A Catalogue of the Butterflies of the United States and Canada with a complete bibliography of the descriptive and systematic literature

2008·328.5 MB·

by  Jonathan P Pelham

HE Journal of Research
N THE LePIDOPTERA
A
Catalogue
of the Butterflies
the United States
of
and Canada
with a complete bibliography
the
of
descriptive and systematic
literature
by
Jonathan Pelham
P.
Volume 40 2008
/
TheJournal of Research on the Lepidoptera
ISSN 0022 4324
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:
A CATALOGUE
OFTHE
BUTTERFLIES OF THE UNITED STATES
AND
CANADA
WITH ACOMPLETE BIBLIOGRAPHY
OFTHE
DESCRIPTIVE AND SYSTEMATIC LITERATURE
by
Jonathan P. Pelham
Curatorial Associate ofLepidoptera
Burke Museum ofNatural History and Culture
Box 353010, University ofWashington
WA
Seattle, 98195 3010
[email protected]
Journal ofResearch on the Lepidoptera
Volume 40
2008
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Twenty-sevenyearshavepassedsincethepublicationofA Catalogue/ChecklistoftheButterfliesofAmericaNorth
ofMexicohyLee D. Millerand F. Martin Brown (1981). Since its publication, hundreds ofbooks andjournal
articles on the taxonomyand systematics ofthe butterflies ofthe United States and Canada have appeared,
newspecies in the region have been discoveredand described, and manynewnames have been proposed to
representgeographicvariationdisplayedbyNorthAmericanbutterflies. Inaddition,previouslyundocumented
specieshave beenfoundinTexas,NewMexico,andArizona, onanalmostyearlybasis,asstraysfrom Mexico,
oras breeding populations at the northern edges oftheir current ranges. The same twenty-seven years has
seentremendousimprovementsinourunderstandingofhigher-levelrelationships (oftribes,subfamiliesand
families) among the world’s butterfly fauna, largely thanks to advances in biotechnology and data analysis,
which have enabled a number ofdetailed molecular phylogenetic studies. This wealth ofnew information
on the butterflies ofthe United States and Canada has made clear the need for an updated systematic and
bibliographic catalogue treating the entire fauna.
Thisvolume is the resultofover20yearsofintensivelaborbythe author,Jonathan Pelham,who hasgone
through the extraordinary effort to personally examine every literature source related to North American
butterflies (save averyfew that proved too rare to be accessed), including the original descriptions ofevery
taxon occurring in the region. As a result,Jon is perhaps more knowledgeable of the literature on North
Americanbutterfliesthananyotherperson,andisalsoextremelyfamiliarwiththeNeotropicalandPalaearctic
literature on butterflies. Coupledwith thisvastbibliographic knowledge,Jon has extensive field experience
withbutterflies (adultsandimmatures) in manypartsofthe NorthAmericancontinent, andhasan intimate
knowledge ofthe Code ofZoological Nomenclature. These unique qualities, in addition to his incredible
determination and perseverance, have all been critical during the compilation ofthis Catalogue.
As editor ofthisvolume, I have enjoyed the undeniable privilege ofbeing able toworkwith the contents
ofthe Catalogue over the past two years, during its peer and editorial review periods, and preparation for
publication. IalreadyconsiderthisCataloguetobeabsolutelyindispensableinmydailylifeasaLepidopterist,
due not only to the details in the body of the Catalogue, but also because of the massive bibliography, an
invaluableresourceforeverystudentofNorthAmerican butterflies, nowandin tbefuture. Itismyhope that
thisCataloguewillfinditswayontothebookshelvesofeveryoneinterestedin NorthAmericanbutterflies,for
nowhere else will you find a modern “all in one” resource on their taxonomy, systematics, and bibliography.
ThisdocumentrepresentsamilestoneinourunderstandingofthebutterfliesoftheUnitedStatesandCanada,
andformsasolidfoundationuponwhichfuturediscoveriesandphilosophiescaneasilybeincorporated. We
are all very much indebted toJonathan Pelham, for bringing this Catalogue to fruition.
Andrew D. Warren, PhD
Editor,/ouma/ofResearch on theLepidoptera
McGuire CenterforLepidopteraand Biodiversity
Florida Museum ofNatural History
University ofFlorida
SW 34th Streetand Hull Road
P. O. Box 112710
Gainesville, FL 32611-2710 (USA)
Museo de Zoologfa “Alfonso L. Herrera”
Departamento de Biologfa Evolutiva
Facultad de Ciencias
Universidad NacionalAutonoma de Mexico
Apdo. Postal 70-399
Mexico, DE04510 Mexico
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V
PREFACE
Thisdocumentisacompilation ofall the namesapplied tobutterfliesoftheUnitedStatesofAmericaand
Canada as ofJune 2006. The names are arranged in taxonomic orderfrom family to species, and ancillary
detailsare given foreach, including the original description, original combination, type-locality, deposition
ofprimary type(s) when known, and annotations when pertinent to their placementwithin the body ofthe
catalogue. Greateffort has been made to include everyname that hasbeen applied to a butterflyoccupying
thisgeographicregion,and toensurethatinformation providedforeachnameisaccurate. More than twenty
years have been spentcompilingthisvolume. Dozensoflibrariesand collections have been consulted, and a
thorough investigation ofthe literature has been conducted, includingthe greatvolume ofrecentliterature.
In addition, I have relied heavily on my colleagues for information and advice.
The immediate predecessor to this catalogue is the “Catalogue/Checklist ofthe Butterflies ofAmerica
North ofMexico”byLeeD. MillerandF. Martin Brown (1981). ThatworkwasprecededbyWilliamF. Kirby’s
“Synonymic Catalogueofthe DiurnalLepidoptera”publishedin 1871 (with itssupplementin 1877), Herman
Strecker’s“CompleteSynonymicalCatalogueofMacrolepidoptera,”publishedin 1878,andSeitz “Die Gross-
,
SchmetterlingederErde,vol. 5” (1907-1924). Theseare theonlycompletecataloguesofthebutterflyfaunaof
the United States and Canada. In preparing this catalogue. Miller& Brown (1981) provided the foundation
uponwhich to build, and theirworkiscited throughout. Some features ofstyle andformatofthis catalogue
have been adopted from Miller & Brown, but the systematic format employed here, including most of the
family-level, genus-level and species-level arrangements, follows Opler&A. Warren (2002). Deviationsfrom
Opler & A. Warren are mostly the result ofinformation published since 2002. Everyyear, new information
on the taxonomy and systematics ofNorth American butterflies is presented, and any taxonomic catalogue
becomes dated upon publication and delivery. I acknowledge this with the expectation that such works,
opinions and contributions rendered in the future may be incorporated in perennial updates, in the hope
that this catalogue may remain current and diverse opinions may be collated and documented.
VI
INTRODUCTION
The purpose of this work is to systematically enumerate all taxa that have been recorded within the
boundaries of the United States of America and Canada. This catalogue is not intended to serve as a
taxonomic revision ofthe butterfliesin this region, and provides no solutions to difficult taxonomic issues; it
isacompilation; abibliographic and Code compliantendeavor. Everyefforthas been made toavoid making
revisionary decisions on the status oftaxa, but in awork ofthis type, some revision is inevitable.
The Merriam-Webster Online Dictionary (http://www.m-w.com/) includes among its definitions of a
catalogue “Acomplete enumeration ofitems arranged systematicallywith descriptive details.” Mayr (1969)
definedataxonomic catalogueas“An index to taxonomic literaturearrangedbytaxa...” Thiscataloguegoes
onestepfurtherbyincludingacompletebibliographyofthesystematicliterature pertainingtothebutterflies
ofthe region. The catalogue enumerates the 800 butterfly species from the United States ofAmerica and
Canada, distributed within 234 genera, including all subspecific and infra-specific names, synonyms, and
emendations. In addition, appendices ofspecies and names no longer included in the covered region are
provided.
NOMENCLATURE
It is impossible to systematically arrange any group oforganisms in a manner agreeable to all, and the
popularityofbutterflies hasamplifieddivergentopinions on theirtaxonomyand nomenclature. There have
been andwill continue tobedisagreementsoverwhatconstitutesafamily,subfamily, tribe, genusandspecies.
However, thanks to the last decade of phylogenetic studies on the higher-level systematics in butterflies,
which have employed molecularand morphologicalsourcesofdata, the family-level nomenclatureforNorth
American butterfly taxa is arguably better resolved than arrangements at the genus- and species-level. The
comments belowprovide some personal and historical perspective on these matters, insofaras they relate to
the present catalogue.
Family-level arrangement
PaulEhrlich’s (1958) “Thecomparativemorphology,phylogenyandhigherclassification ofthebutterflies”
providedthefoundation uponwhichsubsequenthigher-levelstudiesofPapilionoideahaveevolved (although
this study did not include Hesperioidea). This arrangement was popularized in Paul and Anne Ehrlich’s
(1961) “How to Knowthe Butterflies”. In the arrangement offamilies, ifnot sequence, the presentcatalogue
follows Ehrlich (1958) as modified by Eliot (1973), deJong et al. (1996), Ackery et al. (1999), Brower (2000,
2006), Wahlberg and Nylin (2003), Wahlberg et al. (2003b, 2005b), Ereitas and Brown (2004), Braby et al.
(2006), Pena et al. (2006), and Niklas Wahlberg’s website, which is regularly updated. The higher-level
arrangement ofHesperioideafollowsA. Warren (2006). Through these studies, a general consensus on the
number and composition ofbutterfly families has emerged (Wahlberg, et al. 2005a). It seems unlikely that
future arrangementswill diverge significantlyfrom the family-level arrangement presented here.
Genus-level arrangement
Butterflies have received close attention from taxonomists, both professional and amateur, and many
generic names have been generated as a result. Acomparison between butterflygeneric nomenclature and
its structural-morphological foundations, and those ofalmost any other group oforganisms, demonstrates
that there is littlejustification for the dissolution oflarge genera like Callophrys, Plehejusd.nd Papilio, simply
because they are “species rich” and displaya greatdiversity ofwing phenotypes. The generic nomenclature
employed here generallyfollows the works ofCdench (1961, 1970b), Eliot (1973),J. Miller (1987), Gorbunov
(2001),andOplerandA.Warren (2002) amongothers,although somediscrepanciesexist. Someattempthas
been made to accommodate thosewho acceptdivision ofsome large genera, includingPapilio'And Callophrys
intosmallerunitsbyemployingsubgenerathroughoutthecatalogue,despitequestionable monophylyofmany
ofthese subgenera. This is a subjective matter and it is appreciated that the catalogue might be considered
biased in this respect, while allowing for differing interpretations.
,
VII
Species-level arrangement
As noted above, the species-level arrangement in this catalogue largely follows Opler and A. Warren
(2002), as modified by subsequent studies. At the species level, the introductory section of Opler and A.
Warren (2002) is instructive. They noted, “Some ofthe more popularspecies concepts held by taxonomists
todaydon’teven recognizesubspecies.” Thatsaid,acataloguearrangedaccordingtoanycriterion excluding
subspecies would be unsatisfactory to many users. Therefore, the subspecies category is employed in this
catalogue without regard to its validity. It must be noted that defining species and subspecies boundaries
remains contentious, even among those sharing species concepts. Defining an objective measure for “what
is a species?” is remote from the aim ofthis catalogue.
Zoological Nomenclature
The International Commission ofZoological Nomenclature has produced a body ofrules, updated and
revised periodically, defining how names are to be treated in the context ofzoological nomenclature. The
most recent edition is “The International Code of Zoological Nomenclature, Fourth Edition”, published
in 1999 (hereafter referred to as ‘the Code’). These “rules” define nomenclatural protocol and are cited
throughout. Application oftheCodeisguidedbytwomain objectives: “...topromotestabilityanduniversality
in the scientific names ofanimals and to ensure that the name ofeach taxon is unique and distinct.” This
catalogue complieswith the Code in all respectssave one. The single divergence relates to the conformation
ofthe gender ofspecific names with the gender of the genus in a binomen (Code Articles 31.2 and 34.2).
Problemsassociatedwith thesecodearticleswerediscussedindetail byLamas (2004b:xv-xvi) Incaseswhere
.
the emendation ofnames in order to conform withArticles 31.2 and 34.2 ofthe Code seems clearand is not
in dispute, a note is appended to the discussion ofthat name, indicating its conformed spelling (synonyms
and unavailable names are not included in such cases). However, species-level names used in this catalogue
reflect their original orthography.
Catalogue Format
The systematic arrangement ofthe catalogue is intuitive once a few principles are stated. Each specific
entryisnumberedsequentially;generaandsubgeneraare not numbered. Subspecificentriesforeachspecies
are indentedonce, andare precededbylower-case Roman lettersinalphabeticalsequence. Datesin brackets
indicate that these have been subsequently determined from sources external to the original document.
Misspellings are excluded exceptwhere they have beenwidelyused or misinterpreted. All nomina nuda are
included in the bodyofthe catalogue. Annotationsare includedwhere appropriate. Alternative taxonomic
arrangements (e.g., Durden 1982 or Scott 1986), are indicated in many cases. Synonyms are listed under
each valid name.
Genus-level synonymies:
The synonyms ofeach genus-level name are arranged so that:
= subjective synonyms
these are not indented, are preceded byan “=” symbol, and are italicized.
j:permanently invalid synonyms
(e.g., homonyms, objective synonyms, unjustified replacement names, etc.) These are indented once,
preceded byan “=” symbol, a double dagger symbol, and are italicized.
j:”nomina nuda”
theseareindentedonce,placedinquotes, precededbyadoubledaggersymbol (butnotan “=”symbol)
and are not italicized.
Species-level synonymies:
The synonyms ofeach species-level name are arranged so that:
VIII
= subjective synonyms
these are not indented, are preceded by an “=” symbol and are italicized.
= j:permanently invalid synonyms
(e.g., homonyms, objective synonyms, unjustified replacement names, etc.) These are indented once,
preceded byan “=” symbol and a double dagger, and are italicized.
= j:”unavailable names”
(e.g., forms, aberrations, quadrinomials, etc.) These are indented once, placed in quotes precededby
an symbol and a double dagger, and are not italicized,
f’nomina nuda”
theseareindentedonce,placedinquotes, precededbyadoubledaggersymbol (butnotan “=”symbol),
and are not italicized.
Speciesforwhichthenominatesubspeciesdoesnotoccurintheareacoveredhaveanasterisk precedingthe
nominate entry. The symbol “f” is employedwhere a taxon is considered extirpated from the region covered.
Subspecies-level arrangement
The arrangement of subspecies under each polytypic species follows a north to south or east to west
geographic arrangement, similar to that presented in Howe (1975), so that geographically contiguous
populations appear next to each other in the catalogue. This may appeararbitrary in situationswhere taxa
arearrayedalongboth theSierraNevada-Cascade,andtheRockyMountains, butitispreferabletoasequence
that is either alphabetic or based on name seniority.
PRELIMINARY NOTES
IntheprocessofconstructingthiscatalogueIencounteredcertaindifficultiesregardingthepropercitation,
dating,orauthorshipofcertainworks, locationoftypespecimens,andthespellingofoneman’sname. Mostof
these issues have been dealtwith in the body ofthe catalogue and bibliography; others are detailed below.
Location oftypes
Generally, information on the location oftype specimens has been extracted from original descriptions.
However, many older descriptions did not indicate the deposition of types, and in some cases, the actual
deposition of types differs from that noted in the original description. For a large number of taxa, the
location oftype material remains unknown. While no special efforts have been made to verify information
on the deposition oftypes, several colleagues have been ofgreat help in this regard, and two websites have
been useful.
ANSP Insect Type Collection
(http;//data.acnatsci.org/biodiversity_databases/insects.php?mode=highertaxa), and Harvard University’s
MCZ Type Database
http://mcz-28168.oeb.harvard.edu/mcz/FMPro?-db=Species.fm&:-lay=web&-format=search.htm&-view
F. M. Brown (1974b) detailed the holdings ofthe butterfly type material in theANSPand stated “Several
yearsagotheAcademyofNaturalSciences,Philadelphia, exchangeditsbutterflycollectionwith theCarnegie
Museum ofnatural History.” TheANSPwebsite details the specimens thatwould have been involved in such
an exchange as ifthey are still atANSP. I treat these as ifthey are still extant at the ANSP.
John Eatton Le Conte
John Calhoun (2005b) recently published the complete history ofthe misspelling ofthis man’s name.
The citation whichwasfrom Rehn (1954) employed byOplerandA. Warren (2002) is incorrect and this has
been remedied in the catalogue.
The “Entomologische Beytrage...”
This work was published byJohann A. E. Goeze, between 1777 and 1783. The text proposed names for