Papilio machaon aliaska

Cris Guppy cguppy at
Mon Mar 4 22:50:19 EST 2002

Papilio machaon aliaskaSince I was foolish enough to provide "pers. comm.'s" to Harry, and agree to their use in the TILS TC-ISBN list, I am obliged to wade in at this point. Despite getting older every year, I still need to learn to when to keep my mouth shut.:)

(1) Eitschberger's paper provides a body of evidence for deciding that machaon and aliaska are different species.
(2) Felix's 1993 and 1994 papers provide a strong body of evidence that machaon and aliaska are the same species.
(3) I agree with Felix that Eitschberger's paper is weak, due to a the lack of multiple-female replication.
(4) I agree with Felix that his papers are more rigorous and the data has appropriate replication.
(5) I was in error to state to Harry that I was overly "conservative" in not splitting aliaska and machon in the the BC butterfly book. In fact the evidence was not conclusive, especially considering the contary evidence published by Felix, and leaving aliaska as a subspecies of machaon was appropriate given available evidence.
(6) I do not consider the total available evidence to be conclusive as to the appropriate groupings of species and subspecies in the machaon group. The contractictory evidence still needs resolution.
(7) Eitschberger's and Felix's publications should be considered to be simultaneously published for the purpose of the list, because neither author had the opportunity to review the other's work prior to publishing their own work.
(8) How the TILS TC-ISBN list addresses "simultaneous" revisions needs to be determined by Harry and Ron. The "most recent" rule will not work in such a circumstance. The only solution I can think of is to choose one revision as "the best" and use it with a clear statement that the choice was based on the compiler's opinion, and then provide the alternative interpretation.
(9) I suggest the TILS TC-ISBN purge all "pers. comm." comments. They are very tempting, but also very dangerous to the consistency of the list and adherence to the rule of "published evidence".

Cris Guppy

  ----- Original Message ----- 
  From: Felix Sperling 
  To: TILS-leps-talk at ; Harry Pavulaan 
  Cc: cguppy at ; leps-l at 
  Sent: Sunday, March 03, 2002 6:05 AM
  Subject: 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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