Azures in "Spring" in NH - what are they?

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Steve Mirick

Apr 26, 2024, 9:53:27 AMApr 26
to NHButterfly, Mass Leps
I've noticed that several butterfly folks out there are calling the
azures flying around right now (April & May) "Spring Azures" (celastrina
ladon) and entering them as such in iNaturalist.  It used to be that all
early spring azures in the northeast were called "Spring Azures", but
current research suggests that there are between 2 and 3 species of
Azures in April and May in New Hampshire.  These include the confusingly
named (and possibly non-occurring) Spring Azure (celastrina ladon), the
Northern Azure (celastrina lucia), and the enigmatic Cherry Gall Azure
(celastrina serotina).  Identification of these three azures is close to
impossible based on just photos.  It's a real mess out there right now!

Specimens collected in Vermont during their 1st butterfly atlas showed
only Northern Azure and Cherry Gall Azure, but DID NOT show any records
of "Spring Azure"!  The common early spring flying azure in Vermont is
now called the Northern Azure (c. lucia).  This is also the case in
Maine where the most recent publication "Butterflies of Maine and the
Canadian Maritime Provinces" only indicates one species of Azure in the
spring and that is the Northern Azure (they don't show any records of
Cherry Gall Azure). They state that "Spring Azure" has NOT BEEN
CONFIRMED IN MAINE! Both Maine and Vermont now treat nearly all April
and May celastrinas (azures) as Northern Azure (C. lucuia).....with a
few Cherry Gall Azures in Vermont.

Based on this information, it seems likely (or at least possible) that
Spring Azures (C. ladon) do not occur at all in NH!!!!  And everything
flying right now may in fact best be called Northern Azure!!  The key
problem is that Spring Azure is almost impossible to ID from Northern
alone, we have no idea if "Spring Azures" occur in NH.

THIS.  They show Spring Azures (C. ladon) as occurring throughout the
northeast and into southeastern Canada.  THIS IS FALSE as specimens from
Canada have shown that Spring Azures don't occur there except for a
couple of records.

The taxonomy is still evolving and the range of the Spring Azure is
still a mystery and it's actually possible there may be other hidden
species out there.  I've decided to call my early spring azures as just
"Azure" and enter all of my early spring records from April and May in
iNaturalist as "Holarctic Azures" (Celastrina sp.).  If you post to this
list or submit azure photos to iNaturalist during these months, you may
want to follow this procedure until we figure this mess out!  Summer
sightings in late June and July are likely a different species "Summer
Azure" (c. neglecta), but that's another problem to sort out, and
another story...

If anyone out there has any more recent updates on research or opinions,
it would be interesting to know about.

Bryan Pfeiffer does a GREAT JOB in discussing this complex on his web
site.  Highly recommended.  Click on the download PDF file in the first

And if you want to dig deeper, here are some other articles for review:

Steve Mirick
Bradford, MA

Nancy E. Burke

Apr 26, 2024, 10:05:52 AMApr 26
to, NHButterfly, Mass Leps
I think at least we'll have to sort out the "flyby" reports from the photos, to begin with, and just call the flyby's "Azure" as you suggest. A lot of the reports I've seen were flyby's, including mine this week.
Thanks for this clarification.

Nan Burke
Westborough, MA
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Apr 26, 2024, 4:06:02 PMApr 26
to Mass Leps, Steve Mirick
Steve, all:

Unfortunately, many people, especially NABA members who rely solely on Butterflies Through Binoculars or the Swift Guide, still are not aware that the Celastrina genus has been split into several species over the past 20 or so years.  Unfortunately, NABA only recognizes three species in North America.  The Pelham catalogue ( recognizes 11 species, all based on extensive peer-reviewed research.  Thus, many folks posting to iNat or BAMONA still think everything from coast to coast is "C. ladon".  That being said...

To address your email, first, New Hampshire has four Celastrina species: ladon, lucia, neglecta and serotina.  NH does have ladon, or what we are tentatively calling C. ladon.  This is based on the presence of Azures across the southernmost counties of New Hampshire which have the ladon male dorsal wing scale structure that separates ladon from all other Celastrina species.  That "version" of ladon is found throughout southern New England, including the two southmost counties in Maine, but there are no records from Vermont.  This population of ladon actually has hybrid traits with its northern cousin, C. lucia.  The ventral sides of ladon and lucia are virtually impossible to distinguish because each has the same exact range of variation, from spotted venters to heavily-patterned variants, but the males have different dorsal wing scale structures.  This is not "variation", as both scale types breed true as demonstrated by almost 4 decades of research.  Both ladon and lucia fly early, and C. serotina emerges about a full month later than either.  Serotina is easily distinguished by its whitish venter and well-developed spot pattern.

The newest wrench in the gears of Celastrina identification is that the multivoltine ecotype of C. neglecta has apparently established itself well into Massachusetts in recent years.  It previously only ranged north to around New York City and not across southern New England.  I've seen a number of images that clearly show the multivoltine neglecta spring form in Massachusetts.  The problem is that these emerge along with ladon and lucia in regions where each occurs.  Neglecta stands out from lucia and ladon because it is almost pure white beneath with very small spots.  People ask me how the "Summer" Azure can produce a spring flight.  Apparently, this has been the norm throughout the range of multivoltine neglecta - forever.  But we only discovered this in the 1990's.  What many people in the south consider C. ladon, turns out to be spring form neglecta.  This also has been proven out, over 4 decades of breeding and fieldwork.  C. neglecta is the predominant spring flight in many regions.  C. ladon, on the other hand, has a much smaller range than neglecta, as evidenced by examination of male wings.  When I review iNat records of ladon, probably 95% of those images in the U.S. are, in fact, spring brood neglecta.

So what about the traditional Summer Azure in New England?  That is a different univoltine (or partly bivoltine) neglecta ecotype that flies only in July and occasionally in September.  Thus, in southern New England, the multivoltine neglecta would fly in April, June and August, whereas univoltine (or partly bivoltine) neglecta flies in July and occasionally in September.  The two ecotypes leap-frog one another, giving the impression that neglecta flies continuously from June into August or occasionally into September.  This is an amazing situation, which we are still working on.

The recent Butterflies of Maine book, unfortunately, followed a flawed paper by Schmidt & Layberry that discounted C. serotina in Ontario and suggested it might merely be a "late lucia".  Schmidt & Layberry did not recognize C. ladon in Ontario, of which there are historical records around Pt. Pelee.  What we call ladon, based on the male wing scale structure, does occur in southern Maine.  I'm not sure why the authors of the Maine book did not consult with either myself or David Wright on Celastrina distributions.  Schmidt and Layberry also insisted that C. serotina does not occur in Ontario, based on what I consider premature assumptions and flawed research.  The authors of the Maine book followed suit and did not recognize C. serotina in Maine.  I am presently working on a research paper that will demonstrate the presence of serotina in both Ontario and Maine, based on wing color analysis, and will address flaws in the Schmidt & Layberry paper.  

Many of our discoveries with Celastrina come after the publication date of many recent guides, including the Kaufman guide.  The Kaufman guide has the most updated taxonomy, though the Pelham list on is the most current and authoritative.  Maps in virtually all guides are relatively useless.

I am attaching papers which are more recent, but most folks are not aware of.  The ranges of ladon and neglecta are clearly delineated.  Dave Wright and Gordon Pratt performed an extensive electrophosesis-based analysis of North American Celastrina populations, yet unpublished that support our conclusions, as reflected in the 2023 Pelham Catalogue.  The team of Nick Grishin is currently performing extensive allozyme studies of all North American populations, that hope to settle the taxonomic issues.  We are also exploring the presence of at least one, but possibly two new species in the eastern U.S. 

That Schmidt & Layberry paper can be found at: What Azure blues occur in Canada? A re-assessment of Celastrina Tutt species (Lepidoptera, Lycaenidae)  However, I urge readers to take great caution in accepting what is published in this paper.  Neither author consulted with Dave Wright or myself to hash over some of the questions they posed.  The paper is flawed, and I will address this in a paper in work, demonstrating C. serotina in Ontario and Maine.

In the meantime, I am attaching my most recent papers that should help answer some questions.  I welcome questions and inquiries!

Harry Pavulaan


Steve Mirick
Bradford, MA

T REPORT 10(2a) Celastrina ladon and Celastrina neglecta are distinct species.pdf
T REPORT 7-7 Virginia Celastrina lucia.pdf
T REPORT 6-6 serotina.pdf
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Apr 30, 2024, 6:16:31 AMApr 30
to '' via MassLep
Thanks Harry, for your help and information!  Many butterfliers in MA want to do the right thing with ID and reporting, but are constrained by the MA Butterfly Club's association with NABA, and NABA's insistence that only their list can be used for the properties that they own (e.g. journal, website).  I don't know if/when that will change, but it's good to read more about recent published work.  I'm curious - are there any studies that disagree with your azure work and support retaining NABA's, or other conflicting, classification(s)?

Similarly, ongoing updates about tiger swallowtail taxonomy on this listserve would be great!

Bill Benner

Sent from my Verizon, Samsung Galaxy smartphone
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John Calhoun

Apr 30, 2024, 6:16:56 AMApr 30
to MassLep
As a coauthor of the recent Maine and Maritimes butterfly book, I would like to respond to some points in Harry Pavulaan’s discussion of Celastrina.

Although Harry is quick to criticize our treatment of Celastrina in the book, we made a concerted effort to recognize that other species may occur in the region. We state that we "tentatively" follow the arrangement of Schmidt and Layberry (2016), and concede that "As the understanding of Celastrina taxonomy improves, other cryptic species (such as the Cherry Gall Azure) may be detected here." We also include a discussion of the possible presence of the Spring Azure and Cherry Gall Azure in Maine in the section “Butterflies of Possible Occurrence” at the back of the book, where we note that further study is needed to understand the status of such phenotypes. We even state that “It is possible that the Cherry Gall Azure will be confirmed in our region.” We were fully aware that there are different interpretations about the various phenotypes that occur in Maine, and admit that our treatment will not necessarily stand the test of time. This is also true of all the various studies out there, as well as those to come.
Harry also questions why we did not consult with him or David Wright – but we actually did. The treatment of Celastrina in our book was fully reviewed by David. Notice that the C. neglecta account includes the citation “D. M. Wright, pers. comm. to J. Calhoun.” It was David who suggested that the “Spring Azures” in southern Maine are possibly just Northern Azures that present Spring Azure traits, “perhaps the result of past genetic introgression from the Spring Azure at the northern limits of its range." David also reviewed the entire book prior to publication.
We attempted to present a reasonable treatment of Celastrina, but there are so many differing opinions about this group that no matter what is written, someone will disagree. In fact, a recent paper titled “A genetic atlas for the butterflies of continental Canada and the United States” (2024, PLoS ONE 19(4)) suggests that all North American Celastrina represent one species! However, I was told that not all the authors of this paper approve of that conclusion, so there can even be disagreement among coauthors of individual publications.
I think we need to be careful not to suggest that our own opinions are the only correct ones. In the end, it’s all about personal experience and interpretation. We must each decide which treatment to follow and forge ahead. I personally await extensive genomic analyses of all the various phenotypes, cross-referenced with biology and morphology, but that may be years away. Nonetheless, even those results will be open to interpretation.
Finally, one additional point needs to be clarified. Harry states that "Schmidt & Layberry did not recognize C. ladon in Ontario." In fact, these authors mention the discovery of several museum specimens of this species from Ontario, listing them in the text, and even figuring two of them. They ultimately state that “Celastrina ladon is therefore confirmed as part of the Canadian fauna for the first time. Although other literature and even photo records may exist, voucher specimens are needed to verify identification, at least until phenotypic variation and distribution of C. ladon in southern Ontario is better documented.”
Thanks for reading,
John Calhoun

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Apr 30, 2024, 11:51:19 AMApr 30
to MassLep

Sent from my Verizon, Samsung Galaxy smartphone

The difficulty with applying current genomic (DNA) analysis to Celastrina is that it does not work separating recently-evolved species, especially eastern Celastrina and many of the western Blues.  Species like the Appalachian and Dusky Azure, very readily differentiated by their life histories and appearance could be seen as one species if only current genomic analysis was applied, and nothing else.  The parts of their genetic makeup that differentiate them have not yet been found.  The team led by Nick Grishin is working on this.  In the meantime old-fashioned fieldwork is as reliable as ever.  In particular, Celastrina serotina has been studied for its life history, reared and observed by David Wright and myself for over 4 decades.  No more fieldwork is needed on those to demonstate that they are a species, and eventually the molecular makeup that differentiates them will be revealed.  Thus, my surprise that the Maine Butterfly survey and the Ontario lepidopterists did not recognize that species as ocurring in those regions from available data.  I have specimens from Maine and Ontario that match the original description from Rhode Island.



-------- Original message --------
From: 'John Calhoun' via MassLep <>
Date: 4/30/24 10:15 AM (GMT-05:00)
Subject: Re: [MassLep] Azures in "Spring" in NH - what are they?

Sorry, everyone, for the similar postings. 

I originally drafted my comments in Word and copied them over to the group, but this apparently brought some formatting issues that Google didn't like, so it appears that my first posts were held up in the system. The same thing happened when I submitted a similar post to NHButterfly, and Steve had to physically release it. Thinking that my original posts to MassLep were lost, I posted again using the slightly different comments I sent to NHButterfly. In the end, all three of my posts to MassLeps were released, as well as the one to NHButterfly. 

I suppose I should have waited it out a bit longer before posting more comments! Again, I apologize for the confusion.   

John Calhoun    

John Calhoun

Apr 30, 2024, 6:32:41 PMApr 30
to MassLep
Thanks, Harry. 

I agree that DNA analyses need to be combined with morphological and biological observations. For a long time, I considered C. serotina to occur in southern Maine, but the more I looked at the various phenotypes, the more confusing they became. This, and the fact that C. lucia has been seen ovipositing on cherry galls in Nova Scotia made things even more confusing. 

I remember being the first to record C. serotina in Ohio, way back in the 1980s, long before you guys described it in 2005. I thought it was odd that they were fresh when local C. ladon were worn, and they were common around cherry trees that were loaded with galls. I contacted David Wright in 1988 and told him about it. He mentioned that this was likely a new species. The rest is history!   


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