New Papilio described today

[adamcotton]

Congratulations to Chuck!

https://zookeys.pensoft.net/article/142202/ (open access)

A cryptic new species of tiger swallowtail (Lepidoptera, Papilionidae) from eastern North America
Charles J. DeRoller, Xi Wang, Julian R. Dupuis, B. Christian Schmidt

Abstract
In the eastern Great Lakes region of North America, two tiger swallowtail species have previously been recognized, Papilio glaucus Linnaeus, 1758 and Papilio canadensis Rothschild & Jordan, 1906. A third entity, the Midsummer Tiger Swallowtail, has been treated as a P. glaucus × canadensis hybrid, and exhibits a mosaic of both intermediate and unique morphological and biological traits. Here we demonstrate that rather than being a localized, historically recent hybrid phenomenon, the Midsummer Tiger Swallowtail maintains its morphological and physiological distinctness over a large geographic region in the absence of one or both putative parental species, and was first documented in the literature nearly 150 years ago. Papilio solstitius sp. nov. is physiologically unique in delaying post-diapause development, which results in allochronic isolation between the spring flights of P. glaucus and P. canadensis, and the late summer flight of P. glaucus. Similarly, the geographic range of Papilio solstitius spans the region between the northern terminus of P. glaucus and southern limits of P. canadensis, remaining distinct in areas of sympatry. Defining the taxonomic identity of this unique evolutionary lineage provides an important baseline for further inquiry into what has served as an exemplary species group in evolutionary study.

[Chuck]

Don’t forget member Eurytides is also a co author and actually got me involved in this. It was a long journey though collaborating with the professionals Julian and Chris was an amazing learning experience.

Thanks Adam!

[eurytides]

Adam, you must read every article that comes out every day lol. I thought for sure Chuck would post about it first.

[adamcotton]

Don’t forget member Eurytides is also a co author and actually got me involved in this.

Ah, I didn't associate member 'eurytides' with a co-author as I don't know his real name. Congratulations to eurytides, and thanks for involving Chuck too!

Adam, you must read every article that comes out every day lol. I thought for sure Chuck would post about it first.

Actually I found out about it because ResearchGate flagged the citation of one of my papers in my feed so I clicked on the notification and found out about the new paper, otherwise I wouldn't have known about it.

Adam.

[Chuck]

Adam always has some excuse why he’s not as all knowing as he appears, but the sheer volume of such anecdotes makes me think it’s a cover up.

So now that we have Midsummer Tiger presented, but admitted not everything known, I wonder what’s next. I’d do Spring Form but eurytides doesn’t get those up in Canada.

[adamcotton]

So now that we have Midsummer Tiger presented, but admitted not everything known, I wonder what’s next. I’d do Spring Form but eurytides doesn’t get those up in Canada.

I was wondering whether that is in the pipe-line or not. Mind you, if it is the people working on it are unlikely to tell us. I certainly never disclose information about new taxa that I am working on until after publication, and that is the norm otherwise someone will rush in with a poor quality publication to get their name on the taxon.

Adam.

[Chuck]

I was wondering whether that is in the pipe-line or not. Mind you, if it is the people working on it are unlikely to tell us. I certainly never disclose information about new taxa that I am working on until after publication, and that is the norm otherwise someone will rush in with a poor quality publication to get their name on the taxon.

Adam.

At any time, Scriber, Schmidt, Wang, or even most recently Pavulaan could have described MST with enough to meet the “oh yeah well prove me wrong” level. MST was no secret, it was low hanging fruit, and that’s what annoyed me. That and so little was known about it, particularly the range.


Spring Form is low hanging fruit. I think Pavulaan is onto something with bjorkae and “near canadensis.” I’ll bet $100 that our spring form is not glaucus, but is, like MST, closer to canadensis. Problem is getting COI at minimum to show it. The tests are unavailable to most citizen scientists.

Aside from some “low hanging fruit” my Solomon Islands material hasn’t even been closely looked at. The P Ulysses of course are no longer Ulysses. I probably have a dozen or more taxa that wouldn’t be hard to describe as new.

But my Solomons material will probably be donated first, and I’m moving away from Spring Form. There are other Tigers that merit review, beyond CA rutulus and AZ eurymedon. That said, and Trehpr has a point- does anyone care about slightly different Tigers even if scientifically they merit a taxonomic status? I don’t see anyone rushing out to either study or have in their collection both of the north and south split of what we lump under Maynardi.

There’s wierd stuff going on with Speyeria, I’ve seen enough of that in my bycatch, but I’m not touching that.

[JVCalhoun]

I have been chomping at the bit for this paper, and Chuck has patiently answered so many of my questions via email over the past few months. I have been withholding a paper of my own that updates the Maine state list of butterflies, suspecting that this taxon occurs in the state. I am now confident to add this species, along with three others, to the Maine list since the publication of our book, Butterflies of Maine and the Canadian Maritime Provinces, in late 2023. I have collected a dozen of these tiger swallowtails in southern Maine over the past fifteen years, and I always suspected that they were something different. I even placed a label with the series, reading "Pterourus undescribed." Thanks to these authors, it now reads "Pterourus solstitius" (and yes, I prefer the genus Pterourus for this group).

Knowing that the paper was imminent, I had been monitoring the ZooKeys website over the past week. The paper was posted at about 10:45 am (EST) Friday morning, and I alerted Chuck shortly after. I congratulate the authors on such a thorough job of supporting the recognition of solstitius as a discrete species. I look forward to studying it further in Maine.

Nice work!
John

[adamcotton]

John,

One problem that occurs to me is that there are several ancient synonyms of Papilio glaucus Linnaeus, 1758 from eastern USA, some of which may turn out to be senior synonyms of bjorkae Pavulaan, 2024 or the new taxon, solstitius.

The only extant lectotype specimen is pictured here:
https://data.nhm.ac.uk/media/linnean_ty ... efault.jpg
This is the lectotype of Papilio turnus Linnaeus, 1771, designated as lectotype by Honey & Scoble (2001, Linnaeus’s butterflies (Lepidoptera: Papilionoidea and Hesperioidea. Zoological Journal of the Linnean Society, 132: 277–399. doi:10.1006/zjls.2001.0265).
It is a male, and appears to be a summer generation specimen of P. glaucus. I assume this is not solstitius, but no-one really knows for certain whether some of the old names were based on specimens that actually belong to bjorkae or solstitius.

Probably, valid neotype designations should be made for Papilio glaucus Linnaeus, 1758, Papilio antilochus Linnaeus, 1758 and Papilio alcidamas Cramer, 1775 in order to fix these names as synonyms of the species currently known as Papilio glaucus. An attempt to designate neotypes was made by Pavulaan & Wright (2002, The Taxonomic Report, 3(7):1-20. https://lepsurvey.carolinanature.com/ttr/ttr-3-7.pdf) when they described Papilio appalachiensis. Unfortunately the four neotype designations in this publication are all invalid under the ICZN Code, since they do not conform to article 75, governing neotypes.

Adam.

[Chuck]

there are several ancient synonyms of Papilio glaucus Linnaeus, 1758 from eastern USA, some of which may turn out to be senior synonyms of bjorkae Pavulaan, 2024 or the new taxon, solstitius.


Interesting, and that never occurred to me, though it should have, as I’ve questioned the validity of some of the early breeding and back cross tests, given the source material.

To my mind though, the important outcome is that MST has been given a universally agreed name. That allows it to be identified for discussion, and argument. Prior to “soltitius”, as pointed out in the paper, it had several monikers. As do bjorkae, spring form, and “near canadensis”.

Should a type show up that turns out to be solstitius is, to me, just an exercise to satisfy ICZN. It would really be accidental, because the type described then was all-encompassing of Appy, bjorkae, canadensis, etc. and was treated as such for a long time.

I think the greatest element in the paper- which was already known and published, but not common knowledge, is that solstitius is, if stuffed in pre-existing boxes- canadensis, not glaucus. If a type shows up and is solstius, and solstitius is rendered to a form, then the moniker canadensis would disappear.

[adamcotton]

Interesting, and that never occurred to me, though it should have ...

To my mind though, the important outcome is that MST has been given a universally agreed name.

I think the greatest element in the paper- which was already known and published, but not common knowledge, is that solstitius is, if stuffed in pre-existing boxes- canadensis, not glaucus. If a type shows up and is solstius, and solstitius is rendered to a form, then the moniker canadensis would disappear.

Chuck,

Don't worry, types for the ancient names I mentioned won't 'show up', none of them are identifiably still in existence. That is why I recommended that at some time neotypes for these names that definitely belong to the taxon P. glaucus should be designated in order to protect all the newer names. I agree that it is very important that MST should have a scientific name.

Interesting that solstitius is shown to be close to canadensis rather than glaucus. I haven't had time to read the paper in detail yet, so hadn't noticed that, but am very impressed with work judging by the skim-through I did make.

Adam.

[JVCalhoun]

As mentioned by Adam, the four other names involved are antilochus (Linnaeus, 1758), turnus (Linnaeus, 1771), alcidamas (Cramer, 1775) (misspelled as "alcidamus" by some authors, including Pelham 2023), and glaucus (Linnaeus, 1758). A lectotype was designated for turnus by Honey and Scoble (2001), and it does appear to represent the current concept of glaucus. This leaves the neotypes of antilochus, alcidamas, and glaucus, as designated by Pavulaan and Wright (2002), in need of further assessment. Some argue that these neotypes are invalid, but the authors expressly state that "no type specimens are believed to exist," which reflects conventional wisdom, if not the conclusions of prior authors. However, a probable syntype of alcidamas is deposited in NHMUK, setting up a potential lectotype designation to represent that name. Hopefully, it's consistent with the concept of glaucus.

The type locality of antilochus was given as “America septentrionali” (North America). That of alcidamas was given as Jamaica, New York and Carolina (Jamaica is obviously in error), and that of glaucus was "America." Given the time period of these descriptions, the specimens probably originated from somewhere along the Atlantic Coast, between New York City and the Carolinas, which is outside the expected range of the genetic concept of solstitius. Of course, solstitius may be more widely distributed than currently known, but it probably doesn’t reach much farther south, given that it is more genetically aligned with P. canadensis. Its presumed range, north of the blend zone where canadensis and glaucus converge, suggests that what Pavulaan (2024) ascribes to solstitius in southern New England, south of the blend zone, is something else entirely, perhaps more akin to glaucus.

That being said, if the neotypes of antilochus, alcidamas, and glaucus are deemed invalid, and the potential syntype of alcidamas is questioned, then new neotypes should be designated to place all these names within the current concept of glaucus, and comfortably outside the concept of solstitius.

John

[adamcotton]

if the neotypes of antilochus, alcidamas, and glaucus are deemed invalid,

This has already been published by Chainey (2005, Zoological Journal of the Linnean Society, 145, 283–337), who stated that the neotypes are invalid because the authors did not satisfy the requirements of ICZN Code (Article 75.3.4) in that they did not state their reasons for believing the type material to be lost or cite the steps that they took to establish this. Gerardo Lamas confirmed that he also believes all 4 neotype designations are invalid, and I agree. The article governing neotype designation in the ICZN Code is very precise, and requires statements in the publication confirming the various clauses have been met. The reason for this is that neotype designation is a very serious issue for which several hoops need to be jumped through first and those jumps must be confirmed.

Adam.

[adamcotton]

However, a probable syntype of alcidamas is deposited in NHMUK, setting up a potential lectotype designation to represent that name.

This is almost correct, Chainey (2005) listed this specimen:
Possible syntype with Felder label ‘Coll. Lenep’ [no locality] (BMNH(E)#665024), is a fair although not exact match for original plate 21, figs A, A; it has been re-pinned and the apices of the hindwings have been replaced and painted.
Note, Chainey used the word 'possible' rather than 'probable', and it would have to be confirmed as a syntype before lectotype designation, which is rather difficult. The problem is that if the specimen designated as lectotype is subsequently shown not to be a syntype after all it loses its lectotype status.

Adam.

[JVCalhoun]

Please don't get me wrong, I'm not arguing in support of the validity of these designations, but I wanted to point out that there is some confusion over this issue. For example, Pelham (2023) notes that the designations by Pavulaan and Wright (2002) for turnus and alcidamas are invalid, but he does not state the same for those of glaucus and antilochus. I'm sure this is unintentional, but it can lead some researchers to believe that the latter two designations are acceptable.

This has already been published by Chainey (2005, Zoological Journal of the Linnean Society, 145, 283–337), who stated that the neotypes are invalid

Not quite. Chainey (2005) was referring only to the neotype of alcidamas, and he stopped short of declaring that it was an invalid designation. Instead, he remarked that it "does not appear to be valid," leaving it to others to confirm. These designations really need to be more formally evaluated in a paper in which new neotypes are designated. The designation of lectotypes and neotypes is indeed serious business, and those of questionable validity need to be more formally addressed.

The "possible syntype" of alcidamas mentioned by Chainey was subsequently interpreted by Pelham (2008-2023) to be a "probable syntype." Maybe he, or another researcher, had stronger feelings about its status.

John

[Chuck]

So after 250 years, there’s question about where they types of bloody Tiger Swallowtails, captured by the most famed lepidopterist, are and if they exist? I’ve misplaced stuff before, but if they exist I’d think someone would know about it. To expect someone to put in effort to locate them is beyond rediculous. They can’t even find the Holy Grail. And even if they found one it would turn into a case of Longines’ spear.

[JVCalhoun]

Unfortunately, there are numerous historical specimens, including types, missing for various reasons. This contributes to the need for neotypes. In the case of glaucus, Linnaeus did not personally collect the the specimen(s) from which he based his description of the species. In his original description, Linnaeus (1758) cited at least one specimen in "M.L.U.," which refers to Museum Ludovicae Ulricae (Queen Ludovica Ulrika's collection). Carl Clerck is believed to have illustrated one such specimen in the second part of his his Icones Insectorum Rariorum (1764). This illustration is shown below.

Linnaeus (1764) later attributed specimen(s) of glaucus to the Swedish naturalist Peter (Pehr) Kalm, one of Linnaeus' first students who traveled in North America from 1748 to 1751. The fate of the specimen(s) that Linnaeus used for his description is unknown. Honey and Scoble (2001) could not locate any type material.

download/file.php?mode=view&id=4149

[Chuck]

The original type specimen is, by default, THE species it is named to be, so long as it doesn’t turn out to be synonymous

The risk of a neotype is that it could be a different taxon.

For the draft of solstitius I’d picked a very nice, expressive specimen typical of the species. My Canadian colleagues replaced it with a Canadian specimen, correctly arguing it had less risk of being hybrid or a look-alike stray glaucus.

[JVCalhoun]

The risk of a neotype is that it could be a different taxon.

That is a risk, especially for closely-related taxa. In this case, one could designate the specimen in Clerck's illustration of glaucus as the lectotype, arguing that it is likely a syntype. Article 74.4 of the Code states "Designation of an illustration or description of a syntype as a lectotype is to be treated as designation of the specimen illustrated or described; the fact that the specimen no longer exists or cannot be traced does not of itself invalidate the designation."

This could work for glaucus, as neither canadensis nor solstitius are known to have dark forms (though appalachiensis rarely does). The illustration matches Linnaeus' (brief) description, and it seems to be clearly identifiable to species. The only drawbacks are that you don't have any metrics on the missing specimen, you can't sample genetic material from it, and it's geographical origin is uncertain.

John

[adamcotton]

The risk of a neotype is that it could be a different taxon.

I will add that the objective of any potential neotype designation would be to choose specimens that are definitely the same as currently recognised as P. glaucus and ensure that the chosen specimens do not belong to any of the other named taxa currently recognised.

Note that a neotype can be a modern specimen, but it must be housed in an institution named in the publication or be placed in the specified institution on publication (paraphrased from article 75.3.7).

Adam.