Papilio bjorkae (Pavulaan, 2024) Tiger Swallowtail

[Chuck]

Yesterday Harry Pavulaan published a new species description of the New England Tiger Swallowtail Papilio bjorkae

https://doi.org/10.5281/zenodo.13952895

This is the "Spring Form" of the eastern-US population Tiger Swallowtail (though Harry doesn't go so far as to say ALL Spring Form are bjorkae, he focuses on those in New England.)

To summarize, I excerpt the below, bold added:

What is known is that in Rhode Island there are two distinct phenotypes in early spring.
One is clearly the spring phenotype of southeastern, bivoltine Pterourus glaucus glaucus (Figs.
12-17); the other is a univoltine taxon bearing a striking resemblance to Pterourus canadensis
(Figs. 18-21).

Following my past five years of study exclusively the Mid Summer Tiger Swallowtail (MST), my next project was to tackle Spring Form. It appears Harry beat me to it!

That said, while oft repeated and indeed repeated in the bjorkae description, I have found zero evidence genetically confirming that there is a spring flight of Papilio glaucus; there is indeed at least one, or as Harry said, two, phenotypes, but nowhere has this Spring Form been subjected to genetic confirmation that any of them are glaucus.

Given that the eastern (USA and Canada) Tiger Swallowtails appear to be a complex complex (pun intended, I suppose) we simply don't yet have a comprehensive picture of what's what. There's still a lot of research to be done.

Still, at some point somebody has to pull the trigger, even with incomplete data, and Harry Pavulaan finally did it. Notably, he did likewise in breaking up Hemileuca maia in NY & Ontario, and immediately the new taxa were ESA listed- none too soon.

Harry does delve into the Mid Summer Tiger (MST) without naming it as a new taxon. We're working on that now. Interestingly, Harry says of MST "s differentiated from glaucus primarily by the females (Figs. 29-30, 40); the males (Figs. 31-32, 39) are somewhat difficult to differentiate from glaucus." It's interesting because a similar claim was made by Chris Schmidt. Yet I can find the males, not the females, to be more readily differentiated; I can do it with 100% certainty. So it's interesting the observations by differing experts. Of course, perhaps, I have the benefit of having on-hand 174 set specimens of MST, another 30 something on ice, and having sent over 30 to Julian at UK. Plus I have the benefit of travel to glaucus territory regularly.

Now, for me, the question is if it's worth investing more time into Spring Form. To my mind, it's a complex of speciation, hybridization, etc. It's unlikely (I believe) that a new taxon beyond bjorkae will be revealed- or is it? I struggle to think that the Spring Form in JHyatt's VA is the same as the Spring Form in my town, which I know isn't the same as the Spring Form an hour north of here.

Well, anyway, we have Harry's description of Spring Form to digest and build on.

[kevinkk]

apparently this is where collectors are separated from the entomologists. I don't understand how a "spring form" can be a different species.
Or am I misunderstanding the issue?
It seems like the Actias luna form issue, I see the spring form listed sometimes as a different species, rubio...something.

[Trehopr1]

I agree with you kevinkk.
Way too much "over-analyzing" of some well established species.
There's nothing wrong with taking a closer look at some things as Vernon has proven that a couple of Catocala species (established long ago) were in fact 2 different species (4 species in total) which had not been looked at close enough....

However, I think that if you look and you look and you consider the "lines still blurred" then you don't have something that is a distinct species.

I will only take the word of an established, specialized researcher like Adam for true definitive determinations.

[eurytides]

The term “spring form” is misleading. That’s the term we used when we didn’t know what it was and it was meant to convey that it’s the spring phenotype of glaucus. Now Harry is saying that this “spring form” is actually a species on its own, but it’s not a “form” of anything we have previously described. I am also hesitant to take Harry’s word here without genetic data. It’s 2024, and despite the ICZN, I personally don’t think you can claim a new species without genetic data. The Code doesn’t require it, but the Code seems like it needs a modern update.

[adamcotton]

The Code doesn’t require it, but the Code seems like it needs a modern update.

The 5th edition of the Code has been worked on for several years now, and once the draft is ready I understand it will be publicly posted for 1 year in order to receive feedback before finalising for publication. I think it will be at least another year before the first draft is ready for scrutiny.

Adam.

[Trehopr1]

Very well put eurytides !

I am in complete agreement with your thoughts.

[adamcotton]

Note that the neotypes designated in Pavulaan & Wright (2002) for the following 4 names:
Papilio glaucus Linnaeus, 1758, Papilio antilochus Linnaeus, 1758, Papilio turnus Linnaeus, 1771 and Papilio alcidamas Cramer, 1775
are all invalid under the ICZN Code since they did not fulfil the requirements of Article 75,
particularly 75.3.4:

the author's reasons for believing the name -bearing type specimen(s) (i.e. holotype, or
lectotype, or all syntypes, or prior neotype) to be lost or destroyed, and the steps that had been
taken to trace it or them;

At least one of these, Papilio turnus Linnaeus, 1771, still has two extant original type specimens, one designated as lectotype by Honey & Scoble (2001).

Two of these non-neotypes are also illustrated in Pavulaan (2024).

Adam.

[Trehopr1]

So, if I might ask Adam ---- why are these "scientific" papers produced without adequate peer review ? Why are they the least bit meaningful if they're clearly are oversights and inadequate scientific processes having been taken ?

I suppose Vernon could really give us the full effect on this thought.
Although, he has already mentioned considerable issues with many papers that he has seen produced !

[adamcotton]

So, if I might ask Adam ---- why are these "scientific" papers produced without adequate peer review ? Why are they the least bit meaningful if they're clearly are oversights and inadequate scientific processes having been taken ?

I cannot say whether these papers have been peer reviewed or not. One problem I have noticed regularly in scientific papers (I would regard Pavulaan's papers definitely qualify as such), even by major research institutions, that there are very often errors in the work. I would include papers that I have co-authored among these. One problem of peer review is that the reviewers and journal editors themselves may not be aware of certain errors, especially in the case of DNA papers, where reviewers are often experts on DNA analysis methods but know very little about the ICZN Code, for example, or even the taxa in the manuscript they are reviewing.

I reviewed one paper a couple of years ago but specifically declined to review part of the paper on a subject that I was not familiar with, and informed the journal editor accordingly. Hopefully the editor asked someone studying that topic to compensate for my lack of knowledge.

Adam.

[adamcotton]

I should add that the description of Pterourus bjorkae is definitely a good thing. It gives a name to a particular entity (represented by the holotype) and will encourage further more detailed studies, including DNA analysis to find out whether or not this, MST etc are valid species or not. I am sure that this study has already been going on for quite some time, as evidenced by Chuck's thread on Five Finger Lakes glaucus.

Adam.

[Trehopr1]

Thank you Adam for your considerate thoughts on this topic.

I never considered peer review as being quite so complicated but, you certainly evidenced some very valid points regarding its limitations.

[Chuck]

Way too much "over-analyzing" of some well established species.

Such as Papilio maraho?

What is "well established"? If it means "conventional wisdom" or "as currently accepted" well that's the death knell of science.

That's why when I started this I had only about a dozen Papilio glaucus; and so did the Iveson collection, and another local private collection. They were "just Papilio glaucus"- well known, common- so who would need more specimens? Why would Hagen, Lederhouse & Scriber bother to look into Tiger Swallowtails just to describe a smaller form from Canada?

Kevin:
where collectors are separated from the entomologists.

We are all entomologists.

The geneticists are amongst the most important part of modern entomology, yet as Adam said, they don't understand the subject matter. That does not belittle their contribution.

Collectors, even bauble collectors, contribute to the understanding of entomology.

I don't understand how a "spring form" can be a different species....It seems like the Actias luna form issue, I see the spring form listed sometimes as a different species, rubio

Eurytides explained it pretty well.

The take-away from the 1985 Hagen & Lederhouse paper was that there are two flights and the Spring Form makes next year's Spring Form, and the Late Flight makes next year's Late Flight. Contrast that to Eurytides marcellus which any flight can appear next year as any of the three flights. And Samia cecropia which in some areas has two flights, but same thing.

In the case of Hagen's Ithaca-area Tiger Swallowtails you had an interesting event- two groups that looks similar but occupy the same area at different times of the year, and don't inter-mix. Now, we understand (I prefer "believe") that species evolve via genetic changes at a predictable rate. Given the first sentence, that means that those two groups either have, or definitely will, begin to separate (speciate.) The common dragonfly Anax junius is on this same path.

Then we have the case of appalachiensis and MST, for which the parental species split, then recombined a long time ago; ie a recombinant hybrid that has little interface with the parental species, for a long time.

I have long been a "lumper" and not a "splitter." Mainly because so many of the splitter people described new taxa simply to see their names. And, for the most part, Lepidoptery has pushed back from splitting. The aquarium fish people don't - they assign locations to, for example, the "common" Angelfish of the Amazon. They don't give a damn if it's scientifically named, they assign interest and value to naturally occurring varieties- they track the complex themselves. Now, modern genetics of butterflies has shown us that the taxonomy of most all (eg., Tiger Swallowtails) isn't flat like a list, it's branched with differing lengths of branches, like a tree. There are currently (even excluding MST) two big branches under "Papilio glaucus" meaning there are two related groups with a common, extinct ancestor but they are definitely separate. Separate enough to make a bauble collector track down and buy one of each? Probably not. Separate enough to wonder WTF is going on there? You decide.

[Chuck]

On the topic of over-analyzing "well established" species:

It is well established that Danaus gilippus has two subspecies in USA, bernice in the east, and thersippus in the west.

But apparently Edward Pfeiler didn't buy into conventional wisdom, and in his 2024 paper in LepSoc shared his study, which includes genetics.

There are indeed two groups of gilippus, genetically separated by a single significant mutation. Yay, some might jump to say, well established is right! But the COI barcoding of these two groups does not map to the subspecies. Instead, both groups are found, apparently inter-mingled, in both the western and eastern populations. So there goes the two subspecies by region; maybe there are two subspecies (author says no) but they don't map to separated geographical distribution.

Who'd have thought? Such a well established, common species now devoid of subspecies. From Texas to the tip of Florida, it's the same species.

Wait! will say the geneticists. Not so fast. Because, as I've been told a thousand times, COI barcoding only tells PART of the story. Ugh. Now the genetics dudes also use SNP, which doesn't always line up nicely with COI. For God's sake, can it get any more confusing? I suppose it can, because the genetics guys also talk about other genetic markers that mean nothing to me. So maybe there are two subspecies of gilippus, east and west? That will take more study beyond well established.

[Chuck]

why are these "scientific" papers produced without adequate peer review ?

I suppose Vernon could really give us the full effect on this thought.

I would too like to hear Vernon's thoughts on this. I suspect I know the answer.

Pre-publication reviews are always great; even neophytes will catch stuff. I'd proofed one paper a million times, and then my reviewer noted I'd reversed the images & descriptions left/right. Stupid. I also caught this same error in a published paper on Papilio glaucus maynardi.

In some focus fields, I have no peers. I am the authority. So who's to review the subject material?

In cases where the field has peers, and Lepidoptery is a good example, "peers" would include: competitive researchers vying for the same grant money, competitive institutions, the "Old School" famous Lepidopterists who fight change, those with ulterior motives, and the inept. So who the heck do you turn to?

Besides which, peer review take time- usually awaiting someone else's free time. There's a lot of back-and-forth about petty stuff. And, depending on who's doing the review, there are those who just want to nit pick. So there's a significant (year plus) delay to publication. And then, while you're going through peer review, someone else jumps in and describes the species.

Eurytides suggests that all new descriptions employ genetics. In a perfect world, yes. But the same problems with peer review also arise with genetics. I've been waiting years for genetic testing. I've paid for some out of my own pocket, and as Eurytides is aware, a more comprehensive analysis was quoted at $8000. So I or we are supposed to pony up another $8000 on top of the probably $60,000 I've spent on this hobby? So, I can understand those who publish without, or prior to receiving, full genetic analysis.

[adamcotton]

Eurytides suggests that all new descriptions employ genetics. In a perfect world, yes. But the same problems with peer review also arise with genetics. I've been waiting years for genetic testing. I've paid for some out of my own pocket, and as Eurytides is aware, a more comprehensive analysis was quoted at $8000. So I or we are supposed to pony up another $8000 on top of the probably $60,000 I've spent on this hobby? So, I can understand those who publish without, or prior to receiving, full genetic analysis.

I actually don't think that mandating genetic analysis for new taxon descriptions is a good idea. Apart from access of some researchers in far-away places to genetic analysis facilities (and the ability to pay for it) there is also the issue that sometimes different taxa do NOT differ in their genetics, particularly at ssp. level. If two populations are consistently visibly distinct in the vast majority of specimens then they should deserve subspecies status whether or not they are genetically different at all.

Adam.

[bobw]

I believe that genetic analysis is just one of several tools that can be used in taxonomy, and certainly not the most impoortant. It's not the panacea that everybody thought it was going to be, I have seen papers relying on this technique that defy all logic from a morphological and ecological point-of-view. As Adam points out, it is expensive and in many cases requires fresh, perfectly preserved specimens to be reliable. There are only very few labs with sophisticated enough equipment to be able to extract useable samples from old specimens, and they are beyond the reach of most researchers.

[jhyatt]

I believe that genetic analysis is just one of several tools that can be used in taxonomy, and certainly not the most impoortant.

I tend to agree with Bobw: I view CO1 barcoding as just one more character than can, and probably should if feasible, be considered when analysing the degree of relationship between specimens.

My 2 cents' worth,
jh

[Trehopr1]

This topic has certainly turned into a fascinating discussion.

I thank everyone for their insights, knowledge, and expertise in bringing more clarity to the topic.

[kevinkk]

It is very interesting. That's why I like hanging out with you persons:)

I can see a lot goes into deciding new species, I had always thought, that among other factors, differences in the larva, the ability to produce ,or not
to produce offspring were factors, along with microscopic details like genitalia. Not every test is always going to be viable for every type.

[eurytides]

I was not suggesting that descriptions of sp or ssp should rely solely on genetics. If that’s the case, then why study anything else at all (host plants, phenology, hybrid offspring viability…)? However, we live in modern times and much progress has been made in the field of genetics such that I think it warrants inclusion in the Code. Imagine if the Code was written before the invention of microscopes and looking at genitalia didn’t matter. Then, someone invents the microscope and realizes that genitalia differences can be informative when discerning taxa. Would it not be sensible to include that in the toolbox? I believe the same is true of molecular information.

As for bjorkae and Harry, my understanding is that TILS doesn’t really have much of a peer review process (if any). It seems like a conflict of interest for Harry to write a paper and then publish it in a “journal” he runs “out of his basement.” Most good academic journals have a pretty high rejection rate and publications go through only after peer review and revisions. He is bypassing this integral part of the scientific process and automatically approving his own paper for which he is the soul author. I can’t help but view such a publication with some skepticism. It is possible/probable that bjorkae is a legitimate species and I am not saying Harry is definitely wrong or anything. I just hesitate to say he is 100% right either and I personally would like to see more genetic data and some critique from third party entomologists before accepting his paper as gospel.