines that appears in
only aculeate
the
co-authors
shown there
are
(13
long ghost
superfamilies; his
from Brothers and
19 terminal
composite
in
mostly
with
ranges, all additional to those
being
the
geochronological
by thin vertical lines, while long ghost
represented by
deals
concerns
with
the definition of
two
of the time
1-6). Thus, all
5)
particular time
a
the time
range,
epochs (two steps
or
to its
prior
satisfy
longer than,
or
terminal taxon
a
more
present,
ghost
“long”
equal to,
Figs.
ghost
means
additional
two
4) also includes 13 terminal taxa and six
(Fig.
of
sum
This
yields
(Cimbicidae and Cephidae).
cladogram
rea-
and
The cladogram
simple metrics applied
subclade is
the
after
even
so
‘long ghost ranges’ per cladogram.
fossil
ranges
complication.
counting
be
more
only the lower hymenopterans
minal taxa)
phylogeny.
particular
a
when
1. Vilhelmsen’s cladogram (Fig. 3)
Fig.
considers
sophistication
now,
in
seen
the method itself
testing of
a
plest form, it will be the
the
metshidae, Ceraphronidae, and Apidae) above those
preceded by
necessary
Testing cladograms by fossil record
-
Stephanidae, Ichneumonoidea, Trigonalidae, Mai-
if the method is found workable in its sim-
Indeed,
The
this
However,
taxa.
approach does
rather
avoid
to evaluate the absolute time ranges
of the involved
of
To
user.
non-expert
these traps, the calculation should be
Rasnitsyn
Ceraphronidae, and Apidae) appear
(1999: figs. 2,
5
and
9, combined), formatted after Fig.
I.
Contributions
to
Zoology,
69
(4)
-
2000
255
256
A.P.
Fig.
3.
Relation
of the
symphytan
Fig.
4.
Relation
of the
parasitic
Fig.
5.
Relation
of the
aculeate
taxa
taxa
taxa
as
as
as
calculated
Rasnitsyn
by Vilhelmsen (1997: fig. 2),
presented by Whitfield (1992: fig. 5),
calculated
by
Brothers
and
-
Testing cladograms by fossil record
formatted
formatted
after
after Figs.
Carpenter (1993: fig. 11),
Figs.
1,
1,
2
2.
formatted
after
Figs.
1,2,
Contributions
Fig.
Molecular phylogeny
6.
Orussidae
Stephanidae,
because
solely
not
and
daughter
69
Zoology,
to
they
of
and
(4)
-
2000
hymenopteran
Dowton
by
considered
are
et
to
257
taxa
al.
as
calculated
(1997: fig. 2)
be sister and
of Tenthredinidae,
groups
is true for three
dontesidae and
dogram (Fig. 3);
daughter
taxon
cases
Cephidae)
in my
of
cladogram
directly
from
Of six
Scpulcidae.
are
long
(Fig. 4), five
because
phronoidea (s.str.)
Vilhelmsen’s
Cynipoidea
from
-
ghost
and Cerato descend
Megalyridae, Platygastroidea
cidoidea and
the
absent
are
Trigonalidae
suggested there
for
(1994; fig. 3),
formatted
after
Figs.
1,
symphytan families,
2.
Conclusion
the
+
The
above
Chal-
phylogenetic
allow
for
the
set
same
to compare and evaluate
the results of application
standable and
That the
the
explain,
record
than
the
on
sister groups, these latter being the
additional
(Fig. 6),
cladogram
long ghost
ranges
Four of them result from the
In Dowton
there
comparing
same
are
to
causes:
seven
Fig.
in
1.
my
cladogram, Tenthredinoidea originate directly from
Xyclidae, Cephidae
from
from
Sepuicidae, Trigonalidae
Megalyridae, and Evaniidae
+
Gasteruptiidae
from Praeaulacidae.
The
reasons
pear in
in the
why
Figs. 2, 3,
Fig.
1,
are
relevant to the
so
evident.
phylogeny
than to methodology of
will
long ghost
ranges
of
Possibly they
Hymenoptera
ap-
seen
are
rather
phylogenetics, and so they
be considered elsewhere.
groups
At the
cladogram by Brothers
cates
that the
(1988)
high
straightforward
from
result
i.e., the making of
the
automatic
preciation
Several
range
one
and
ways
simple
fits better
of others,
the
only legitimate
indi-
by Rasnitsyn’s
the present
is not
test
method
a
used there,
cladogram by the method of
pencil” approach) instead of
an
ap-
ancestor and descendant taxa.
to
further
may
of
to
Carpenter (1993)
of the
of
appreciation
time, testing of the
received
be
development of
proposed.
is the calculation of the
count
under-
are
parsimony calculation, and
of the
method
a
same
and
score
cladogram
Flennig (“paper
other
4 and 6 in addition to those
not
in cladistics.
ones
Moreover,
in additional
monoidea + Aculeata from
daughter
does
at least in part.
majority
the
partially
least
at
mother and
Ephialtitidae.
taxa
cladograms
trees.
proctotrupoid family Mesoserphidae, and Ichneu-
and Austin’s
the
total
different
by
of
of the method
to
the
cladogram from Rasnitsyn (1988)
fossil
depends
possible
that
displayed
ranges
trees
forming the background of these
to
ancestral
us
indicate
observations
of ghost
amount
cla-
cladogram Cephidae is
ranges in Whitfield’s tree
in my
The
(Cimbicidae, Megaloin
and Austin
apocritans,
Praesirici-
dae, Megaspilidae, and Sphecidae, respectively.
same
Dowton
by
for the
ghost
The
the
most
ghost
evident
length rather than the
ranges:
this would make the
258
A.P.
method
As
possibility.
there is
precise. However,
more
the
proposed,
another
yet
method
range
ghost
Rasnitsyn
Molecular
designed
to
in
competing cladograms
test
their congruence with the fossil
record. Instead,
it
might be found useful
geochronologi-
cal
data
in
directly
to
the
the
use
Duncan
PA,
parsimony calculation,
sil
the
it its
even
Norrel
Ass.
for
test
ghost
method
range
of
effect
a
Quicke
Acknowledgements
D.
D,
1996.
Lu
Guo
L,
Zoological Institute, Uppsala
versity, Uppsala, Sweden, made it possible
March 4-7,
was
where
1998)
presented.
the
Dmitry
Dr.
E,
Institute, Russian Academy
to the
attention
am
thankful to
State
were
Ji
Sh.
and to
for
useful
the
anonymous
The work
me.
attracted
Hitchin
my
(1997).
I
reviewers whose advises
supported
was
Trust to D.
from The Leverhulme
grant F58AQ
Paleontological
Zoological Museum, Moscow
Pavlinov ofthe
Igor
(Uppsala,
in
part by
Quicke
a
and M.
Fitton.
Moscow:
pterous
AP.
Rasnitsyn
archaic
Bull.
Benton
MJ,
Nat.
llitchin
R.
(Lond.) (B)
Brischke
D.
1886.
Congruence
the
of
53-58.
phylo-
of life. Proc.
885-890.
des
N.
Danzig,
JM.
F.
6:
1993.
Bernsteins. Schr.
278-279.
Phytogeny
of Aculeata:
Hym.
Res.
227-304.
Darling
Bull.
Dlussky
D
Sharkey
Amer.
Mus.
Nat.
GM.
1999,
The
menoptera)
in
the
MJ.
Hist.
first
Lower
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