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<div style="font-family:arial,helvetica;font-size:10pt;color:black"><font size="2">After publication of Glassberg's article, several people have inquired with me regarding the status of the Appalachian Tiger Swallowtail (Pterourus appalachiensis), thinking it is now just a "subspecies" of the Eastern Tiger Swallowtail because that is Dr. Glassberg's opinion based on viewing two single specimens during a brief trip! Sorry, but this is just plain wrong.</font></div>
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<div class="MsoNormal">Commentary on <i>Searching for ‘Appalachian’ Tiger
Swallowtails in the North Carolina Mountains </i>by Jeffrey Glassberg (American
Butterflies 29(3), page 22, by Harry Pavulaan.<o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">In this article, Dr. Glassberg’s <u>personal</u> view that
“it’s not clear that Canadian and Eastern Tiger Swallowtails are actually two
species…” contradicts what several experts in the field have concluded many
decades ago with numerous scientific studies.
I grant him his <u>personal</u> view that these are a single species,
but he’s in the minority view and does not concur with the actual experts on
the subject who have done the research.
So much for scientific research.<o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">At issue here is that many species are no longer considered clear-cut
“black and white” entities by those doing actual research. There are gray, mosaic or checkerboard areas
of many degrees between many species. Species
boundaries are apparently being discovered to be fluid or porous, leading to
exchange of genes between closely-related species (introgression). Some taxa fly together in some areas and
behave as separate species, while in other areas, they comingle and interbreed
freely. Tiger Swallowtails aside, the
western Fritillaries and Blues all present a similar challenge to the biologist. Dr. Glassberg cites his doubt of species
status of Canadian and Eastern Tiger Swallowtails as “they appear to hybridize
over a large area in the northeast”.
It’s not that simple. Several studies
have demonstrated that the purported northeastern hybrid zone consists of at
least four distinct taxa: <i>Pterourus glaucus, P. canadensis</i>, an “early
flight” (EF) taxon and a “late flight” (LF) taxon. The EF and LF taxa occur north (or above) of
where <i>glaucus</i> is able to produce two annual generations and where it is generally
too cold for <i>glaucus</i> to survive the winter. The EF and LF taxa are stable entities
produced over thousands of years of hybridization between <i>glaucus</i> and <i>canadensis</i>
during the last glacial maxima. They may
not fit traditional characterizations as “species” but are uniquely evolved taxa
that may or may not be considered species, depending on which “species concept”
you follow. Yet they are not active
hybrid populations, though some research points to the LF taxon as a possible recent
development of introgression of <i>glaucus</i> genes into <i>canadensis</i> in
places like Vermont. In Ontario, it is
believed that the LF taxon represents a valid species-level taxon – the
“Mid-summer Tiger Swallowtail”. Whatever
the case, one must be careful not assume that two species, <i>glaucus</i> and <i>canadensis</i>,
are individually cross-mating throughout a purported “hybrid zone”.<o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">I will admit that there appears to be active hybridization
going on in New England, but all evidence I’ve seen shows it is NOT <i>glaucus </i>x
<i>canadensis</i>. It is likely that <i>glaucus</i>
is hybridizing with the LF (Mid-summer Tiger Swallowtail) taxon in southern New
England, as there are appears to be a broad range of intermediates between the
two in summer, but it’s still not so simple.
There are other factors at work.
Thus, species are not “black and white” and we have to accept that there
are other classes of taxa that are of unique status. The EF Tiger appears quite distinct from <i>glaucus</i>
in New England, and could be much older than the LF Tiger that could be a much
more recent development.<o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">The Appalachian Tiger Swallowtail is one such “hybrid” taxon,
though it has fully speciated since the last ice age. It is not, as Dr. Glassberg suggested: a
“subspecies” of the Eastern Tiger Swallowtail.
It is accepted by biologists that two subspecies cannot be fully (100%)
sympatric, and do not occupy the same range, habitats, and flight period. The Appalachian Tiger inhabits higher
elevations of the southern and central Appalachian Mountain region. Unfortunately, Dr. Glassberg did not spend
enough time in the N.C. mountains to see the females, that are strikingly
different from Eastern Tiger females.
Females of both species can be found together, and over many years of
studying these, I have found no “intermediates” suggestive that they are the
same species, nor clear evidence of hybridization. Dr. Glassberg’s two sightings of males in
North Carolina does not weigh against decades of research by the “experts” on
the subject.<o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">For those of you still doubting the status of the
Appalachian Tiger Swallowtail, do your own research. See what the experts (and some observant
authors) are finding. Here is some help:<o:p></o:p></div>
<div class="MsoNormal"><u><o:p> </o:p></u></div>
<div class="MsoNormal"><u>BIBLIOGRAPHY OF <i>PTEROURUS APPALACHIENSIS </i>LITERATURE<o:p></o:p></u></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Belth, J.E. 2013. Butterflies of Indiana – a Field Guide. Indiana University Press, Bloomington,
IN.: 328 pp. <a href="https://books.google.com/books?hl=en&lr=&id=c9B4x7j6cJEC&oi=fnd&pg=PP1&dq=papilio+appalachiensis&ots=15RTJIUunp&sig=nI_RcB7WojxYV0SirwkjkzfF2FY#v=onepage&q=papilio%20appalachiensis&f=false">https://books.google.com/books?hl=en&lr=&id=c9B4x7j6cJEC&oi=fnd&pg=PP1&dq=papilio+appalachiensis&ots=15RTJIUunp&sig=nI_RcB7WojxYV0SirwkjkzfF2FY#v=onepage&q=papilio%20appalachiensis&f=false</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Borchelt, R.
2014. Side by Side
Swallowtails: Eastern and Appalachian
Tigers. LepLog. May 20, 2014.<o:p></o:p></div>
<div class="MsoNormal"><a href="https://leplog.wordpress.com/2014/05/20/side-by-side-swallowtails-eastern-and-appalachian-tigers/">https://leplog.wordpress.com/2014/05/20/side-by-side-swallowtails-eastern-and-appalachian-tigers/</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Bright, S.B., V. Charny & W.M. Howell. 2013.
Butterflies: New records for Alabama.
Journal of Alabama Academy of Science 84(1): 37-42.
<a href="https://alabama.butterflyatlas.usf.edu/static/documents/JAAS_Vol_84_No1_37-42.pdf">https://alabama.butterflyatlas.usf.edu/static/documents/JAAS_Vol_84_No1_37-42.pdf</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Chapman, P.D., S.P. Bradley, E.J. Haught, K.E. Riggs, M.M.
Haffar, K.C. Daly & A.M. Dacks.
2017. Co-option of a
motor-to-sensory histaminergic circuit correlates with insect flight
biomechanics. Proceedings of the Royal
Society B 284: 20170339. <a href="http://dx.doi.org/10.1098/rspb.2017.0339">http://dx.doi.org/10.1098/rspb.2017.0339</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Cong, Q., D. Bork, Z. Otwinowski & N.V. Grishin. 2015.
Tiger Swallowtail genome reveals mechanisms for speciation and
caterpillar chemical defense. Cell
Reports 10(6): 910-919.<o:p></o:p></div>
<div class="MsoNormal"><a href="https://doi.org/10.1016/j.celrep.2015.01.026" target="_blank" title="Persistent link using digital object identifier"><span style="font-size:10.5pt;font-family:"Arial",sans-serif;color:#0C7DBB">https://doi.org/10.1016/j.celrep.2015.01.026</span></a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Kunte, K., C. Shea, M.L. Aardema, J.M. Scriber, T.E.
Juenger, L.E. Gilbert & M.R. Kronforst.
2011. Sex chromosome mosaicism
and hybrid speciation among Tiger Swallowtail butterflies. Plos Genetics 7(9): <span style="font-size: 10pt; font-family: Helvetica, sans-serif; color: rgb(32, 32, 32); background-image: initial; background-position: initial; background-size: initial; background-repeat: initial; background-attachment: initial; background-origin: initial; background-clip: initial;">e1002274. </span><a href="https://doi.org/10.1371/journal.pgen.1002274"><span style="font-size: 10pt; font-family: Helvetica, sans-serif; color: rgb(3, 113, 101); background-image: initial; background-position: initial; background-size: initial; background-repeat: initial; background-attachment: initial; background-origin: initial; background-clip: initial;">https://doi.org/10.1371/journal.pgen.1002274</span></a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Mavárez, J. & M. Linares.
2008. Homoploid hybrid speciation
in animals. Molecular Ecology 17:
4181-4185. doi:
10.1111/j.1365-294X.2008.03898.x <o:p></o:p></div>
<div class="MsoNormal"><a href="https://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.531.8795&rep=rep1&type=pdf">https://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.531.8795&rep=rep1&type=pdf</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">North Carolina Dept. of Parks. 2021.
Butterflies of North Carolina: their Distribution and Abundance. Online:
<a href="https://auth1.dpr.ncparks.gov/nbnc/a/accounts.php?acctID=6">https://auth1.dpr.ncparks.gov/nbnc/a/accounts.php?acctID=6</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Ogard, P.H.
2010. Butterflies of Alabama –
Glimpses into Their Lives. University of
Alabama Press<o:p></o:p></div>
<div class="MsoNormal"><a href="https://books.google.com/books?hl=en&lr=&id=TtgtDwAAQBAJ&oi=fnd&pg=PT12&dq=butterflies+of+alabama&ots=CYa8yIplf5&sig=cc9kU5Au0qpYOk15E3Y836L1zLI#v=onepage&q=butterflies%20of%20alabama&f=false">https://books.google.com/books?hl=en&lr=&id=TtgtDwAAQBAJ&oi=fnd&pg=PT12&dq=butterflies+of+alabama&ots=CYa8yIplf5&sig=cc9kU5Au0qpYOk15E3Y836L1zLI#v=onepage&q=butterflies%20of%20alabama&f=false</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Ording, G.J., R.J. Mercader, M.L. Aardema & J.M.
Scriber. 2010. Allochronic isolation and incipient hybrid
speciation in tiger swallowtail butterflies.
Oecologia: DOI 10.1007/s00442-009-1493-8. 9 pp.<o:p></o:p></div>
<div class="MsoNormal"><a href="https://d1wqtxts1xzle7.cloudfront.net/43328971/Allochronic_isolation_and_incipient_hybr20160303-26273-q7ohdq.pdf?1457046190=&response-content-disposition=inline%3B+filename%3DAllochronic_isolation_and_incipient_hybr.pdf&Expires=1631839179&Signature=Grnec6NqNsIe66MYEdcB2KTTIWfvti~cD6Cy-p0xp~ow2hDsbQQzo96CBH7ZklPTC--ueTx5paEjgjOaxdBUBH2iQGt9BzhwuZcHTGxsr9ArCzEo0QXcv1n~uyugb84~YR9ZqJfGmLQa2PDWWjJ5g-rISo2WJWLnwEIFJYLFzB2X9RnCCKx3RoBRueSGGAL-e716pNUClC6BaismukGZly-9k~3p5GViUjx~OQcyJ0QxCLdf1HTwE1Zw35GJivYyEj~S5pVkNJJKivlN7CC-Y9-0E81mTuXNXLsRIs6yUdeFqrB569RiRiln1jKXdxuACzlfDoIYmUiq9Pb2itztLA__&Key-Pair-Id=APKAJLOHF5GGSLRBV4ZA">https://d1wqtxts1xzle7.cloudfront.net/43328971/Allochronic_isolation_and_incipient_hybr20160303-26273-q7ohdq.pdf?1457046190=&response-content-disposition=inline%3B+filename%3DAllochronic_isolation_and_incipient_hybr.pdf&Expires=1631839179&Signature=Grnec6NqNsIe66MYEdcB2KTTIWfvti~cD6Cy-p0xp~ow2hDsbQQzo96CBH7ZklPTC--ueTx5paEjgjOaxdBUBH2iQGt9BzhwuZcHTGxsr9ArCzEo0QXcv1n~uyugb84~YR9ZqJfGmLQa2PDWWjJ5g-rISo2WJWLnwEIFJYLFzB2X9RnCCKx3RoBRueSGGAL-e716pNUClC6BaismukGZly-9k~3p5GViUjx~OQcyJ0QxCLdf1HTwE1Zw35GJivYyEj~S5pVkNJJKivlN7CC-Y9-0E81mTuXNXLsRIs6yUdeFqrB569RiRiln1jKXdxuACzlfDoIYmUiq9Pb2itztLA__&Key-Pair-Id=APKAJLOHF5GGSLRBV4ZA</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Pavulaan, H. & D.M. Wright. 2002. <i>Pterourus
appalachiensis </i>(Papilionidae: Papilioninae), a new swallowtail butterfly
from the Appalachian region of the United States. The Taxonomic Report of the International
Lepidoptera Survey 3(7): 20 pp. <a href="http://www.lepsurvey.carolinanature.com/ttr/ttr-3-7.pdf">http://www.lepsurvey.carolinanature.com/ttr/ttr-3-7.pdf</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Pavulaan, H. & D.M. Wright. 2004.
Discovery of a black female form of <i>Pterourus appalachiensis </i>(Papilionidae:
Papilioninae) and additional observations of the species in West Virginia. The Taxonomic Report of the International
Lepidoptera Survey 6(1): 10 pp. <a href="https://www.lepsurvey.carolinanature.com/ttr/ttr-6-1.pdf">https://www.lepsurvey.carolinanature.com/ttr/ttr-6-1.pdf</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Schmidt, B.C.
2019. More on Ontario Tiger
Swallowtails. Ontario Insects 2019: 11 pp.<o:p></o:p></div>
<div class="MsoNormal"><a href="https://www.ontarioinsects.org/publications/Summaries/2019_tigers.pdf">https://www.ontarioinsects.org/publications/Summaries/2019_tigers.pdf</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Scriber, J.M.
2010. Impacts of climate warming
on hybrid zone movement: Geographically diffuse and biologically porous
“species borders”. Insect Science
18(2): 121-159. <a href="https://onlinelibrary.wiley.com/doi/10.1111/j.1744-7917.2010.01367.x">https://onlinelibrary.wiley.com/doi/10.1111/j.1744-7917.2010.01367.x</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Scriber, J.M.
2013. Climate-driven reshuffling
of species and genes: Potential conservation roles for species translocations
and recombinant hybrid genotypes.
Insects 5(1): 1-61.<o:p></o:p></div>
<div class="MsoNormal"><a href="https://www.mdpi.com/2075-4450/5/1/1/htm">https://www.mdpi.com/2075-4450/5/1/1/htm</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Scriber, J.M.
2015. Tales of Three Tigers: A
50-Year Career-Shaping Journey Chasing Swallowtail Butterflies. <i>In: </i>Dyer, L. & M. Forister (eds),
The Lives of Lepidopterists. Springer
International Publishing, Switzerland. <a href="https://doi.org/10.1007/978-3-319-20457-4_10"><span style="font-size:10.5pt;font-family:"Source Sans Pro",sans-serif;letter-spacing:.2pt;background:#FCFCFC">https://doi.org/10.1007/978-3-319-20457-4_10</span></a><span style="font-size:10.5pt;font-family:"Source Sans Pro",sans-serif;color:#333333;letter-spacing:.2pt;background:#FCFCFC"><o:p></o:p></span></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Scriber, J.M. B. Elliot, E. Maher, M. McGuire & M.
Niblack. 2014. Adaptations to “thermal time” constraints in <i>Papilio</i>:
Latitudinal and local size clines differ in response to regional climate
change. Insects 5: 199-226. <a href="file:///C:/Users/harry/Downloads/insects-05-00199-v2%20(2).pdf">file:///C:/Users/harry/Downloads/insects-05-00199-v2%20(2).pdf</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Scriber, J.M. & G.J. Ording. 2005.
Ecological speciation without host plant specialization; possible
origins of a recently described cryptic <i>Papilio </i>species. Entomologia Experimentalis et Applicata
115(1): 247-263. <a href="https://doi.org/10.1111/j.1570-7458.2005.00285.x"><span style="font-size:10.5pt;font-family:"Arial",sans-serif;color:#005274">https://doi.org/10.1111/j.1570-7458.2005.00285.x</span></a><span style="font-size:10.5pt;font-family:"Arial",sans-serif;color:#767676"><o:p></o:p></span></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Scriber, J.M., G.J. Ording & R.J. Mercader. 2008.
Introgression and Parapatric Speciation in a Hybrid Zone. Chapter 6.
<i>In: </i>Tilmon, K. (ed.).
Specialization, Speciation, and Radiation: The Evolutionary Biology of
Herbivorous Insects. California
Scholarship Online: 69-87. <a href="https://california.universitypressscholarship.com/view/10.1525/california/9780520251328.001.0001/upso-9780520251328">https://california.universitypressscholarship.com/view/10.1525/california/9780520251328.001.0001/upso-9780520251328</a><span class="MsoHyperlink"><o:p></o:p></span></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">University of Texas at Austin. 2011.
Appalachian tiger swallowtail butterfly is a hybrid of two other
swallowtails, scientists find. Phys Org
online, Sept. 8, 2011. <o:p></o:p></div>
<div class="MsoNormal"><a href="https://phys.org/news/2011-09-appalachian-tiger-swallowtail-butterfly-hybrid.html">https://phys.org/news/2011-09-appalachian-tiger-swallowtail-butterfly-hybrid.html</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Zhang, W., K. Kunte & M.R. Kronforst. 2013.
Genome-wide characterization of adaptation and speciation in tiger
swallowtail butterflies using <i>de novo</i> transcriptome assemblies. Genome Biology and Evolution 5(6):
1233-1245. <a href="https://academic.oup.com/gbe/article/5/6/1233/619586?login=true">https://academic.oup.com/gbe/article/5/6/1233/619586?login=true</a><o:p></o:p></div>
<div class="MsoNormal"><o:p> </o:p></div>
<div class="MsoNormal">Seeing is believing.
If anyone is interested in viewing this magnificent swallowtail
firsthand, please write to me and I’ll consider organizing a field trip to the
best location to see it in West Virginia, or provide directions.<o:p></o:p></div>
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