Papilio machaon aliaska

Felix Sperling Felix.Sperling at
Sun Mar 3 09:05:54 EST 2002

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 

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 

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 

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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