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Papilio. New Series

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Papilio (New Series), an entomology journal, is a scientific journal that covers the systematics and taxonomy and biology of butterflies, mostly from Colorado. There are about 700 species of butterflies in North America and about 270 in Colorado, New discoveries are made every year on the Colorado species. Systematics is the study of the kinds of butterflies that exist on our planet, and taxonomy involves the names of butterflies, including the description and naming of species new to science. Papilio (New Series) started in 1981 to name the butterflies in single publications. This digital collection includes the single issues as they are published.

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  • ItemOpen Access
    Chaetotaxy of first-stage butterfly larvae, with improved homologies and nomenclature for lepidoptera setae and sensilla
    (Colorado State University. Libraries, 2020) Scott, James A., author; James A. Scott, publisher
    This paper has two goals. The first goal, of interest to all Lepidopterists, is to improve the names given to the setae and olfactory pores and sensillae: new homologies of the setae are presented including those on the last abdomen segment (good homologies on this segment are presented for the first time); a name is assigned for every seta and sensilla of the entire body including mouthparts, building upon Hinton (1946); different names are assigned to non-homologous setae that formerly were confused by having the same name; figures and an alphabetical glossary of structures are presented for users. The second goal is to present new data on butterflies, including setal maps for every subfamily known (only Pseudopontiinae and Calinaginae now lack setal maps), an improved key that includes additional subfamilies and genera, and diagnostic characters for each family, subfamily, and tribe. Scott (1986a) presented setal maps for selected first stage butterfly larvae and presented a key to all known North American subfamilies. Scott (1985, 1986b) and Scott and Wright (1990) used characters of first stage larvae as well as many other characters to deduce the phylogeny of butterflies.
  • ItemOpen Access
    Butterflies of the southern Rocky Mountains area, and their natural history and behavior: photos of mostly eggs larvae pupae. Part IV. Lycaenidae
    (Colorado State University. Libraries, 2020) Scott, James A., author; James A. Scott, publisher
    These four issues of Papilio (New Series) are photos for my book "Butterflies of the Southern Rocky Mts. Area, and their Natural History and Behavior", (https://hdl.handle.net/10217/200723) showing some adults but mostly early stages (eggs, 1st-stage, mature larvae, and pupae) of as many of the species as possible, primarily from the Southern Rockies area (I added a few other interesting species that do not occur in the area). They have been cropped and downsized to illustrate just the butterflies and conserve kilobytes, rather than serve as artistic images. They are arranged by evolutionary relationship, as in the book text.
  • ItemOpen Access
    Butterflies of the southern Rocky Mountains area, and their natural history and behavior: photos of mostly eggs larvae pupae. Part I. Hesperiidae
    (Colorado State University. Libraries, 2020) Scott, James A., author; James A. Scott, publisher
    These four issues of Papilio (New Series) are photos for my book "Butterflies of the Southern Rocky Mts. Area, and their Natural History and Behavior", (https://hdl.handle.net/10217/200723) showing some adults but mostly early stages (eggs, 1st-stage, mature larvae, and pupae) of as many of the species as possible, primarily from the Southern Rockies area (I added a few other interesting species that do not occur in the area). They have been cropped and downsized to illustrate just the butterflies and conserve kilobytes, rather than serve as artistic images. They are arranged by evolutionary relationship, as in the book text.
  • ItemOpen Access
    Butterflies of the southern Rocky Mountains area, and their natural history and behavior: photos of mostly eggs larvae pupae. Part III. Nymphalinae (Anaeini to Melitaeini)
    (Colorado State University. Libraries, 2020) Scott, James A., author; James A. Scott, publisher
    These four issues of Papilio (New Series) are photos for my book "Butterflies of the Southern Rocky Mts. Area, and their Natural History and Behavior", (https://hdl.handle.net/10217/200723) showing some adults but mostly early stages (eggs, 1st-stage, mature larvae, and pupae) of as many of the species as possible, primarily from the Southern Rockies area (I added a few other interesting species that do not occur in the area). They have been cropped and downsized to illustrate just the butterflies and conserve kilobytes, rather than serve as artistic images. They are arranged by evolutionary relationship, as in the book text.
  • ItemOpen Access
    Butterflies of the southern Rocky Mountains area, and their natural history and behavior: photos of mostly eggs larvae pupae. Part II. Papilionidae, Pieridae, Nymphalidae (Libytheinae to Satyrinae)
    (Colorado State University. Libraries, 2020) Scott, James A., author; James A. Scott, publisher
    These four issues of Papilio (New Series) are photos for my book "Butterflies of the Southern Rocky Mts. Area, and their Natural History and Behavior", (https://hdl.handle.net/10217/200723) showing some adults but mostly early stages (eggs, 1st-stage, mature larvae, and pupae) of as many of the species as possible, primarily from the Southern Rockies area (I added a few other interesting species that do not occur in the area). They have been cropped and downsized to illustrate just the butterflies and conserve kilobytes, rather than serve as artistic images. They are arranged by evolutionary relationship, as in the book text.
  • ItemOpen Access
    Butterflies of the southern Rocky Mountains area, and their natural history and behavior
    (Colorado State University. Libraries, 2022-03) Scott, James A., author; James A. Scott, publisher
    This book reports the biology of the butterflies of the southern Rocky Mountains area, including all the species in Colorado, although surrounding areas are also discussed, especially the rest of the Southern Rocky Mts. in Wyoming and New Mexico and into Utah. This book presents what is known of the biology of the butterflies of Colorado and vicinity, including hostplants, eggs/larvae/pupae appearance and habits, behavior including flight habits and migration and mate-locating and mating and basking and roosting, and the flowers and other foods of adult butterflies, and natural history aspects of their biochemistry, plus mimicry, flight periods and number of generations, etc. It also includes taxonomic matters to assist identification of all the species and subspecies and forms. Much research on the biology of Colorado area butterflies has been done recently, but it has been published in many scattered publications and scientific journals and is not readily available, and some good research is unpublished; this book attempts to make it available, and provides the sources for good published research.
  • ItemOpen Access
    New taxa and geographic variation of western North American butterflies: based on specimens in the C. P. Gillette Museum of Arthropod Diversity, Colorado State University
    (Colorado State University. Libraries, 2017-05) Scott, James A., author; Davenport, Ken E., author; Kondla, Norbert G., author; Opler, Paul A., author; Fisher, Michael S., author; James A. Scott, publisher
    During the two winters of 2016-2017, Scott volunteered to curate the butterflies in the C. P. Gillette Museum of Arthropod Diversity (CSUC), at Colorado State University, Fort Collins, Colorado. After more than 700 hours of identifying and sorting ~30,000 butterflies, all are arranged by species and identifiable subspecies, except for various tropical and Palearctic groups beyond Scott's expertise. Studying the specimens revealed some unnamed (identifiable) subspecies, and documented the geographic variation of many species. New research discoveries are reported here.
  • ItemOpen Access
    Ernest J. Oslar, 1858-1944: his travel and collection itinerary, and his butterflies
    (Colorado State University. Libraries, 2016) Scott, James A., author; James A. Scott, publisher
    Ernest John Oslar collected more than 50,000 butterflies and moths and other insects and sold them to many taxonomists and museums throughout the world. This paper attempts to determine his travels in America to collect those specimens, by using data from labeled specimens (most in his remaining collection but some from published papers) plus information from correspondence etc. and a few small field diaries preserved by his descendants. The butterfly specimens and their localities/dates in his collection in the C. P. Gillette Museum (Colorado State University, Fort Collins, Colorado) are detailed. This information will help determine the possible collection locations of Oslar specimens that lack accurate collection data. Many more biographical details of Oslar are revealed, and the 26 insects named for Oslar are detailed.
  • ItemOpen Access
    Mead’s butterflies in Colorado, 1871
    (Colorado State University. Libraries, 2016) Scott, James A., author; James A. Scott, publisher
    Theodore L. Mead (1852-1936) visited central Colorado from June to September 1871 to collect butterflies. Considerable effort has been spent trying to determine the identities of the butterflies he collected for his future father-in-law William Henry Edwards, and where he collected them. Brown (1956) tried to deduce his itinerary based on the specimens and the few letters etc. available to him then. Brown (1964-1987) designated lectotypes and neotypes for the names of the butterflies that William Henry Edwards described, including 24 based on Mead's specimens. Brown & Brown (1996) published many later-discovered letters written by Mead describing his travels and collections. Calhoun (2013) purchased Mead's journal and published Mead's brief journal descriptions of his collecting efforts and his travels by stage and horseback and walking, and Calhoun commented on some of the butterflies he collected (especially lectotypes). Calhoun (2015a) published an abbreviated summary of Mead's travels using those improved locations from the journal etc., and detailed the type localities of some of the butterflies named from Mead specimens. Unfortunately, the data published to date is a mess, because bits and pieces were published here and there, but the whole has not been systematically collected and organized and fully analyzed. To organize, all the information from every available source of data was computerized, combined into an organized whole, discrepancies and contradictions were fixed, and that was used to produce several useful products. The current study will form a solid foundation onto which can be added the last major potential improvement: a detailed listing of all the Mead 1871 specimens in museums that contain month and date on the labels.
  • ItemOpen Access
    About Papilio (New Series)
    (Colorado State University. Libraries, 2014-10) Scott, James A., author; James A. Scott, publisher
    This entomology journal is a scientific journal that covers the systematics and taxonomy and biology of butterflies, mostly from Colorado. There are about 700 species of butterflies in North America, and about 270 in Colorado, and new discoveries are made every year on the Colorado species. Systematics is the study of the kinds of butterflies that exist on our planet, and taxonomy involves the names of butterflies, including the description and naming of species new to science. The word Papilio comes from the scientific name Papilio of Swallowtail Butterflies, very large butterflies common in Colorado. I started Papilio (New Series) in 1981 when I was working on a book on the biology of North American butterflies for Stanford Univ. Press (see Scott 1986 in the list of publications below) and found several dozen butterflies that needed to be in the book but lacked names, so I decided to name them in one publication rather than go through the onerous process of getting several dozen separate papers published.
  • ItemOpen Access
    Identification of Phyciodes diminutor, P. cocyta, and P. tharos in northeastern U.S. (Nymphalidae)
    (Colorado State University. Libraries, 2014) Scott, James A., author; James A. Scott, publisher
    This issue of Papilio (New Series) consists of three studies on the identification of the closely-related species Phyciodes diminutor diminutor Scott, P. cocyta selenis (W. Kirby)and P. tharos tharos (Drury), from Vermont, Ohio, and Michigan (studies labeled A, B, and C below). Identification problems have hindered the study of these butterflies in the northeastern U.S., and misidentifications have even occurred in recent DNA studies. Difficulties in distinguishing diminutor & selenis, and their status, are discussed. Vermont contains all three taxa in this group. P. tharos has two generations and occurs in southern and central Vermont, north to Woodsville on the Connecticut River on the E edge of Vermont, and occurs also on North and South Hero Islands in Lake Champlain in extreme NW Vermont. P. cocyta has one generation and evidently occurs in the cooler areas throughout Vermont except in the two southern counties Bennington and Windham. P. diminutor has two generations and evidently occurs throughout Vermont if it belongs to a separate species P. diminutor as studies elsewhere suggest. Michigan has all three species, while northern Ohio (including Lucas Co.) has P. tharos and P. diminutor. Few of the specimens of P. cocyta, P. tharos, and P. batesii used in the mtDNA study of Proshek & Houghton (2006) were misidentified (among all specimens examined, a few "tharos" and "cocyta" are P. diminutor, some "batesii" females from one site are P. cocyta, and all "cocyta" from S Mich. are P. diminutor). mtDNA is not usable to identify any of these species. The proper future research plan is described, which includes rearing numerous families found on wild identified hostplants from localities with multiple taxa.
  • ItemOpen Access
    Systematics and life history studies of Rocky Mountains butterflies
    (Colorado State University. Libraries, 2014) Kondla, Norbert G., author; Scott, James A., author; Gray, Richard E., author; James A. Scott, publisher
    This note discusses previously-unrecognized adult wing pattern forms of Papilio multicaudata Kirby, a discussion of ecological causes of these forms, and the reasons that the name pusillus Austin & J. Emmel is an early seasonal form and not a subspecies. Form minimulticaudata is tiny in size, and is caused by starvation of second-generation larvae in the lower mountains the previous late summer. This research was started when Michael S. Fisher was studying the forms he was observing. As a result, the tiny form minimulticaudata was named by Fisher & Scott in Fisher (2012), who summarized the distribution and flight periods of the forms.
  • ItemOpen Access
    Argynnis (Speyeria) nokomis nokomis: geographic variation, metapopulations, and the origin of spurious specimens (Nymphalidae)
    (Colorado State University. Libraries, 2014) Fisher, Michael S., author; Scott, James A., author; James A. Scott, publisher
    Geographic variation in wing pattern within ssp. nokomis is documented across its range from northern New Mexico to southern Colorado and southeastern Utah and northeastern Arizona. This variation allows one to pinpoint the origin of collected specimens within that range, and defines the maximum possible areas of metapopulations. All of the numerous specimens of ssp. nokomis labeled from Ernest Oslar from southwestern and central Colorado, including the mislabeled neotype, are actually part of a hundred nokomis collected by Wilmatte and Theodore Cockerell from Beulah, New Mexico, so that is the true nokomis type locality. Ssp. tularosa is an invalid synonym because it is identical to Beulah ssp. nokomis, the provenance of all specimens is dubious as all were mislabeled from Sacramento Mts. but evidently actually collected at Beulah by the Cockerells or Henry Skinner, the purported altitude is too low, known collectors did not find it at the mislabeled Sacramento Mts. sites when it supposedly occurred there, numerous other mislabeled nokomis exist, and the only valid specimens from those mountains are another subspecies coerulescens.
  • ItemOpen Access
    Biological catalogue of North American butterflies
    (Colorado State University. Libraries, 2008-12-03) Scott, James A., author; James A. Scott, publisher
  • ItemOpen Access
    Geographic variation and new taxa of western North American butterflies, especially from Colorado
    (Colorado State University. Libraries, 2008-12-03) Garhart, Matthew C., author; Stout, Todd, author; Kondla, Norbert G., author; Spomer, Stephen M., author; Fisher, Michael S., author; Scott, James A., author; Wright, David M., author; Marrone, Gary M., author; James A. Scott, publisher
    Michael Fisher is currently updating the 1957 book Colorado Butterflies, by F. Martin Brown, J. Donald Eff, and Bernard Rotger (Fisher 2005a, 2005b, 2006). This project has emphasized the necessity of naming certain butterflies in Colorado and vicinity that are distinctive, but currently have no name, as part of our goal of applying correct species/subspecies names to all Colorado butterflies. Eleven of those distinctive butterflies are named here, in the genera Anthocharis, Neominois, Asterocampa, Argynnis (Speyeria), Euphydryas, Lycaena, and Hesperia. New life histories are reported for species or subspecies of Neominois & Oeneis & Euphydryas & Lycaena that were recently described or recently elevated in status. Lycaena florus differs in hostplant, egg morphology, and somewhat in a seta on 1st-stage larvae. We also report the results of research elsewhere in North America that was needed to determine which of the current subspecies names should be applied to other butterflies in Colorado, in the genera Anthocharis, Neominois, Apodemia, Callophrys, Atlides, Euphilotes, PlebeJus, Polites, & Hylephila. This research has added additional species to the total of Colorado butterflies. Nomenclatural problems in Colorado Lycaena & Callophrys are settled with lectotypes and designations of type localities and two pending petitions to suppress toxotaxa. Difficulties with the ICZN Code in properly applying names to clines are explored, and new terminology is given to some necessary biological solutions.
  • ItemOpen Access
    Portable (six-drawer) cabinets for California Academy Drawers
    (Colorado State University. Libraries, 2006-04-28) Scott, James A., author; James A. Scott, publisher
  • ItemOpen Access
  • ItemOpen Access
    Corrections/reviews of 58 North American butterfly books
    (Colorado State University. Libraries, 2008-12-03) Scott, James A., author; James A. Scott, publisher
    Corrections are given for 58 North American butterfly books. Most of these books are recent. Misidentified figures mostly of adults, erroneous hostplants, and other mistakes are corrected in each book. Suggestions are made to improve future butterfly books. Identifications of figured specimens in Holland's 1931 & 1898 Butterfly Book & 1915 Butterfly Guide are corrected, and their type status clarified, and corrections are made to F. M. Brown's series of papers on Edwards; types (many figured by Holland), because some of Holland's 75 lectotype designations override lectotype specimens that were designated later, and several dozen Holland lectotype designations are added to the J. Pelham Catalogue. Type locality designations are corrected/defined here (some made by Brown, most by others), for numerous names: aenus, artonis, balder, bremnerii, brettoides, brucei (Oeneis), caespitatis, cahmus, callina, carus, colon, colorado, coolinensis, comus, conquista, dacotah, damei, dumeti, edwardsii (Oarisma), elada, epixanthe, eunus, fulvia, furcae, garita,hermodur, kootenai, lagus, mejicanus, mormo, mormonia, nilus, nympha, oreas, oslari, philetas, phylace, pratincola, rhena, saga, scudderi, simius, taxiles, uhleri. Five first reviser actions are made (albihalos=austinorum, davenporti=pratti, latalinea=subaridum, maritima=texana [Cercyonis], ricei=calneva). The name c-argenteum is designated nomen oblitum, faunus a nomen protectum. Three taxa are demonstrated to be invalid nomina nuda (blackmorei, sulfuris, svilhae), and another nomen nudum (damei) is added to catalogues as a "schizophrenic tax on" in order to preserve stability. Problems caused by old scientific names and the time wasted on them are discussed.
  • ItemOpen Access
    Butterfly hostplant records, 1992-2005, with a treatise on the evolution of Erynnis, and a note on new terminology for mate-locating behavior
    (Colorado State University. Libraries, 2006-04-28) Scott, James A., author; James A. Scott, publisher
    Hostplants of larvae, based on 1,014 records (including 474 records of ovipositions and 540 discoveries of eggs, larvae, or pupae in nature) from 1992 through 2005, are presented for butterflies (including skippers), mostly from Colorado, and some from Wyoming, Nebraska, and Minnesota. New life histories are given, including many notes on egg placement, overwintering stage, behavior, and ecology. Larvae and pupae of Colo. Cyllopsis pertepida can be either green or tan, and thus retain a seasonal polyphenism that is present in other Cyllopsis even though only one generation occurs in Colo. Erebia magdalena oviposits on large boulders. Phyciodes picta evidently eats an annual gummy aster in much of the northern part of its range. Still another bog butterfly has been found to be polyphagous (Pyrgus centaureae), adding to the many polyphagous bog butterflies previously known (many Boloria, Colias scudderii); Speyeria mormonia eurynome might be semipolyphagous as well, though conclusive evidence is unavailable. Cercyonis (sthenele) meadii oviposits in shade north of pine trees near its sedge host that grows in that shade. Coenonympha tullia has green and brown larval forms, and striped and unstriped pupal forms. Erebia epipsodea oviposits high on its grass hosts in the foothills, low on its grass hosts in the alpine zone, to moderate the temperature of the eggs. The pupa of Chlosyne palla calydon is black-and white, versus brown in Calif. C. palla palla. Thorybes pylades and Everes amyntula specialize on tendril-bearing (pea "vine") herbaceous legumes. Stinga morrisoni is the only known butterfly that chooses large bunch-grasses (seven species) of many grass taxa. Paratrytone snowi eats only Muhlenbergia montana. Erynnis icelus oviposits only on seedlings. The evolution of Erynnis is discussed, using many new characters of larvae and pupae and valval flexion. Mature larvae of some Pyrginae (Pyrgus communis, Pholisora catullus) that diapause become reddish in color, whereas non-diapausing mature larvae remain greenish. An appendix provides new terminology for describing mate-locating behavior.
  • ItemOpen Access
    Phyciodes (Phyciodes): more progress
    (Colorado State University. Libraries, 2006-04-28) Scott, James A., author; James A. Scott, publisher
    Recent modest findings in Phyciodes are reported. P. tharos tharos older larvae are not always blacker than P. cocyta, although they average darker. P. tharos tharos in eastern U.S. sometimes has orange antenna nudum, even as far south as SW Mississippi where orange is slightly more frequent than black. Study of mitochondrial DNA by Wahlberg et al. suggests that P. tharos is a distinct species that has received little genetic influx from P. cocyta, while P. cocyta, P. batesii, and P. pulchella have enormous overlapping variation in mtDNA due to ancient variability plus some later introgression with each other and with P. tharos. P. (cocyta) diminutor has been found in northern Ohio, based on are examination of P. "tharos" specimens used in an electrophoresis study by Porter & Mueller. That study came to the erroneous conclusion that P. tharos and P. cocyta are conspecific, but actually found that Mich. P. cocyta could be conspecific (based on electrophoresis alone) with (misidentified) Ohio P. (cocyta) diminutor. Orange-antenna populations previously reported from Penn. and vicinity are probably mostly P. (cocyta) diminutor. The species/subspecies status of three cocyta-group taxa are discussed: P. diminutor could be a species, and diminutor and incognitus could be conspecific, and all three could be ssp. of P. cocyta (a lot of rearing will be needed to determine their status). A neotype of P. cocyta selenis is designated. P. cocyta arenacolor is a valid local subspecies. A synonym of P. batesii batesii (maconensis) is discussed. P. batesii lakota rarely has orange antenna club. A conclusive history of the travels and owners of the lectotype/neotype Phyciodes pulchella has now been finalized, including the discovery that Foster Hendrickson Benjamin actually wrote the "a/c [according to] Hofer [Carl Hofer]" labels found on numerous Boisduval specimens, in 1925. Other corrections involving Boisduval types are made. The pupal cremaster width of P. pulchella pulchella is studied further and averages not as small as previously thought, though it may average smaller. P. pulchella near pulchella populations from the E slope of the southern Sierra Nevada show past introgression from ssp. camillus, as do some intergrading synonyms in Nevada. Other recent synonymies are discussed and presented in a completely annotated synonymic checklist. More larval/pupal descriptions are presented. Improved identification/description tables are given.