[leps-talk] Papilio machaon aliaska - ???

[Barb Beck]

Hi,

I am not trying to defend Eitschberger's conclusions because I do not know
enough about the subject and do not have the original paper but I have a
problem with some of Felix's statements.

I am just trying to follow the logic here with my poor little old female
brain.

Felix states:
    	"The only evidence cited in favor of elevating aliaska is contained in
Eitschberger (1993, Atalanta 24: 15-32), which is a description
	(written in German - which I do read) of egg microsculpture variation
	in the P. machaon complex."

Then he concludes that the studies are flawed because:
	" The obvious conclusion is that eggs from a single female was
	used to characterize each taxon, at least for the three that have the
	same locality and date."

But how is it the OBVIOUS conclusion that because there is the same locality
and date that the eggs are from only a single female?  More than one female
could have been captured at each location on each date.  You have no
evidence one way or the other.  You disagree with the conclusion of the
paper (possibly good reasons) and you SUSPECT that their results MAY be due
to the fact than they only used one female per taxon but it seems to me that
OBVIOUSLY you do not seem to have hard evidence to show that it was the
case.  Have you bothered to contact the author to inquire about this?

Oh well - I'm blond so not supposed to be able to follow logic - there is
lots of "literature" on that.

Barb Beck
Edmonton, Alberta, Canada


-----Original Message-----
From: Felix Sperling [mailto:Felix.Sperling at ualberta.ca]
Sent: March 3, 2002 7:06 AM
To: TILS-leps-talk at yahoogroups.com; Harry Pavulaan
Cc: cguppy at quesnelbc.com; leps-l at lists.yale.edu
Subject: [leps-talk] Papilio machaon aliaska


I haven't had the time to really go over the TILS TC-ISBN list except
for the Papilio machaon group, but from what I see I am seriously
worried about the quality of the decisions that have been made. In
fact this case provides a strong argument for the practice of *not*
splitting species as a default whenever someone says it should be
done.

Splitting "Papilio aliaska" off as a separate species from Papilio
machaon is seriously misguided for two reasons. First, it is
supported by extremely weak evidence, which is accepted uncritically
while Pavulaan appears to dismiss contradicting evidence without
examining it. Second, the reasoning used to support elevation of
aliaska to species status in the  TILS TC-ISBN list is a
contradiction of the principles laid out in the preamble to the list.

The only evidence cited in favor of elevating aliaska is contained in
Eitschberger (1993, Atalanta 24: 15-32), which is a description
(written in German - which I do read) of egg microsculpture variation
in the P. machaon complex. Nonetheless, Pavulaan's notes accompanying
the TILS list state that "Various studies have demonstrated that
machaon is limited to the Palearctic region."  This is a gross
misstatement. One study is not various studies.

Pavulaan also cites McCorkle and Hammond (1988(89)), but this paper
did not address the relationship between aliaska and Eurasian
machaon, and furthermore is based solely on a restatement of old
evidence without presenting any new evidence.

But let us consider the evidence in Eitschberger (1993). Of the 18
small-format pages in this paper, only 3 pages comprise the body of
the text, and there is 1 page of figure legends.  Eitschberger used
scanning electron microscopy to examine the micropyle of eggs from P.
alexanor (not relevant here), and 6 members of the machaon group:
aliaska, saharae, rathjensi, mauretanica, hippocrates and hospiton.
In spite of copious figures showing various eggs, it is obvious that
all mauretanica, hippocrates, and aliaska have the same locality and
collection date. The saharae have 2 dates, and the rathjensi and
hospiton have no date. In all cases there is one locality given per
taxon. The obvious conclusion is that eggs from a single female was
used to characterize each taxon, at least for the three that have the
same locality and date.

Thus Eitschberger's taxonomic conclusions about aliaska rely on the
eggs of a single female butterfly, with no attempt to sample
population variation or geographic variation within that taxon, much
less any of the other taxa that are found in North America. In
Eurasia, the only sampling from outside the Mediterranean is the eggs
from a single female of hippocrates from Japan.  In spite of this
extraordinarily limited sampling, it is evident from the figures that
there is substantive variation within each egg batch (taxon) and that
this amount of variation approaches that of differences between taxa.
At least one of the egg batches (hippocrates) was collected in
6.VIII.1952. There is no information on whether these are eggs
extracted from a pinned female or whether the eggs were preserved
separately since 1952, but in any case there has also been plenty of
time for artifacts in the wax layers of the eggs to have arisen.

Based on this evidence, I concluded that this is an interesting
character system that is well worth pursuing further, but that the
number of independent specimens sampled for this particular study is
simply inadequate to draw any taxonomic conclusions.

The conclusions that Eitschberger draws, based on a single character
system and the eggs from a single female per taxon (at least in most
cases) are that aliaska should be elevated to species status. He also
states that aliaska has fewer teeth in the ridge on the male
genitalia of aliaska versus machaon. He shows the left and right
sides of one specimen of aliaska, and cites Eller (1936) in support
of a contention that aliaska has fewer teeth than machaon. Here
Eitschberger is simply wrong. Seyer (1974, 1976a, 1976a, 1977, [all
Mitteilungen der Entomol. Gesellschaft Basel] and 1982 Dissertation)
exhaustively examined variation in the male genitalia of P. machaon
but is not cited by Eitschberger.  Although he described numerous
subspecies, Seyer also showed that there was extensive overlap
between subspecies in all aspects of the male genitalia. In any case,
Eitschberger makes only vague statements comparing aliaska genitalia
with other taxa. Interestingly enough, Pavulaan's notes cite a "pers.
comm." from Guppy that he has done a survey of the genitalia of the
machaon group and found that "The number of teeth was slightly
(28-30) variable within each taxon, and the taxa aliaska, oregonius,
pikei, polyxenes and zelicaon all had the same range and mean number
of teeth (allowing for the within-taxon variation)."

Eitschberger (1993) presents no evidence whatsoever on the placement
of P. machaon hudsonianus and P. m. pikei. Nonetheless, he states
that both of them should be considered a new formal taxonomic
combinations - P. aliaska hudsonianus and P. aliaska pikei.
Furthermore, he states that hudsonianus was observed ovipositing on
Petasites palmatus by Sperling (1986), which is simply wrong. In
addition, he relies on my report of Muting's (1972) observation of
larval feeding on Artemisia by P. machaon in Afghanistan, without
checking that report himself. In short, the bald statements of
taxonomic reassignment by Eitschberger are either completely
unsupported or severely undersupported by real evidence, and the
paper itself shows other signs of a disinclination to examine
evidence directly.

Despite the fact that only there were only 3 short pages that needed
to be read in Eitschberger (1993), I get the impression that Pavulaan
has not actually read the paper himself either. If he is unable to
read German, then he should admit it and say so in his notes. It is
misleading to do otherwise, just as it is misleading to state that
"various studies" have supported something when it is really a single
study.

The recent, published evidence in favor of the conspecificity of
aliaska and machaon is contained in Sperling (1993  Mem. Ent. Soc.
Canada 165: 233-242) and particularly Sperling and Harrison (1994
Evolution 48: 408-422). In short, this work demonstrates that there
is extensive shared polymorphism in mtDNA between Eurasian P. machaon
(9 specimens and 5 taxa sampled) and North American P. machaon (29
specimens and 5 taxa). In addition, there was virtually no difference
between the European P. machaon and P. machaon oregonius that were
sampled (and later supported with sequencing studies).

However, just as Pavulaan appears not to have read Eitschberger's
work, he gives no evidence that he has actually read my work either.
Instead, he dismisses this work on bizarre legalistic grounds. He
states that "We find it interesting that both McCorkle & Hammond
(1998(99)) and Eitschberger (1993) cited Sperling's work on nearctic
members of the P. machaon complex. Yet, Sperling (1993), Sperling &
Harrison (1994), Sperling & Feeny (1995), Caterino & Sperling (1999)
and Reed & Sperling (1999) continued to treat all Nearctic members of
this complex as "P. machaon", and have cited neither McCorkle &
Hammond (1988(89)) nor Eitschberger (1993)."

Considering that Eitschberger's work was published in 1993, how
exactly was I supposed to have seen his work and cited it in my 1993
or 1994 paper? For my 1994 work (which is the only paper where it
might have been appropriate to cite Eitschberger), it is stated
plainly on the first page that it was submitted  in July 1992 and
accepted April 1993. The 1995 work concerned only foodplant records,
while the later work was published in molecular evolution journals
and dealt with molecular evolution issues that precluded any
diversion into taxonomy even if I had wanted to.  Furthermore, why
would I be obligated to cite an opinion piece by McCorkle and Hammond
that presented absolutely no new evidence, and in fact simply
restated opinions that were already widely available elsewhere? This
certainly wasn't a case of actively ignoring evidence on my part, or
even dissenting opinions. It was a case of these papers not having
been relevant to the theme and format of the papers I was writing. In
fact, I wasn't even aware of Eitschberger's (1993) paper until about
1996 or 1997, when he sent me a copy of the work. The journal
Atalanta is not subscribed to by any of the online databases or the
library I was using at the time (UC Berkeley - otherwise one of the
best entomology libraries in the U.S.).

In any case, the way in which Pavulaan has phrased his "notes" leaves
the reader with the impression that I was doing something shady by
not citing these other paper. I find this completely inappropriate.

However, this very strange line of reasoning has allowed Pavulaan to
avoid actually considering the contents of my papers. This brings me
to my second line of argument for why Pavulaan's treatment of aliaska
is so seriously misguided. He has quite simply and openly flaunted
his own guidelines in setting up this list. Even though he hides
behind a royal "we" in the note on machaon, it is obvious from an
earlier posting by Ron Gatrelle that Pavulaan is the one who has
compiled this list. Considering that, it is a really pity that he has
not shown the courage to admit to that fact, but instead says in a
recent posting that "Thus, I may add a disclaimer to the TC-ISBN:
"The taxonomic arrangements presented in the Index represent the
published work of authors and NOT the personal VIEW of the Index
editor", although in rare cases it may appear that the editor is
taking sides.  Remember, the editor may personally disagree with the
arrangement being used in the index.  I hope nobody takes this
personally."  If Mr. Pavulaan didn't write those notes, then who did?
Stating that no one should take offense and the editor is just a
compiler anyway just doesn't jibe with the facts. It does show a lack
of ability to owe up to what has been written.

Whoever the author of the notes is, there was a clear judgement made
to ignore contemporaneously published evidence (mine) which presented
better sampling and clearer characters (mtDNA restriction sites) that
contrasted with that of Eitschberger. The preamble to the TILS list
says that "The purpose of this list is not to compete with, or claim
legitimacy over, any other established or proposed lists, but to
present an objective treatment of scientific butterfly names. In
fact, it is not called a "list". It is intended to be an index of
available names for anyone interested in obtaining an "unfiltered"
record of described taxa, and for researchers and authors intending
to construct lists of their own. The TC-ISBN follows a strict set of
guidelines and chronological progression as much as possible. The
guiding premise is that all listed taxa are presented accurately with
respect for the work of researchers chronological time frame. Thus,
each taxa is first viewed as it was originally described, then
reexamined in light of more recent research which compels changes in
the nomenclature. In many cases, there was never any actual published
work to justified a change in usage (authors or listers simply made
the changes). This may mean that some taxa, currently treated
generally as subspecies of one or another species, may be re-elevated
to the status of their original description. If the status of a
described taxon is changed or revised by means of a published study,
that change is reflected in the index. With every published change or
revision, the most recent one is reflected, and it is assumed that
researchers will use every available tool and published source upon
which to refine our knowledge of butterfly taxonomy. The editor of
the TC-ISBN will not make any changes based on unpublished "expert
opinion" or "common knowledge". The bottom line is that no changes
are allowed to the index unless published in a scientific journal,
bulletin or other forum intended as a monograph or similar-level
treatment."

Instead of taking my 1994 paper (which is the key paper) as
contemporaneous, Pavulaan has chosen to fault me for not citing
Eitschberger and McCorkle and Hammond, thereby providing an excuse to
not even discuss the contents of my paper. Instead of consulting me
about the chronology of the work, Pavulaan has chosen to consult
instead solely with a variety of others who have openly stated that
they have differing taxonomic interpretations. Instead of following
his own principles of considering only published evidence in regular
journals, Pavulaan has instead used "pers comm." sources (e.g.
Guppy's remark that he was overly conservative in considering aliaska
to be conspecific with machaon in his book). Instead of weighing the
evidence of Guppy's "pers. comm." that there are no genitalic
differences between aliaska and machaon in the context of the fact
that this is an independent assessment of Eitschberger's evidence in
general, Pavulaan has instead chosen to assume that egg micropyles
are somehow more worthy of taxonomic consideration. Instead of
considering that the egg micropyle evidence cited in favor of the
phoebus/smintheus split was part of a larger package with more
sampling, Pavulaan has simply accepted the reasoning that anyone who
accepts a split between phoebus and smintheus must accept a split
between machaon and aliaska.

So what the point of all this anyway? Whatever it is, it does not
inspire confidence in the TILS TC-ISBN list.

Felix Sperling








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