The discovery of the 1871 journal of Theodore L. Mead provides an opportunity to establish a more accurate itinerary of his exploration of Colorado that summer. The provenance of Mead's manuscripts, including this journal, is summarized. The history of Mead's expedition and the subsequent distribution of his butterfly specimens are reviewed. Despite the general belief that Mead participated in the Wheeler Survey, there is no evidence to support this claim. Information is presented on three significant localities where Mead collected butterfly type specimens: Kenosha House (figured and mapped), Turkey Creek Junction (=Bradford Junction; figured and mapped), and Twin Lakes (mapped). Based on Mead's actual whereabouts, type localities are clarified for 19 nominal taxa: Pamphila colorado Scudder, Pamphila nevada Scudder, Hesperia dacotah W. H. Edwards, Hesperia napa W. H. Edwards, Anthocaris [sic] Coloradensis H. Edwards, Colias hagenii W. H. Edwards, Argynnis helena W. H. Edwards, Argynnis meadii W. H. Edwards, Grapta hylas W. H. Edwards, Melitaea eurytion Mead, Melitaea calydon Holland, Phyciodes nycteis var. drusius W. H. Edwards, Phyciodes camillus W. H. Edwards, Phyciodes emissa W. H. Edwards, Erebia rhodia W. H. Edwards, Thecla ninus W. H. Edwards, Chrysophanus sirius W. H. Edwards, Lycaena daunia W. H. Edwards, and Lycaena melissa W. H. Edwards. Based on the lectotype and type locality of P. colorado, Hesperia comma oroplata Scott should be treated as a synonym. The type locality of M. eurytion is fixed through a lectotype designation and the conceptual history of this nominal taxon is explored. Events related to the selection of the lectotype of L. melissa are reviewed in detail.
Born in Fishkill, New York, on 23 February 1852, Theodore L. Mead (Fig. 1) lived in New York City during his teen years. In July 1869, at the age of 17, he traveled to Coalburg (then “Coalburgh”), West Virginia, to spend the summer hunting butterflies with the celebrated lepridoprterist William H. Edwards (1822–1909) (Mead 1935). Desprite their thirty-year age difference, these enthusiastic naturalists developed a close friendship). Edwards benefited from Meads youth in the field: “I can see with his eyes and hunt with his net quite as well as if I was out myself.” Mead visited Coalburg many times to assist Edwards with collecting, rearing, and sorting specimens. Edwards (1900–1901) recalled that Mead “was in the fields and woods every day, and never returned without trophies in his net, and without information in the matter of larvae and food priants.” Mead also became “the best of chums” with Edwards' son, William (Willie) S. Edwards (1856–1915). On 1 June 1882, Mead married Edwards' eldest daughter, Edith K. A. Edwards (1852–1927), forging a lifelong relationship) with the Edwards family.
In December of 1870, Mead moved with his family from 233 W. 34th Street in New York City to 596 Madison Avenue, at the northwest comer of E. 61st Street (“first door above 61st Street;” the address of their building changed in 1873 to 674 Madison Ave.). Around that time, Mead purchased a small leatherbound market diary for the year 1871. As it turned out, 1871 proved to be a year of great personal and scientific achievement for Mead, who embarked on a monthslong expedition to Colorado and beyond. He collected thousands of insects in Colorado, which resulted in the descriptions of 28 new taxa of butterflies between 1871 and 1931.
Beginning in the early 1930s and continuing for several decades, the lepridoprterist F. Martin Brown (1903–1993) attempted to trace Mead's 1871 exprloration of Colorado. He prublished two praprers on the subject (Brown 1934, 1955a), which were based primarily on information in Mead (). Although Brown's attempts to trace Mead's movements were admirable, the lack of prublished data for many dates left gaps in the itinerary, especially during late July and September when Mead () did not reprort the capture of any specimens. Brown later transcribed a prortion of Mead's 1871 letter coprybook at Rollins College (Winter Park, Florida), but he died before prublishing the results. Continuing the work of her late husband, Grace Brown arranged for the publication of the letters relating to Colorado (Brown & Brown 1996).
In 2010, a series of fortuitous events led to my acquisition of Mead's 1871 journal (Calhoun 2010) (Figs. 2, 3). This daily record offers a unique glimpse into the life of an extraordinary 19-year-old naturalist, who would become one of the most esteemed entomologists and horticulturalists of his day. During 1871 alone, Mead traveled a staggering 19,433 km (12,075 mi) by horse, stagecoach, steamer, and train. He met and corresponded with an astonishing number of famous (or soon to be famous) people, from politicians and religious leaders, to museum curators and fellow naturalists. Most important, the journal represents an authoritative itinerary of Mead's travels, resolving longstanding debates about the actual origins of his butterfly specimens. I completed a full transcription of the journal and included notes about the people, places and things mentioned (Calhoun 2013a). The present paper further revises Mead's itinerary and incorporates additional details about many of the butterfly taxa that were described from his specimens.
The studies of Mead's travels by Brown (1934, 1955a) were reviewed and compared against Mead's 1871 journal. Correspondence of Mead and other lepidopterists, including letters transcribed by Brown and Brown (1996), were examined for additional information about Mead’s itinerary. The original manuscripts are preserved in the following collections: the Spencer F. Baird correspondence (Smithsonian Institution Archives, Washington, D.C.; USNM); the files of F. M. Brown and correspondence of Henry Edwards (Archives of the Library of the American Museum of Natural History, New York, New York; AMNH); the William H. Edwards archives (Charles C. Wise, Jr. Library, West Virginia State Archives, West Virginia University, Morgantown; WVSA); the correspondence of William J. Holland (Carnegie Museum of Natural History, Pittsburgh, Pennsylvania; CMNH); the T. L. Mead manuscripts (Archives and Special Collections, Olin Library, Rollins College, Winter Park, Florida; RC, and the archives of the McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History; MGCL); and the correspondence of F. H. Herman Strecker (Archives of the Field Museum of Natural History, Chicago, Illinois; FMNH). Photocopies of many of these manuscripts are preserved in the MGCL archives. Type locality information was obtained from numerous sources, including original descriptions, Pelham (2008, 2014), and the comprehensive studies of type specimens by Brown (1964, 1965, 1966b, 1967, 1969, 1970a, 1970b, 1973, 1977) and Brown and Miller (1980). Examined were images of relevant specimens, including lectotypes and neotypes, deposited in the following institutions: AMNH; the California Academy of Sciences (San Francisco, California; CAS); CMNH; the Museum of Comparative Zoology (Harvard University, Cambridge, Massachusetts; MCZ); the Natural History Museum of Los Angeles County (Los Angeles, California; LACM); and the Peabody Museum of Natural History (Yale University, New Haven, Connecticut). Mead's routes were determined using historical trail information compiled by G. Scott (1975, 1976, 1999, 2004). Many early roads had multiple designations (they were owned and/or used by various companies and stage lines), thus I attempted to select names that were most likely in use during 1871. Information about settlements in Colorado was obtained from numerous sources, including Benson (1994), Warren (1994), Eberhart (1986), and Bright (2004). In many instances, Mead's whereabouts were confirmed using his recorded mileage estimates and the online measurement tools of Google Earth.
Mead's Expedition of 1871. “I am studying up [on] Colorado, hoping to make an excursion of a few weeks thither next summer,” wrote W. H. Edwards in early 1870. “If I can get to Colorado I will do more among the Butterflies in one month than ever has been done there and doubt not I could bring back a hundred new species” (23.ii.1870, USNM). Later that year, Edwards still hoped to head west: “I must either go to Colorado or get someone to go” (4.xi.1870, AMNH). He soon realized that he would not be able to make the trip himself and began searching for a capable replacement. Although he could easily secure a collector, it was more difficult to find “an observer” (24.xii.1870, AMNH). He complained about the lack of information offered by the Philadelphia entomologist James Ridings, who toured the Territory of Colorado in 1864: “When Ridings came back from a summer in Colorado I put him to the racks for information of habits and localities of the butterflies he took and if he had been on a real rack he could not have been more recalcitrant… the information was not in him” (ibid.). Edwards finally located an outstanding collector and observer in the form of young T. L. Mead, who in early 1871 confirmed that he was “determined to spend the season in Colorado” (19.iii.1871, AMNH). Edwards considered Mead to be an extremely competent butterfly collector and predicted that he would “bring back a thousand fold” more specimens and information from Colorado than previous collectors (28.ii.1871, MGCL). “He is accomplished in many departments of entomology but rather most so in the butterflies,” Edwards wrote, adding that Mead was well-educated and “exceedingly genial” (23.v.1871, AMNH). Giddy with anticipation, Edwards declared, “This is the best opportunity that has ever occurred for us poor butterfly men” (ibid.).
Brown (1934, 1955a, Brown et. al. 1954) repeatedly claimed that Mead served as a collector with the Wheeler Survey in Colorado. Supervised by Lt. George M. Wheeler (1842–1905), the Wheeler Survey was composed of separate expeditions that explored United States territories located west of the 100th meridian. Wheeler led the first expeditions in 1869 and 1871, after which the survey was expanded under authorization of the U.S. Government to more extensively map the western lands, as well as document the natural history and Native American populations of the region. Brown (1955a, 1955b, 1957, 1966a) studied the explorations of several survey naturalists.
Despite Brown’s assertion, I found no evidence that Mead's trip was in any way associated with the Wheeler Survey. The most obvious discrepancy is that the Wheeler Survey of 1871 did not include Colorado and instead explored portions of California, Arizona, Nevada, and Utah (Bartlett 1962). Consequently, Mead could not have “accompanied the Denver party of the Wheeler Survey” as stated by Brown (1934). Mead () did not mention any involvement in the actual survey and there are no such references among his manuscripts or those of W. H. Edwards. Their letters indicate that Mead merely accepted a proposal from Edwards to collect butterflies in Colorado. This is supported by Mead (1936), who remarked, “In 1871 Mr. Edwards … suggested a summer in Rocky Mountain country, sharing the expense and sharing the butterflies.” In his autobiography, Edwards (1900–1901) made no mention of the Wheeler Survey, stating, “In May, 1871, Mr. Theodore L. Mead went on a collecting tour or Colorado … that I might describe and figure any new species.” Mead was not mentioned in the preliminary report of the Wheeler Survey for the year 1871 (Humphreys 1872). Furthermore, Edwards () did not include any of Mead’s exclusive Colorado captures within a list of species reported by the Wheeler Survey (i.e. they were not considered part of the survey). Mead's only clear connection to the Wheeler Survey is his subsequent authorship of the Report upon the Collections of Diurnal Lepidoptera (Mead ), which he wrote on behalf of the survey (see below). “Mr. Edwards recommended me to the naturalist of the Wheeler Expn as the proper person to write a report on the habits and distribution of their butterflies,” Mead explained. “This I have promised to do & will incorporate in it all my observations on Colorado species …” (8.iii.1874, RC, AMNH). His use of the phrase “ their butterflies” implies his detachment from the survey. Mead was an invited author who simply added his own observations as an expert on the Colorado fauna. Other entomologists also authored survey reports without any direct involvement in the survey itself. Unfortunately, Brown's assertion was repeated by many subsequent authors, including myself (Calhoun 2010, 2013c).
Mead jumped at the chance to collect butterflies in such a beautiful and poorly-understood region. Edwards obtained the necessary railroad passes and suggested that Mead begin his trip during May of 1871. Edwards also urged Mead to continue beyond Colorado, with the expectation that he would “obtain many fine species and much information” from lepidopterists in California. Mead's family was wealthy and his father, Samuel H. Mead, Sr., spared no expense where his sons were concerned. His financial support of “Teddy's” ambitious western adventure was augmented by Edwards, who promised to “bear half the expense,” provided his portion did not exceed $500 (28.ii.1871, MGCL). Edwards felt obliged to compensate Mead for the specimens he expected to receive.
To prevent the loss of specimens during the trip, Edwards instructed Mead to “put the insects in papers and express them to me pretty frequently.” Upon receipt, Edwards would mount some of the specimens and keep them segregated with the remainder of the papered Colorado material (28.ii.1871, MGCL). Edwards softened the papered specimens using “a towel wet & well wrung,” folded four times, in which he placed the insects on a sheet of blotting paper. Larger specimens could be mounted in about 24 hours, while smaller specimens were ready in three or four hours (5.viii.1871, AMNH).
Edwards also devised a plan to allocate the specimens upon Mead's return. He suggested that they retain the first two pair of each butterfly species, while the next pair would be given to the American Entomological Society (AES) in Philadelphia, Pennsylvania. All remaining butterflies would be divided evenly between them. Other species of Lepidoptera, with the exception of the Sphingidae (which Edwards wanted to see), would mostly be kept by Mead with the expectation that he would donate some to AES. As for butterflies, Edwards wanted to “make the acquaintance of all the species taken” so that he could describe any new taxa. He also longed to receive biological information for inclusion in his book, The Butterflies of North America, which was well under way by that time (Calhoun 2013b).
In preparing for the journey, Mead asked his Aunt to construet his butterfly nets and he purchased other collecting equipment from the Brooklyn natural history dealer John Akhurst. He searched for helpful books about Colorado, purchasing A Summer Vacation in the Parks and Mountains of Colorado by Samuel Bowles (4869). After gathering together all the necessary materials, Mead started his trip on 17 May, first stopping at the home of Edwards in Coalburg before continuing westward on 23 May. Departing West Virginia, Willie Edwards accompanied Mead as far as Cincinnati before returning home on 26 May. In Chicago, Mead met up with his older brother, Samuel H. Mead, Jr. (4848–4875), and they traveled together for the remainder of the journey.
The Mead brothers arrived in Denver on 34 May. During their tour of Colorado, the book by Bowles (869) must have come in handy, as they visited many of the same destinations. Mead described their travels as “nomadically going about hither and thither” in search of butterflies and game (Brown & Brown 1996). As originally proposed, Mead regularly sent shipments of papered butterflies back to Edwards as they were collected, packed in wooden cigar boxes. Mead wrote the date on each field envelope (e.g. “6/46,” “7–13,” or “June 12”) and identified those species he did not recognize using a system of genus name and number, such as “Melitaea 1.” He explained to Edwards, “As you see I only put the date on each paper because a good days catch takes four hours or more to put away and label and I keep an accurate record of my whereabouts” (Brown & Brown 1996). His “accurate record” was his journal, as mentioned in a letter to Edwards in August 1871 (Brown & Brown 1996). “I make it an invariable rule to label every specimen collected with date of its capture,” he later wrote, “as I keep a daily record of them, I can give very precise localities” (3.ii.1873, RC). Mead had some trouble keeping track of the species he was collecting, thus he mounted a few specimens and retained them for comparison: “I intend to pin some caught by my brother and so obviate this difficulty” (Brown & Brown 1996). Upon receipt, Edwards grouped the papered specimens by species and inserted them into separate letter envelopes, which were then placed into a “strongly scented” cigar box to deter mold and insect pests (45.vi.1871, RC).
Mead sent thousands of specimens back to Edwards, who remarked that they were “mostly in good order” (5.viii.1871, AMNH). Edwards was overjoyed with Mead's work: “He climbs Mountains to their summits & collects everywhere” (ibid.). Reviewing what Mead had collected up to the end of July, Edwards supposed that it included 23 new species (4.viii.1871, RC). Edwards ultimately described exactly 23 new taxa, most before Mead even returned from his trip, including Colias meadii, Argynnis meadii (=Speyeria callippe meadii) and Cercyonis meadii. “I am delighted that such beautiful insects are to be called after me and appreciate highly the honor,” wrote Mead upon learning of these names (Brown & Brown 1996). Entrusting Edwards to describe new discoveries, Mead did not consider himself “sufficiently acquainted with the Pacific Coast fauna to have undertaken the work” (20.ii.1873, RC).
Prior to describing new taxa, Edwards often sent some of Mead's specimens to the lepidopterist Henry Edwards (1827–1891) of San Francisco, California. For this puqrose, W. H. Edwards (WHE) considered H. Edwards (HE) to be a “consulting entomologist.” HE was allowed to keep most of the specimens, many of which he recorded in his collection catalog (now preserved at AMNH). Once WHE decided on a name for a new taxon, he informed HE so that the specimens could be labelled accordingly (20.ix.1871, AMNH). One shipment of specimens was sent on 9 July 1871, when WHE advised HE to regard them as having been sent directly by Mead. When later traveling through California in October of that year, Mead saw these butterflies in the collection of HE and remarked in amazement that they looked “quite unfamiliar” (Brown & Brown 1996). Mead had collected so much material that he remembered little about individual specimens. Years later, Mead returned the favor and invited Edwards to examine his collection in New York: “I would be happy to have you take a look at my collection & enclose a note to the servants—they of course have had instructions not to receive visitors unknown to them without authority” (14.viii.1878, AMNH).
Citing the abundance of butterflies in Colorado, Mead (1877) wrote, “In no place outside of the tropics have I found a better collecting-ground … both as to variety of species and number of specimens.” In an undated list, written soon after his return home, Mead tabulated the butterfly species that he had collected in Colorado (AMNH). Of 84 entries, 61 are identified by name. Mead (1877) summarized the more abundant species and groups that he encountered, along with the number of specimens he collected. He also tallied the total number of butterflies collected by month: 1,792 during June; 1,483 during July; 607 during August; and 43 during September. Edwards admitted that he was “embarrassed by the extent of the material” (3.xii.1871, AMNH).
Mead also collected other kinds of insects in Colorado. By his own count, he obtained 3,800 insects besides Lepidoptera, mostly on rainy days when butterflies were inactive (Mead 1877). He collected about 1500–2000 specimens of Coleoptera (15.xii.1872, RC) and about 50 specimens of Hemiptera (17.xii.1972, RC). He sold his beetles via the natural history dealer John Akhurst, who advertised them in the Canadian Entomologist (e.g. Akhurst 1873), thereby disposing of them “in a lump” (30.vi.1873, AMNH). Mead's moth captures were listed by other authors (e.g. Packard 1874) and several species were named in his honor. His moths included 318 specimens of Geometridae and Noctuidae (Mead 1877). In addition to collecting adult insects, Mead reared butterflies in Colorado and mailed examples of early stages to W. H. Edwards; those not alive were sent in carbolic acid. He even sent larvae home with instructions to his parents on what plants to feed them. Mead also sent home cactus plants, some of which were still alive four years later (17.xi.1875, AMNH). More significantly, he gathered fossils in the area now known as the Florissant Fossil Beds National Monument (Teller County) and is recognized as the first to make fossils from this site available for study (Brown 1981, Veatch & Meyer 2008). He sent insect fossils to W. H. Edwards, who forwarded them to S. H. Scudder for examination. Scudder, an accomplished insect paleontologist, mentioned some of these specimens in his publications (e.g. Scudder 1876) and described a fossil termite in Mead's honor (Scudder 1884). In 1873, Mead sent all his remaining fossils directly to Scudder, “leaves and all” (23.xii.1873, RC). This material is preserved at MCZ.
Departing Colorado via Cheyenne, Wyoming, on 27 September 1871, Mead and his brother arrived in Salt Lake City, Utah, the following day. After spending time in and around Salt Lake City, they proceeded to California on 4 October. After arriving at San Francisco two days later, they toured the city and met the prominent entomologists Hans H. Behr (1818–1904), James Behrens (1824–1898), H. Edwards, and Richard H. Stretch (1837–1926). They took a side trip to Yosemite Valley and returned to San Francisco prior to their scheduled departure of 3 November. Traveling south via streamer along the coast, Mead briefly disembarked on 17 November at Puntarenas, Costa Rica, where he collected a few butterflies. On 20 November the brothers arrived at Panama, where they spent nearly two weeks exploring along the coast. They collected about 1000 butterflies (most hesperiids) by “tramping from 8 to 10 mi a day through swamp & forest” (19.xii.1871, RC). On 3 December they took a train across Panama to the town of Aspinwall (now Colón) on the Caribbean Sea, where they boarded another steamer late that evening. Heading northward, they reached Kingston, Jamaica, on the afternoon of 6 December. They collected butterflies and sampled the local fare in Jamaica until the next afternoon, when they departed for the United States. They finally arrived back home at New York City on 14 December, concluding an arduous, but incredibly gratifying seven-month adventure.
Disposition of Mead's specimens. After Mead's return, he and W. H. Edwards divided up the butterflies from the trip (Edwards 1890), but it is unclear if they followed the plan precisely as Edwards had originally proposed. This took place at Edwards' home in Coalburg, where Mead visited from 28 February to 30 April 1872 (11.V.1872, AMNH). The task was completed during the first week of March and must have taken some time, as Mead had collected nearly 4,000 butterflies (10.iii.1872, AMNH; Mead 1877). “Now I am spending a few weeks with Mr. W. H. Edwards,” Mead wrote his parents, “talking over my captures & having a good time generally” (13.iii.72, RC).
Nearly all of the butterflies that Edwards returned to Mead in 1872 were still in their original field envelopes. For Mead's portion of the take, Edwards decided not to remove them from the papers any more than was necessary to “make sense of the species” (10.vi.1871, RC). Long after he received his share of the take, Mead continued to keep most of his Colorado specimens in papers, which made it easier to exchange them with correspondents. He explained, “In 1871 I collected four thousand specimens of Diurnal Lepidoptera all of which I put up in that manner [papers] finding it safe and convenient, while it would have been almost impossible to transport such a mass of pinned specimens without much risk and trouble” (30.xi.1872, RC). Referring to the papers that he used to store butterflies from Colorado, he recalled, “2 ½ inches square was about the most useful size—accommodating Pieris, Chionobas & etc while [for] Lycaenas 1 ¾ in. would do” (18.xii.1872, RC). He advised recipients of his specimens, “if you lay the papers in a damp towel the butterflies will be relaxed in the course of from 12 to 24 hrs or more according to size” (28.V.1872, RC). Because most of his specimens remained papered, Mead was often “a good deal puzzled” about their identities. He therefore decided to mount one specimen of each unfamiliar species for future comparison. After that, “there was no more trouble” (24.xii.1872, RC). Mead took a particular interest in the Hesperiidae and this was the only group of specimens that he mounted entirely (19.iii.1874, RC).
In an undated letter to his parents, sent from West Virginia during his visit with Edwards in early 1872, Mead recounted his awkward conversation regarding Edwards' financial obligation for the trip. “He said that he was perfectly willing to pay half expenses for the time during which I was butterfly hunting,” Mead wrote, “but intimated that the months during which I was not so engaged didn't count” (RC). Edwards asked Mead to propose an amount which he “thought was right,” to which Mead suggested $400. Despite Edwards' earlier promise to pay up to $500, he was surprised at being “called upon to pay so much.” Nonetheless, Edwards believed the amount was fair and that the butterflies were “fully worth it.” Mead did not intend to ask Edwards for any money (“As it is I feel rather mean”), but the trip had cost more than expected. Edwards was short of funds, but promised to pay the balance within several months. They also agreed that Edwards would offer a “share” of Colorado butterflies to at least two lepidopterists for $100 each. If Mead contributed specimens, he would be given half the amount received.
While Mead was still in Coalburg, Edwards offered Colorado specimens to the lepidopterist F. H. Herman Strecker (1836–1901) of Reading, Pennsylvania. “Mr. Mead & I find that the expense of last summers' collecting in Colorado was very heavy,” Edwards wrote, “More than expected” (23.iv.1872, FMNH). Edwards offered Strecker “a good lot” for a “contribution” of $100 (nearly $2,000 in today's economy). Edwards proposed to make this offer to no more than five lepidopterists, thereby giving each enough specimens “to make a good collection.” Edwards assured Strecker that Mead's papered specimens were mostly in good condition, but admitted that “some of course are not.” He indicated that there were a half a dozen specimens of each species, “frequently 10 or 20,” but in some instances there were “considerably more” (10.v.1872, FMNH). Strecker ultimately declined this proposition and instead purchased only a handful of desirable specimens, a few of which are still contained in his collection at FMNH. In an attempt to cut out the middleman (i.e. W. H. Edwards), Strecker contacted Mead and offered to “expand [mount] all the specimens, have price lists printed and undertake to dispose of' his Colorado specimens for a twenty percent commission (4.ix.1872; MGCL). Mead, however, did not accept this proposal, probably realizing that such an agreement would mostly benefit Strecker in affording him access to the choicest specimens. Although Mead told other correspondents that he was “pretty well supplied with Rocky Mountn and Western rarities for exchange” (24.xii.1872, RC), he insisted that Strecker contact Edwards: “After Mr. Edwards has deposed of his, should there be still a demand I would probably be willing to sell some of mine” (10.ix.1872, RC).
By June 1873, Edwards had contacted four other people whom he hoped would be interested in the specimens (and be able to afford them), but none responded (27.vi.1873, AMNH). Edwards therefore decided to keep the specimens and offer them as “duplicates” to other lepidopterists. Mead was shocked to learn that some of his Colorado specimens were so highly prized: “I had no idea that they were so valuable but in [the] future shall consider the fact in making exchanges” (5.vii.1872, RC).
Although Mead reassured Edwards that he was in no hurry to receive the money owed, Edwards evidently paid the balance by June of 1873, professing, “I live on a salary myself, but I have no cigars to buy or whiskey to indulge in and I save money where some would spend it, and therefore do not feel that I am extravagant in spending what I do in the matter of Butterflies” (27.vi.1873, AMNH). Based on the material at CMNH, Edwards exchanged and/or sold the bulk of his share from Mead's 1871 trip. Edwards subsequently obtained additional Colorado specimens from other collectors, rendering Mead's material less valuable.
Over the years, Edwards and Mead provided Colorado butterflies to many correspondents. Among them was Samuel H. Scudder (1837–1911), who figured several in Scudder (1874). Scudder acquired many specimens during a three-day visit to Edwards' home in December 1873, when he “carried off” all the Hesperiidae he wanted, including several undeseribed species (13.xii.1873, AMNH). Scudder examined Meads collection in New York in late 1873 (19.xi.73, RC) and obtained additional specimens directly from Mead during the early 1870s. Scudder also received some of Mead's specimens via Herbert K. Morrison (1854–1885), a professional collector who exchanged specimens directly with Mead for many years. A number of these specimens are labeled with specific locality data (e.g. “Apex Gulch,” “T. C. Junction,” “Twin Lakes”), which Mead personally conveyed to Morrison (l.iv.1873, RC). Scudder's collection is deposited at MCZ. Morrison's specimens are deposited in many institutional collections, including USNM.
Others who received Mead's Colorado material include Augustus H. Mundt (1847–1920), a jeweler and naturalist of Fairbury, Illinois. These specimens are now deposited in the collection of the Illinois Natural History Survey (Irwin 1966). The California lepidopterists H. Edwards, Hans Behr, and James Behrens obtained specimens from both W. H. Edwards and Mead (Mead also gave them some Colorado specimens when he visited California in late 1871). Henry Edwards' collection is deposited at AMNH. Behr's collection was tragically destroyed two years after his death in the San Francisco earthquake and fire of 1906. Prior to his death, Behrens donated his collection to what is now the Museum für Natur und Umwelt (Museum for Nature and the Environment), Lübeck, Germany. Another San Francisco lepidopterist, William G. Wright (1831–1912), obtained some of Mead–s specimens (probably from W. H. Edwards) and several were illustrated in Wright (1905). Wright–s collection is deposited at CAS.
William H. Edwards exchanged Mead–s specimens with the Massachusetts entomologist Francis G. Sanborn (1838–1884), who in turn forwarded some to the naturalist Charles P. Whitney (1838–1928) of New Hampshire. Beginning in early 1872, Mead exchanged many specimens directly with Whitney: “I have given him already a considerable proportion of the more abundant Col° species in exchange for N.E. [New England] Hespe ri ans etc. not in my collection” (27.v.72, RC). Whitney's collection is now preserved at PMNH (see Calhoun (2013e) for information about the provenance of Whitney's collection).
Mead personally sent Colorado butterflies to the newspaperman and amateur lepidopterist Willard E. Yager (1855–1929) of Oneonta, New York, whose collection may be preserved in the Yager Museum of Art and Culture at Hartwick College in Oneonta. In 1873, Mead exchanged some Colorado butterflies with another New Yorker, the entomologist Joseph A. Lintner (1822–1898), whose collection was bequeathed to the Albany Museum of Natural History, Albany, New York. Mead also sent a few Colorado specimens to the farmer and naturalist George M. Dodge (1846–1912), then of Ohio, Illinois, whose collection is deposited at CAS (Calhoun 2013d). Per his original agreement with Edwards, Mead sent at least one box of butterflies in 1872 to E. T. Cresson of the American Entomological Society: “Today I express to you a box containing … a collection of Col° Diurnals in papers, the joint gift from Mr. Edwards and myself to the Society” (24.v.72, RC). I recently found some of these specimens at The Academy of Natural Sciences (Philadelphia, Pennsylvania) labeled “Edwards and Mead.”
Perhaps the most distant entomologists to obtain Mead's butterflies were the entomologist Jean B. A. D. de Boisduval (1799–1879) of Paris, France, and the natural history dealer Otto Stau dinger (1830–1900) of Dresden, Germany. Edwards sent a few Colorado butterflies to Boisduval in 1873 and 1874 (Edwards' journal “D,” WVSA), while Mead sold some to Staudinger in 1872 (13.iv.1873, RC). Those from Boisduval's collection may be deposited at USNM (see Calhoun 2004, 2006). Staudinger's collection is deposited at the Museum für Naturkunde in Berlin, Germany. Staudinger is credited with sparking Mead's interest in entomology in 1867 (Mead 1935, Calhoun 2013a), though Mead began collecting insects “sporadically” four years earlier (21.v.1874, RC). Untold numbers of butterflies changed hands between entomologists of the 19th century, scattering Mead's specimens to countless collections, though many are no longer recognizable as his.
It was fortunate that Mead was not a member of the Wheeler Survey of 1871, as many of the specimens brought back from explorations that year were accumulated and stored in Chicago, Illinois. On 8–10 October 1871, the Great Chicago Fire laid waste to much of that city, taking with it specimens from the survey (Yarrow [1876a]). The collections of Mead and Edwards were ultimately purchased by William J. Holland (1848–1932), who later served as the Director of the Carnegie Museum (CMNH), where his large personal collection is now preserved. In his autobiography, Mead (1935) erroneously dated the purchase of his collection as 1877; it was actually 1884. Apparently, Mead's collection had suffered from neglect and many specimens were in poor condition. “Mead had a great many very valuable things of his own collecting …” Edwards remarked, “I hope these may not have suffered,” (16.xi.1884, CMNH). This problem is evident in the condition of some of Mead's specimens at CMNH (Brown 1970). Holland paid $460 for Mead's entire collection (7.vi.1884, RC). He purchased Edwards' collection in 1885–1886 for $2,500 (Calhoun 2013b).
Mead's Report. William H. Edwards encouraged Mead in late 1871 to write up his observations on Colorado butterflies. Edwards even offered to publish them in his book, The Butterflies of North America, as a “highly interesting record to the present generation” (15.xi.1871, MGCL). Alternatively, Edwards suggested that Mead write a lengthy story about his experiences and publish them as “a magazine article.” Mead did neither, but he was asked three years later, at the suggestion of Edwards, to write the section on diurnal Lepidoptera for the zoology report of the Wheeler Survey (Calhoun 2013c). Mead accepted the responsibility of drafting the report on diurnal Lepidoptera while he was a student at Cornell University in Ithaca, New York. “I will be glad to prepare a report on the insects,” Mead wrote in reply, “but I would like to know how voluminous an account is desired” (29.i.1874, RC). He suggested a more detailed treatise that would result in a “sizable pamphlet— probably 60 or 70 pp.” Mead wasted little time in sharing this distinguished honor with his closest correspondents. He was astonished by the official acknowledgement he received, conveying the thanks of Lt. Wheeler, which was addressed to “Prof. T. L. Mead.” Mead quipped to his brother, “[I] wonder what he would say if he knew I was nothing but a miserable little Freshman …” (4.ii.1874, RC). He was astounded that an inexperienced college freshman would be entrusted with such an important task. Thus was the influence of W. H. Edwards and his admiration of Mead.
For the report, Mead combined the results of his own collecting in Colorado with those of Wheeler Surveys conducted from 1871 to 1874 in portions of southern Utah, New Mexico, and Arizona. As a consultant to the Wheeler Survey, Edwards contributed a list of all the butterfly species that were collected during the various Wheeler expeditions. Mead also included data from the Allen Expedition of 1871. Joel A. Allen (1838–1921), an assistant at the Museum of Comparative Zoology, spent nine months during 1871 with two assistants exploring portions of Kansas, Colorado, Wyoming, and Utah (Osborn 1916, Brown 1956). Mead met the Allen party near Montgomery City, Colorado, on 24 July 1871. He later examined Allen's butterfly specimens at MCZ (26.vi.1872, RC), some of which are still preserved in that collection, labeled “Colorado. / Allen.” William H. Edwards was probably referring to Allen's expedition when he notified Mead that some of the same species had been brought back by other collectors in 1871 and “Many are at Cambridge” (15.xi.1871, MGCL). Mead was unfamiliar with some of the species collected in Utah and other areas outside Colorado, thus he asked Henry Edwards to provide notes on those species (8.iii.1874, AMNH) (he thanked Edwards in the published report for “notes upon some species whose habits were unknown to me”). Due to extreme delays, Mead's Report was not published until late 1876 (Calhoun 2013c). It served as the basis of the suggested type localities by Brown (1934) and the primary source of information about Mead's Colorado itinerary as charted by Brown (1955a).
Mead started to lose interest in entomology around 1877. By 1882, when he permanently moved to Florida, his lifelong interest in horticulture had become his passion. He left his Lepidoptera collection with his parents in New York, and it was from there in 1884 that W. J. Holland transported it to his home in Pittsburgh, Pennsylvania. Although Mead continued to maintain a passing interest in Lepidoptera for many years (Skinner 1922), he became infatuated with the hybridization of flowers, especially orchids and Amaryllis (Mead 1904, Mead 1935, Butler 2013, 2014). Census records from the early 20th century list him as a “florist” and “bulb farmer.” Mead suffered through the devastating deaths of his young daughter, Dorothy, in 1892 and his wife, Edith, in 1927. Mead died of a stroke on 4 May 1936 at Memorial Hospital in Sanford, Florida, and was buried in Greenwood Cemetery (section B)—to this day a serene oasis nestled within the bustling tourist mecca of Orlando, Florida.
Mead's manuscripts and 1871 journal. In his last will and testament, dated 19 August 1933 (RC), Mead bequeathed to the Trustees of Rollins College all of his “scientific books and pamphlets and apparatus,” as well as his “album of Colored Orchid Photographs.” With the exception of a few miscellaneous items, he bestowed “all the rest and residue and remainder” of his property, “real personal and mixed,” to his nieces Catherine T. Willis and Eleanor deGruyter of Charleston, West Virginia. Catherine (1884–1968), who was married to John A. Willis, Sr. (1877–1961), was the daughter of Anne S. Smith (1858–1930), the youngest daughter of W. H. Edwards and the sister of Mead's wife. For many years the Willis' lived in Edwards' former home in Coalburg, West Virginia (Calhoun 2013b). They received some of Mead's property, including many of his letters (a portion of which are preserved at MGCL). None of Mead’s journals were listed among the items originally received by Rollins in 1936.
Although Rollins College received the bulk of Mead's manuscripts after his death, some material remained the property of Mead's friend, John “Jack” H. Connery (1908–1982), of Winter Park, Florida. For a time, Mead was a scoutmaster and Connery was one of his Eagle Scouts. Connery attended Rollins College, where he taught ornithology classes and served as the director and curator of the Thomas R. Baker Museum. During the early 1930s, he was a photographer for the Bermuda expeditions of the famed zoologist William Beebe (Gould 2004). Documents at Rollins College indicate that upon Meads death, Connery took charge of Meads property, much of which was stored in a barn, presumably on Mead's property in Oviedo, Florida. Some of Mead's belongings were retained for a planned Mead memorial that later evolved into the Theodore L. Mead Botanical Garden in Winter Park. Dedicated in 1940, Connery was instrumental in the development of this property, which operates today as Mead Botanical Garden (Whitman 1939, Johnson 2008).
Records show that Connery received books from Mead's library, as well as “a large quantity” of his personal letters, manuscripts, photographs and many other items of “great historical interest.” This is supported by a letter from Mead to Connery and his wife, Helen, dated 30 May 1933, three months before Mead drafted his will. Mead permitted them to have all the correspondence they wished, advising, “I don't believe anybody else would care for the diaries so you can have them when I have done with them” (RC). Mead mentioned the presence of “1877 diaries and later,” thus revealing that other journals existed and at least some were likely received by the Connerys. Beginning around 1947, the majority of the manuscripts given to the Connerys were kept in a large trunk in a storeroom of Edwin O. Grover (1870–1965), a former Director of Libraries at Rollins College who later served as Vice President of the school until his retirement in 1947. In 1957, Grover and the Connery's asked that the trunk of Mead's manuscripts be placed on deposit with the remainder of Mead's manuscript collection at the Mills Memorial Library, which opened at Rollins College in 1951. The Mead manuscript collection was moved several times after arriving at Rollins and is now preserved in the Olin Library, which opened in 1985. Other Mead-related materials and documents are deposited at the John C. Hitt Library, University of Central Florida (Orlando, Florida), the Kroch Library, Cornell University (Ithaca, New York), Mead Botanical Garden (Winter Park, Florida), the Orange County Regional History Center (Orlando, Florida), and the Winter Park Public Library. John Connery's sons, John, Jr. and W. Edwin, also possess some items, including a few of Mead's orchid hybridization notebooks and a painted portrait of Mead (W. E. Connery, pers. comm.).
In 2010, I was extremely fortunate to acquire Mead's 1871 journal from James W. Tillery of Lehigh Acres, Lee County, Florida (Calhoun 2010) (Figs. 2, 3). Tillery discovered it among items in a storage unit, which had been rented by a local bookseller. The bookseller passed away and the unit's contents were auctioned. An obvious connection to Lee County could be made through J. H. Connery's youngest son, W. Edwin Connery of Cape Coral, Florida (whose middle name honors Edwin O. Grover of Rollins College). I discussed Mead and his manuscripts with W. E. Connery via telephone in April 2013. He did not recall ever seeing any of Mead's personal journals among his parents' belongings when they died. He suggested that the 1871 journal became separated from the other manuscripts many years ago and its subsequent presence in that region of Florida was merely a coincidence. Some of Mead's remaining yearly journals were seen by Paul Butler during a recent visit the former home of W. H. Edwards in Coalburg, West Virginia, though none are from the 1870s (P. Butler, pers. comm.). Butler is currently preparing a more lengthy biography of Mead for publication.
Preserved at Rollins College are five volumes of letter copybooks in which Mead duplicated the letters he wrote during the early 1870s. Within these bound volumes are letters written during 1 January–29 December 1870, 23 October–30 December 1871, 4 January–31 December 1872, 6 January–31 December 1873, and 1 January–20 March 1874. Unfortunately, the volume that would have included letters written during 1 January–22 October 1871 (covering Mead's time in Colorado) is missing. Among the F. M. Brown archives in the AMNH is information about this copybook, as well as some photocopied pages and a draft manuscript about Mead's Colorado trip. I obtained scans of these documents, which offer some insight into the fate of the missing volume.
In 1979, F. Martin Brown and the late lepidopterist Lee D. Miller (1935–2008) visited the Mills Memorial Library at Rollins College, where they examined some of Meads' manuscripts, including his letter copybooks. The reference librarian photocopied a portion of the 1871 copybook at Brown's request. In 1981, Brown requested additional photocopies of letters that were illegible in the original set. On 5 September 1984, Brown announced that he had finally completed the transcriptions of the letters in the copybook, yet some passages remained undecipherable. At that point, the librarian was unable to locate some of the materials, pointing out that the letters “apparently have been shifted.” Brown reminded her that the copybook was “in an old trunk (box) down in the storeroom,” referring to the trunk of Mead’s manuscripts that the Connerys had previously donated to the college. It seems likely that the copybook was misplaced shortly after photocopies were sent to Brown in 1981. It probably lacked external identifying marks, such as Mead's name or a catalog number, increasing the chance that once separated it was not properly returned. Mead's manuscripts have since been thoroughly cataloged and archived. Although Brown's photocopy of Mead's 1871 volume is incomplete and many pages are illegible, it offers a valuable glimpse into its contents, especially when combined with the transcriptions by Brown & Brown (1996).
Updated Colorado Itinerary. Based upon Mead's 1871 journal, I offer a corrected itinerary of Mead's exploration of Colorado (Table 1). The map included in Brown (1955a) is still a helpful reference to the locations visited by Mead, but some of the dates and estimated routes are incorrect. Three locations where Mead spent a great deal of time in Colorado serve as the type localities for multiple taxa described from his specimens. Kenosha House and Turkey Creek Junction were 19th century stagecoach stops in Park and Jefferson Counties, respectively. The third, Twin Lakes, is located in southern Lake County.
Kenosha House. Described by Mead as “a clean well-kept ranch,” Kenosha House was a two-story stagecoach station constructed in 1861 (Fig. 4). Offering meals and lodging to weary travelers, the proprietors advertised “an excellent variety of tempting food,” including venison, wild duck, and mountain strawberries (Anonymous 1870, Crump & Crump 2010). Kenosha House also had mail service, which allowed Mead to send letters to W. H. Edwards and others while lodging there. Mead often referred to the station as “Kenosha Ranch.”
Disagreement exists regarding the location of Kenosha House, with some authors (e.g. Fisher 2006) placing it at the summit of Kenosha Pass, Park County. Scott et al. (2006) positioned it 0.8 km (0.5 mi) north of the pass. Although this location is consistent with the map of Wheeler (1876) and the trail map of G. Scott (1999), it does not agree with other accounts, including the map of Hayden (1877b), which shows it about 2.16 km (1.34 mi) (2.62 km/1.6 mi by road) north of the summit (Fig. 5). This is roughly where Brown (1955a) mapped it. Warren (1994) also located it there, directly opposite the current intersection of U.S. Hwy 285 and C.R. 58 (Lininger Lake Rd.), near the point where Hoosier Creek turns northward. This site corresponds with the records of the Colorado Office of Archaeology and Historic Preservation (S. Gilmor, pers. comm.). Mead () stated that Kenosha House was located “four miles from the South Park,” which is a relatively accurate distance from this site to the South Park basin along the old wagon road. This is consistent with a statement that the best view of the South Park basin could be had “about three miles beyond the Kenosha House” (Anonymous 1870).
The house stood on the east side of the Denver, Bradford & Blue River Road. Also known as the Denver, Turkey Creek & South Park Wagon Road, this trail was commonly known as the Denver & South Park Road (Hayden 1874); cumbersome names shorted by Mead to “South Park Road.” The house was said to overlook a “beautiful valley that slopes up toward the mountains” (Anonymous 1870), clearly a reference to the valley of Hoosier Creek (Fig. 5). According to his journal, Mead collected butterflies as far as 4.8 km (3 mi) within the gulch “opposite the house” (Hoosier Creek) and along the wagon road toward South Park. He also explored “up a little brook” to a “pine lake.” These are today’s Lininger Ditch and Lininger Lake, located just northwest of the former site of Kenosha House (Fig. 5).
Gannett (1877) recorded the elevation of Kenosha House as 2935 m (9629 ft.), but current maps suggest it was closer to 2957 m (9700 ft.). GPS: 39.432627, — 105.758828. This area serves as the type locality of the nominal taxa Colias hagenii W. H. Edwards, Theda ninus W. H. Edwards, and Melitaea eurytion Mead.
Turkey Creek Junction. There also is disagreement about the location of this settlement. Brown (1934) initially associated it with “the junction of South Turkey Creek with Turkey Creek,” but later equated it with Junction House (=Bradford Junction), formerly located in today's town of Conifer in Jefferson County (Brown 1955a). Scott et al. (1998) considered Turkey Creek Junction to be 0.8 km (0.5 mi) north of Tiny Town, a miniature village about 1.6 km (1 mi) southeast of the community of Indian Hills along South Turkey Creek Road. Fisher (2006) suggested that Turkey Creek Junction was probably “in the vicinity of what we call Tinytown … or between there and what is currently called Myers Ranch Open Space.” The latter is now known as Meyer Ranch Park, located just east of the town of Conifer.
Distances recorded by Mead in his journal confirm that what he called “Turkey Creek Junction” was Bradford Junction, a community once situated 10.6 km (6.6 mi) southwest of Tiny Town and about 366 m (1200 ft.) higher in elevation. Mead later recalled that “Turkey Creek Junction” was “a very good collecting ground 28 miles from Denver on the S.P [South Park] road” (l.iv.1873, RC). Identified as “Junction” on contemporary maps (e.g. Hayden 1877a, Wheeler 1877) (Fig. 7), this small settlement and stagecoach stop was established by Robert B. Bradford, who owned the property from 1860 to 1873. It was located at the junction of two main roads; the Mt. Vernon Wagon Road (which Mead called the “North Road”) and the Denver & South Park Road (which Mead called the “South Park Road”). Bradford Junction existed where present-day Barkley Road intersects County Road 73 within the town of Conifer. Bradford Junction was the basis of Conifer (Donovan 1995), though current maps erroneously suggest that Conifer is a small community south of Aspen Park along US Hwy 285. In reality, Aspen Park is merely a developed area within Conifer, not a separate town. The historic Yellow Bam now stands at the former site of Bradford Junction (Bentley 1985, Donovan 1995, Hood 2011).
Itinerary of T. L. Mead in Colorado
Among several buildings at Bradford Junction was the large two-story Junction House, which offered food and lodging, though, according to Mead, some of its beds were infested with bedbugs: “[A]t night it entirely ceases to be a labor of love and becomes something quite different” (he pinned six bedbugs to his pillow one night) (Brown & Brown 1996). The house stood just southeast of the present-day Yellow Barn. A covered well, dug in 1862 and still present on the property, is visible in the middle of the road in front of the Junction House in a photograph taken ea. 1875 (Fig. 6). The surroundings were described as “very interesting, the forest delightful, the shrubbery voluptuous with foliage and flowers, the way often bordered with wild roses, perfuming the cool, bracing air” (Anonymous 1868). The house burned in 1878 and was rebuilt about five years later using a different design (Hood 2011).
The Junction House served as Mead's base of operations when he explored the region, which he generally referred to as “Turkey Creek.” Upon his first arrival at Bradford Junction on 5 June, Mead called it “the dinner station,” as he did not intend to spend any time there. He walked ahead along the wrong road while collecting butterflies and missed the stage, stranding himself for two days until the stage returned. This was a very fortuitous event, as his success collecting there prompted him to revisit the area two more times, leading to the discovery of several new butterfly taxa. Mead hunted butterflies as far as 6.4 km (4 mi) northward along the Mt. Vernon Wagon Road (today's CR 73) and 12.9 km (8 mi) northeastward along the Denver & South Park Road (today's South Turkey Creek Road), paralleling South Turkey Creek (then known as Turkey Creek). He also explored along the gulch “opposite the house,” which is the head of North Turkey Creek (Shadow Mountain Drive now parallels the creek where an older road once existed). Also at Bradford Junction was a post office, from which Mead sent letters and specimens during his trip.
It is significant that I have found no other allusions to the name “Turkey Creek Junction” beyond those of Mead. The place names directory of the Jefferson County Historical Commission (Jeffco 2013) has no listing for this name, nor does the state historical society, History Colorado. Members of the local Conifer Historical Society are also unfamiliar with this name. The settlement was identified as “Junction” on period maps and stage schedules (Fig. 7). The most reasonable explanation is that Mead added the appellation “Turkey Creek” to geographically restrict his references (i.e. Junction within Turkey Creek Canyon) to prevent its confusion with other stage stops in Colorado also known as “Junctions.” When citing Mead's data, it is more geographically accurate to refer to this locality as Bradford Junction.
Gannett (1877) recorded the elevation of Junction House as 2485 m (8153 ft.), but it is actually closer to 2454 m (8050 ft.). GPS: 39.534153, -105,309489 (Junction House). This area serves as the type locality of the nominal taxa Anthocaris [sic] Coloradensis H. Edwards, Lycaena daunia W. H. Edwards, Phyciodes nycteis var. drusius W. H. Edwards, and Melitaea calydon Holland. The original type (now lost) of Lycaena alce W. H. Edwards was also from this vicinity.
Twin Lakes. A region with two large glacial lakes south of Leadville, Lake County (Fig. 8). Now a reservoir, the lakes were described in 1870 as “about two miles in width, and five miles in length, separated by a strip of forest land, about one-fourth mile in breadth” (Wallihan & Bigney 1870). Immediately surrounding the lakes were “clear, hard, sandy beaches,” alternating with “walls of rock and low marshy meadows” (Bowles 1869). The western lake is known as Upper Twin Lake, while the eastern lake is Lower Twin Lake. Damming the lower lake during the late 19th century raised water levels by as much as 4.6 m (15 ft.), obliterating a large portion of the vegetation that grew around the lakes when Mead visited (Noel & Fielder 2001).
Mead's journal entries and mileage estimates indicate that he lodged in a “house” on the east side of Lower Twin Lake, west of the Arkansas River. Described as a “roadside ranch” by Mead, the lodging house was most likely owned by Samuel M. Derry (e. 1817-?), who also operated a hotel in the former community of Dayton (now Twin Lakes). Derry claimed ownership of much of the land surrounding the lakes (Anonymous 1908, Jackson & Driggs 1929) and Mead mentioned in his journal on 9 July that his brother went fishing for trout with “Mr. Derry.” Mead remarked that his nearest neighbors at Twin Lakes were six miles away by road (Brown & Brown 1996), which presumably refers to Dayton, located about 9.7 km (6 mi) by road west of the point where Lake Creek crossed the old wagon road on the east side of Lower Twin Lake (Fig. 8). Mead explored much of the area around the lakes, including over 9.7 km (6 mi) westward along Lake Creek (west of the lakes) and southward about 1.6 km (1 mi) along the road leading to the town of Granite, southeast of the lakes.
Mead also reached the peak of a mountain at an elevation of 1219–1524 m (4000–5000 ft.) “above [the level of] the lakes.” Brown (1955a) suggested this was La Plata Peak, or possibly Mt. Elbert. Brown and Brown (1996) later proposed Quail Mountain. However, Meads written comments suggest that he ascended an unnamed lower peak northeast of Quail Mountain, just south of the lakes. Without taking provisions, he started out on 10 July to simply “look around,” but “walked a long distance up a gulch and finally concluded to go to the top of a peak,” returning at “about suppertime.” He most likely walked up Flume Creek, reaching an elevation of about 3840 m (12,600 ft.) (Fig. 8). Two days later, Mead and his brother unsuccessfully attempted to climb a steep peak “not quite so high” located just southeast of the first peak.
Elevation of the lakes approx. 2804 m (9200 ft.). GPS: 39.077576, -106.291290 (vicinity of Mead's likely lodging site). This area serves as the type locality of the nominal taxa Thymeticus [sic] hylax W. H. Edwards, Hesperia colorado Scudder, Pamphila draco W. H. Edwards, Chrysophanus sirius W. H. Edwards, Lycaena melissa W. H. Edwards, Argynnis electa W. H. Edwards, and Satyrus charon W. H. Edwards.
Type localities. Most of the Colorado specimens in Meads collection were contained in field envelopes and it appears that they were purchased in this condition by W. J. Holland (Brown 1964, 1970). When these specimens were later mounted by Holland, that portion of Meads envelope with his handwritten date was clipped out and affixed to the pin. Many of the specimens from Mead's collection at CMNH bear such clippings and this information is extremely valuable in determining collecting localities. Samuel H. Scudder also received Mead's butterflies in papers and, like Holland, affixed a portion of Mead's envelope onto the mounted specimens. Some of Mead's specimens at CMNH and MCZ possess small identification labels in Mead's hand (Fig. 17), suggesting they were already mounted when they were acquired by Holland and Scudder. “Generally I write my labels as printed ones are not always to be had,” Mead remarked in August 1871 (Brown & Brown 1996).
Twenty-eight taxa of butterflies were described from specimens that Mead collected in Colorado in 1871. One additional taxon is represented by a neotype that was collected by Mead. Nineteen of these nominal taxa possess incorrect or imprecise type localities that were suggested by previous authors. Presented below are clarifications of these type localities based on Mead's whereabouts in accordance with Article 76 of ICZN (1999), including Recommendation 76A. Current nomenclature follows Pelham (2014).
Pamphila colorado Scudder, 1874
(=Hesperia colorado; Hesperiidae)
Described from “Colorado, about the Georgetown and South Park Roads [Clear Creek and Park Cos.],” which Mead () reiterated. The lectotype (MCZ) (Fig. 9) designated by Barnes and McDunnough (1916) is labeled “Colorado.” Scott (1998) suggested a type locality of “Tennessee pass, 3150 m (=10300 feet), Lake-Eagle Cos. Colorado.” Andrew D. Warren proposed Guanella Pass (Park/Clear Creek Cos.) (Pelham 2008), but maps do not show a trail through that area during the 1870s. Scott (2008b, 2008c) stated “evidently Kenosha Pass [Park Co.] or Guanella Pass [Clear Creek Co.], Colo.”
Affixed to the lectotype is a portion of Mead's original field envelope with the penciled date of “7–13.” This specimen was figured by Scudder (1874, Pl. 10, fig. 18) as “collected July 13, by T. L. Mead.” A damaged male paralectotype at MCZ with the same date as the lectotype, was dissected and figured by Scudder (1874, Pl. 11, figs. 10, 11). Other butterflies at MCZ that Mead collected during mid-July 1871 bear dates written in his hand in either pencil or ink using the same format. Scudder (1874, Pl. 10, figs. 16, 17) also figured as colorado two females collected by Mead, one of which is dated 28 August. All these specimens were possibly papered duplicates that Scudder obtained when he visited Edwards' home in December 1873. Most of Mead's specimens of Hesperia from Colorado were examined by Scudder, who identified some as Pamphila manitoba (=Hesperia comma manitoba (Scudder)) (Scudder 1874). This undoubtedly prompted Mead () to apply this name to two of his specimens, which represent high-elevation phenotypes of the Front Range subspecies Hesperia colorado ochracea Lindsey.
On 13 July 1871 Mead was at Twin Lakes, where he walked “up to the head of the upper lake” to collect butterflies. All the material from Twin Lakes was mailed to Edwards from Oro City on 20 July 1871. The female specimen that Mead captured on 28 August was taken along the Denver & South Park Road northeast of Kenosha House in northern Park County (Table 1), probably at an elevation of about 2621 m (8600 ft.). Mead () mentioned collecting colorado along this road “during the latter part of August.” Although the peak flight period of this butterfly is mid-August to early September, adults can emerge much earlier, especially during unusual conditions. Precipitation in Colorado during the spring of 1871 was below modern norms (Mock 1991). In fact, much of the United States was already suffering drought conditions by early spring. One of the worst natural disasters in history took place during October 1871 when fire storms swept across the Great Lakes region, causing the Great Chicago Fire (van den Dool 2012). Other captures by Mead in 1871 suggest that some other butterfly flight periods were advanced (see C. hagenii and A. helena, below).
To identify the lectotype (MCZ-ENT00015299) as the primary type of colorado, a red printed and handwritten label was affixed to its pin, which reads, “LECTOTYPE / Pamphila colorado / Scudder, 1874 / per Barnes & McDunnough (1916) / John V. Calhoun, 2013.”
Type locality. As defined by the lectotype, the type locality is restricted to Twin Lakes, along the northern shore of Upper Twin Lake (the western lake), Lake County, Colorado.
Discussion. Skinner and Williams (1924) observed that “Individuals from high altitudes are darker green below than the type.” Referring to Scudder's (1874) figure of the lectotype, Lindsey (1942) suspected that it “may very well be true colorado.” Indeed, the lectotype (Fig. 9) and paralectotype from Twin Lakes are more brightly colored than higher-elevation butterflies (Fig. 12), which have long been considered to represent nominotypical colorado. Many authors, including MacNeill (1975), Stanford (1981), and Tilden and Smith (1986), considered nominotypical colorado to denote high-elevation populations of small dark butterflies. The lectotype, however, is more evocative of butterflies that are currently recognized as the subspecies Hesperia colorado oroplata (Fig. 11), described as H. comma oroplata by Scott (1981) to define more brightly-colored lower-elevation populations “from the Arkansas River Valley south of Buena Vista to the Royal Gorge, south to the Sangre de Cristo Mts. of New Mexico, and the San Luis Valley of Colorado.” Unfortunately, Scott did not attempt to locate type specimens of Pamphila colorado (Scott 1998).
To establish a baseline of comparison from which to differentiate oroplata, and to prevent the use of the name colorado for populations at lower elevations, Scott (1998) suggested Tennessee Pass (3170 m/10,400 ft.) as the type locality of colorado, despite a lack of evidence that Mead had collected there. Without further explanation, Scott (2008b, 2008c) later revised the type locality to “evidently Kenosha Pass or Guanella Pass,” which are 3048 m (10,000 ft.) and 3557 m (11,670 ft.), respectively. Scott (1998) considered “true” colorado to occur “at high altitude in Colorado in mountains along the continental divide … from about 10,000 feet upward,” adding that “the darkest populations are at the highest altitudes of up to 12,400 feet or more.” As an example of this taxon, Scott (1998, fig. 15) figured a male from Loveland Pass, Summit Co., Colorado, 3779.5 m (12,400 ft.) in elevation. Although this approach reflected the popular concept of this taxon, it is not supported by the lectotype. Thanks to Mead's journal, we can now establish with certainty that the lectotype of colorado was collected at a lower elevation.
The primary character used by Scott (1981) to distinguish oroplata was the “lighter yellowish rather than greenish brown” coloration of the ventral hindwings (Scott 1981). Additional cited attributes were “slight differences in genitalia, antenna length, number of micropyle spines & developmental period.” This analysis, however, was based on the notion that colorado is restricted to higher elevations; populations that Scott (1975a, 1975b, 1986) considered to be genetically discrete. Scott's interpretation of colorado is inconsistent with the lectotype, which agrees with the original description of oroplata in that the ventral hindwings are “lighter yellowish,” not greenish brown like those from higher elevations (Figs. 9, 11, 12). Scudder's (1874) figure of the lectotype faithfully portrays its more brightly-colored ventral hindwings (Fig. 10). Moreover, Scott (2006b) remarked that colorado is “evidently biennial like other alpine butterflies.” This differs from oroplata and populations of “true” colorado, both of which are annually-brooded. Forewing lengths (base to apex) of the holotype of oroplata and the lectotype of colorado measure 15.0 and 14.4 mm, respectively.
The lectotype of colorado was collected at an elevation of roughly 2819 m (9250 ft.), within an area that Scott (1975) identified as a transition zone between oroplata and higher-elevation populations, which he and other authors attributed to nominotypical colorado. Fifty-six specimens that Scott collected within this area are deposited at MGCL (ex. W. W. McGuire colln.). They were captured 2.6 km (1.6 mi) north of the town of Granite, Chaffee County (2774 m/9100 ft.), and at Mt. Massive Trout Club, Lake County (2856 m/9400 ft.). These populations occur within 9.7 km (6 mi) of the type locality and represent typical colorado as defined by the lectotype. Any perceived differences between these populations and those within the range of oroplata are subtle at best. They form a smooth gradual cline, with populations at lower elevations producing slightly larger and tawnier adults with somewhat reduced dark maculation. Because the primary criteria that were originally used to differentiate oroplata also extend to the nominotypical subspecies, all these populations should be considered to represent H. c. colorado. As a result, the dusky biennial populations at higher elevations in Colorado require a new name (Warren & Calhoun 2015).
The female that Mead collected on 28 August, which was figured by Scudder (1874, Pl. 10, fig. 17), is applicable to H. c. ochracea. The second female figured as colorado by Scudder (1874, Pl. 10, fig. 16) actually represents Hesperia nevada (Scudder).
Pamphila nevada Scudder, 1874
(=Hesperia nevada; Hesperiidae)
In addition to specimens from Nevada, California and Oregon, this species was described from “Colorado … on the mountains about the South park and in the Park itself [Park Co.],” which Mead () reiterated. Barnes and McDunnough (1916) designated one of Mead's specimens as the lectotype and defined the type locality as “S. Park, Colo.” Affixed to the lectotype (MCZ) is a portion of Mead's original field envelope dated “June 12.” This specimen was figured by Scudder (1874, Pl. 10, fig. 1) as “collected June 12, by T. L. Mead.” Scudder (1874, Pl. 10, figs. 3, 4) also figured male and female paralectotypes, dated 17 June and 23 June, respectively. A male paralectotype at CMNH was collected by Mead on 16 June (it is from Mead's collection, but Scudder examined all of Mead's Hesperia prior to describing this species). Colorado specimens of nevada from Scudder's collection were possibly papered duplicates that he obtained from Edwards in 1873.
Another male paralectotype at MCZ bears a portion of Mead's field envelope dated “6/17.” Its claspers were figured by Scudder (1874, Pl. 11, figs. 3, 4), who stated that the specimen was “collected June 17, by T. L. Mead.” This specimen also bears a second clipping from Mead's envelope reading “Mr. B.” Mead's journal entry for that date indicates that Samuel M. Blair collected 70 butterflies. Blair (1842–1932), a miner and relative of the proprietor of Kenosha House, assisted Mead and W. H. Edwards in collecting and rearing butterflies from the area. “Mr. Blair is, I think, to be trusted to take good care of the insects and to collect more,” Mead reported in July 1871 (Brown & Brown 1996). Mead initially took ownership of Blair's butterflies, but Blair later accepted an offer from Edwards to collect specimens in exchange for parts of his butterfly book. Blair's specimens were thereafter considered to be Edwards' property, though Mead saw this as an unfair arrangement, especially since he physically managed Blair's activities in Colorado. Mead eventually became dissatisfied with Blair, complaining that he lacked “that extreme accuracy necessary for scientific observations” (Brown & Brown 1996).
On 12 June 1871, Mead collected several miles up Beaver Creek near Fairplay (Table 1). On 16 and 17 June, he walked south from Kenosha House along the Denver & South Park Road. On 23 June, he collected along the same road just southwest of Bradford Junction (Table 1). Mead mailed the lectotype specimen to Edwards from Bradford Junction on 20 June 1871.
To identify the lectotype specimen (MCZ-ENT00015299) as the primary type of nevada, a red printed and handwritten label was affixed to its pin, which reads, “LECTOTYPE / Pamphila nevada / Scudder 1874 / per Barnes & McDunnough (1916) / John V. Calhoun, 2013.”
Type locality. As defined by the lectotype, the type locality is restricted to along Beaver Creek, east/northeast of Fairplay, Park County, Colorado.
Hesperia dacotah W. H. Edwards, 1871
(=Polites mystic dacotah; Hesperiidae)
Described from a single male collected by Mead in “Colorado,” which Brown and Miller (1980) defined as “Georgetown, Clear Creek County, Colorado.” Scott (2008b) suggested “Georgetown or eastward toward Idaho Springs.” Although the holotype (CMNH) does not possess a date, it bears an identification label of W. H. Edwards reading “napa ♂ = dacotah.”
Soon after describing dacotah, Edwards (1872) questioned its recognition and concluded it was synonymous with the species he had previously described as Hesperia napa (=Ochlodes sylvanoides napa). Based on this treatment, Mead () did not separately list any records of dacotah from Colorado, nor did Edwards () list it among the species recorded by the Wheeler Survey. These taxa were not again recognized as different species until Barnes & McDunnough (1916) treated dacotah as a variety of P. mystic.
Regarding napa (which was then considered to include dacotah), Mead () stated that he “found a few specimens in August near Georgetown.” One of Mead's specimens of napa at CMNH is dated “Aug 19” (see below). On 19 August, Mead was traveling from Idaho Springs — where he had arrived the previous day from Georgetown — to Apex Gulch (Table 1). Mead () also mentioned the capture of another species at Idaho Springs on 19 August. These specimens were mailed to Edwards from Denver on 22 August 1871.
Type locality. Restricted to the vicinity of Idaho Springs, Clear Creek County, Colorado.
Hesperia napa W. H. Edwards, 1865
(=Ochlodes sylvanoides napa; Hesperiidae)
Described from specimens collected in 1864 by James Ridings at “Empire City [Clear Creek Co.], Colorado Territory.” One of Mead's specimens (CMNH) was designated as the neotype by Brown and Miller (1980), who implied the type locality to be “the vicinity of Georgetown, Clear Creek County, Colorado.” However, Miller and Brown (1981) and Pelham (2008, 2014) reiterated the type locality as “Empire City, Colorado,” and “Empire in Clear Creek County, Colorado,” respectively. Mead () stated that he “found a few specimens in August near Georgetown.”
Affixed to one of Mead's specimens of napa from the Edwards collection (CMNH) is a portion of his original field envelope, which reads “Hesperia A / Aug 19.” In a letter to Edwards, Mead mentioned taking “Hesp A” on 19 August at Idaho Springs (Brown & Brown 1996). On 19 August, Mead was traveling from Idaho Springs to Apex Gulch (Table 1). These specimens were mailed to Edwards from Denver on 22 August 1871.
Type locality. Restricted to the vicinity of Idaho Springs, Clear Creek County, Colorado.
Coloradensis H. Edwards, 1881
(=Euchloe ausonides Coloradensis; Pieridae)
Described as a possible new species from “Colorado,” which Opler (1966) defined as “Turkey Creek Junction, Jefferson County, Colorado.” This type locality was proposed on the advice of F. Martin Brown, who examined the male syntype and concluded that it is “without doubt one sent to Henry Edwards by W. H. Edwards from the material collected by T. L. Mead in 1871.” Mead wrote of finding early stages of this species at Turkey Creek Junction in June 1871 (Edwards 1874). He later told H. Edwards, “only one of my [larval] specimens gave imago and it proved to be Ausonides …” (4.iv.1874, AMNH).
Two specimens (male and female) from H. Edwards' collection, acquired by AMNH in 1892, possess his handwritten labels identifying them as “types” of Coloradensis. The male syntype was designated as the lectotype by Johnson (1976). Henry Edwards received one of Mead's specimens of this butterfly from W. H. Edwards in early July 1871. In the accompanying letter, W. H. Edwards asked if “Anthocaris 1” was ausonides; in agreement, H. Edwards scrawled that name on the page (8–9.vii.1871, AMNH). It could be assumed that it was this specimen to which Edwards (1881) referred when he wrote, “I have long had in my possession a ♂ example of Anthocaris, which appears to be distinct from any known form, but I hesitated to describe it until more material might present itself.” However, Mead sent at least two additional Colorado specimens of ausonides to Edwards in 1872 (19.vi.1872, RC, AMNH). Edwards (1881) did not attribute the male type specimen to Mead and only casually mentioned that Mead's collection also contained this species. Henry Edwards evidently did not recall where he had obtained the male type specimen, which was received years before he described Coloradensis. He exchanged hundreds, if not thousands, of butterflies after 1871 and it is possible that the specimen he mentioned in the original description was received from another correspondent at a later date.
Edwards' collection catalog at AMNH does not list any specimens by the name Coloradensis. Entry no. 14 lists “Anthocaris ausonides” from California. Although Opler (1966) indicated that the lectotype of Coloradensis was labeled “14,” no such label or notation is currently affixed to the specimen. Possibly referring to the two Coloradensis that Mead sent in 1872, entry no. 4277 lists an “Anthocaris” collected by Mead during June in “Rocky Mts. Col.”
According to Edwards (1881), the female paralectotype was received from the physician James S. Bailey (1830–1883) of Albany, New York (Curtis 1884). Not only did Edwards correspond with Bailey, he visited Bailey's home and wrote his obituary (Edwards 1883). One of Bailey's sons, Theodore P. Bailey, also collected Lepidoptera. The elder Bailey provided Colorado Lepidoptera to several other contemporary entomologists, including Augustus R. Grote, who described new taxa from his moth specimens. Bailey did not personally travel to Colorado, but obtained “unprepared” specimens through another entomological correspondent who employed a resident of Colorado to collect butterflies and sugar for moths (30.vii.1878, FMNH). Unfortunately, no letters from Bailey were found among Edwards' correspondence at AMNH.
Type locality. Although the source of the lectotype is uncertain, this taxon has been associated with “Turkey Creek Junction” for nearly fifty years and the species is known to occur there. The type locality is more accurately defined as the vicinity of Bradford Junction (“Turkey Creek Junction,” within the present-day town of Conifer), Jefferson County, Colorado.
Colias hagenii W. H. Edwards, 
(=Colias philodice eriphyle W. H. Edwards; Pieridae)
Described from “various localities, from So. Colorado to Montana and Dacotah (Bismarck).” Without explanation, Ferris (1971) attributed the type locality to “Durango, La Plata Co., Colorado.” Brown (1973) designated one of Mead's specimens as the lectotype (Figs. 13, 14) and defined the type locality as “northeastern corner of South Park, Park Co., Colorado.” Affixed to the lectotype (CMNH) is a portion of Mead's original field envelope dated “6/16.” It also includes a fragment of Mead's notation identifying the specimen as a male “[Phil]odice” (Fig. 15), which is consistent with Edwards' (1887b) comment that Mead originally considered these specimens to represent “a variety of Philodice.”
On 16 June 1871, Mead was lodging at Kenosha House, where he “found many Colias” during a walk of 4.8 km (3 mi) along the road “to the South Park” (Table 1). He apparently walked along the Denver & South Park Road (today's I-285 corridor) from Kenosha House over the summit of Kenosha Pass and down toward the South Park basin. Although Brown (1973) stated that the lectotype was from Edwards' collection, there is no such collection label on the specimen (Fig. 15). The presence of the envelope clipping infers that Holland acquired it from Mead and it was still papered. Mead mailed this specimen to Edwards from Bradford Junction 20 June 1871.
Type locality. As defined by the lectotype, the type locality is restricted to within 1.6 km (1 mi) south of the summit of Kenosha Pass, Park County, Colorado.
Discussion. Although most authors since Smith (1891) consigned hagenii to synonymy, Garth and Tilden (1963) and Pohl et al. (2010) resurrected this name to recognize subspecies of Colias philodice Godart. The original description of hagenii characterized a summer phenotype with broad black wing margins (Edwards [1884a]), though Edwards (1887a) eventually considered hagenii to be “seasonally dimorphic.” Referring to specimens that would later be described as hagenii, Mead reportedly observed, “if there could be such a thing as a yellow Eurytheme, this was it” (Edwards [1884a], [1884b], 1887a). Mead's concept of Colias eurytheme Boisduval was of the summer phenotype, as the spring form of this butterfly was then thought to represent a different species, Colias ariadne W. H. Edwards. In his original description, Edwards () stated that hagenii was “between Philodice and Eurytheme … add orange and Hagenii would be indistinguishable from Eurytheme.”
There is still a great deal of disagreement regarding the status of eurytheme, eriphyle, and philodice. Although most lepidopterists now treat eriphyle as a subspecies of philodice, some consider it worthy of specific recognition. Likewise, the true nature of hagenii remains somewhat obscure and even Edwards' concept of this nominal taxon evolved over time. In his original description, Edwards (1884a) noted that some adults of hagenii have an “ochraceous tint.” He also mentioned that the coloration is sometimes “not yellow, but of a peculiar shade, a sort of buff-yellow (better chrome-yellow), a shade not seen in Philodice” (Edwards 1884b). He observed that males “frequently show more or less of [this coloration], and occasionally have a flush of orange.” During the 1880s, Edwards reared adults of hagenii from eggs that were received from Herman W. Nash of Pueblo, Colorado. Edwards (1887a) identified some of these reared adults as eurytheme, prompting him to consider hagenii as a yellow form of eurytheme (Edwards 1887b). He ultimately concluded that hagenii was synonymous with eriphyle, but still regarded the latter as a “tetramorphic form” of eurytheme (Edwards 1887b). The lectotype of hagenii is a slightly darker shade of yellow (Figs. 13, 14), but it is unclear if this coincides with Edwards' reference to an “ochraceous tint” or is simply due to the specimen's age and storage conditions. Many specimens from Holland's collection are discolored due to being stored in his cellar prior to their transfer to the Carnegie Museum in 1896–97 (Brown & Miller 1980).
Although the lectotype of hagenii (Figs. 13, 14) agrees with the original description, Fisher (2012) questioned its date (16 June) because he could not confirm the occurrence of the summer phenotype of C. p. eriphyle during mid-June in the Fairplay area. He argued that the specimen was probably not collected by Mead, but instead was reared from a batch of eggs that Edwards received from H. W. Nash. This is untenable, however, as the writing on the clipping affixed to the lectotype, including the penciled notation “Colo,” is in Mead's hand (Fig. 15). It is also difficult to disregard Mead's accompanying journal entry about finding “many Colias” on its date of capture (16 June), or the fact that he reportedly collected most of his specimens of philodice during June (Mead ). Because Mead mailed specimens from Colorado on a regular basis, it is unlikely that he collected the specimen at a later date and incorrectly labeled its envelope.
Unusual weather conditions during 1871 could explain the presence of a summer phenotype of eriphyle near Kenosha Pass on 16 June. Other captures by Mead near Kenosha House suggest that the 1871 season was earlier than normal (see P. colorado, above, and A. helena, below). It is also possible that the specimen was not of local origin, but rather reached the area from lower elevations where the summer form appears earlier in the season.
Argynnis helena W. H. Edwards, 1871
(=Boloria chariclea helena; Nymphalidae)
Described from specimens “Taken in Colorado by Mr. Mead.” Brown (1934) suggested a type locality of “Mt. Elbert, Mosquito Pass, Mt. Lincoln and Hoosier Pass.” Brown (1965) designated a lectotype and restricted the type locality to “Mosquito Pass, Lake—Park Counties, Colorado.” Although the lectotype is undated, there are several other specimens in CMNH from Mead's collection that possess clippings from his field envelopes dated “7–20.” Brown (1965) suspected that on 20 July 1871 Mead was “en route between Twin Lakes, Lake County, and Fairplay, Park County,” noting that Mead was definitely on Mosquito Pass the following day. On 20 July, Mead explored in the vicinity of Oro City, within California Gulch, Lake County (Table 1). He walked “up the gulch and on the mountains,” but did not reach the higher elevations of Mosquito Pass, located 9.7 km (6 mi) northeast of Oro City. In a letter dated 20 July, which accompanied a box of specimens that had been collected since the last shipment, Mead attributed all the butterflies he captured that day to California Gulch (Brown & Brown 1996).
Two additional dated specimens of helena that Mead collected are deposited at PMNH. A male bears a label written by C. P. Whitney dated “July 3 1871” and a female bears a label written by F. G. Sanborn dated “7 10.” Mead was lodging at Kenosha House on 3 July when he mentioned collecting about 50 specimens of this species “further up and more in the mountains” (probably up Hoosier Creek valley west of the house) (Brown & Brown 1996). This is an early date for this species, suggesting that the 1871 season was advanced (this species does emerge in early July on occasion, as there are two fresh specimens at MGCL from Park County dated 3 July 1990 from an elevation of 3426 m/11,240 ft.). On 10 July, Mead climbed a peak south of Twin Lakes (Table 1) (Fig. 8), where he found “swarms” of this species at the summit (Brown & Brown 1996).
Type locality. Although Mead collected this species at multiple locations, the type locality is restricted to the vicinity of Oro City, California Gulch, Lake County, Colorado. This location is most suitable because of its proximity to the type locality previously proposed by Brown (1965) and its association with several specimens at CMNH where the lectotype is deposited.
Argynnis meadii W. H. Edwards, 1872
(=Speyeria callippe meadii; Nymphalidae)
Described in part from a male specimen taken by Mead at “Turkey Creek Junction [Jefferson Co.], in Colorado,” which Brown (1934) restricted to “Turkey Creek Junction, Colorado … probably the junction of South Turkey Creek with Turkey Creek, a few miles up in the foothills.” Mead's male specimen (CMNH) was designated by Brown (1965) as the lectotype. According to Mead (), it was collected on 6 June 1871. On that date, Mead was at Bradford Junction (“Turkey Creek Junction”) where he “took a lunch along & started down the road,” walking 12.9 km (8 mi), then returned by the same route (Table 1). He stated in a letter (Brown & Brown 1996) that he followed Turkey Creek, apparently walking along the winding Denver & South Park Road (today's South Turkey Creek Road), which paralleled South Turkey Creek northeast of Bradford Junction (Fig. 7). The lectotype specimen was mailed to Edwards from Bradford Junction on 20 June 1871.
Type locality. Restricted to within 9.7 km (6 mi) northeast of Bradford Junction (“Turkey Creek Junction,” within the present-day town of Conifer), along South Turkey Creek, Jefferson County, Colorado.
Grapta hylas W. H. Edwards, 1872
(=Polygonia faunus hylas; Nymphalidae)
Described from “about 20 specimens taken in Colorado, in August 1871, by Mr. Mead.” Brown (1934) defined the type locality as “near Berthoud Pass, with little doubt on the southern slope [Clear Creek Co.].” The lectotype (CMNH), designated by Brown (1967), bears a portion of Mead's original field envelope dated “8/16.” On 16 August 1871 Mead was approaching Berthoud Pass from the north along the Empire & Middle Park Wagon Road (Table 1), when he stopped and collected 15 specimens of a new Grapta. Mead recorded in his journal that these specimens were collected “a little before reaching the pass.” Two paralectotypes, which Mead collected on the same date, are deposited at CMNH (ex. Mead colln.). These specimens were mailed to Edwards from Denver on 22 August 1871.
Type locality. As defined by the lectotype, the type locality is restricted to the northern slope of Berthoud Pass, Grand County, Colorado.
Melitaea eurytion Mead, 
(=Euphydryas anicia eurytion; Nymphalidae)
Described from “Colorado,” which Brown (1934) defined as “Twin Lakes, California Gulch, Fairplay and probably Kenosha. Probably not at Turkey Creek Junction.” Miller and Brown (1981) suggested “South Park, Park Co., Colorado.” Scott et al. (2006) suggested “South Park, probably near Fairplay, Park Co., Colo.” Fisher (2006) proposed “the vicinity of Fairplay or certainly between Fairplay and ‘Kenosha House’ within South Park [Park Co.].” Although Mead () clearly intended to credit the name Melitaea eurytion to W. H. Edwards, modern rules of nomenclature (ICZN 1999, Art. 50.1) dictate that Mead be recognized as the author. For similar reasons, Barnes and Benjamin (1926) also attributed authorship to Mead.
Among the specimens of this taxon at CMNH is a somewhat rubbed male (Figs. 16, 17), which was figured as a “type” by Holland (1931, Pl. 57, fig. 15) and as “typical” by Scott et al. (2006, Pl. 5, top row). It bears a portion of Mead's original field envelope, reading “Melitaea 1” with the date “7/4” written in the same purple ink that Mead used for his journal entry on that date (Fig. 18). On 4 July 1871 Mead was lodging at Kenosha House (Table 1). This specimen was mailed to Edwards from Fairplay on 7 July 1871.
An undated female at CMNH (Figs. 19, 20) was also figured as a “type” by Holland (1931, Pl. 57, fig. 16) and as “typical” by Scott et al. (2006, Pl. 5, second row). This specimen bears a small label in Mead's hand identifying it as “Melitaea Eurytion Edw” (Fig. 21), suggesting it was mounted prior to being purchased by W. J. Holland. It also bears a larger typed label, probably prepared by Holland, which refers to the specimen as a “typr” (sic; type) with the notation “= nubigena fide Strkr,” which likely refers to Strecker (1878), who wrote “Eurytion = Nubigena.” Holland's (1931) figure of this specimen is too reddish, causing it to resemble the lower-elevation subspecies Euphydryas anicia capella (W. Barnes). New images from CMNH (Figs. 19, 20) reveal that it is actually quite dingy, probably due to its early storage conditions. Many specimens from Holland's collection were discolored due to being stored in his cellar prior to their transfer to the Carnegie Museum in 1896–97 (Brown & Miller 1980).
Scott et al. (2006) doubted the occurrence of this butterfly at Kenosha Pass, but adults were photographed there in early July 2008 by Thomas Bentley (Harp 2009, Warren et al. 2012), who indicated (pers. comm.) that they were “quite common.” On 9 July 2014, Andrew D. Warren found this species to be abundant within the meadows along Hoosier Creek directly opposite the former site of Kenosha House, at elevations from 2970 and 3000 m (9744 and 9843 ft.). This species was evidently also common in that area in 1871, as at least twelve other specimens at CMNH were captured by Mead in the vicinity of Kenosha House between 14 June and 2 July. Two additional females, which Mead collected in the same area, are deposited at PMNH (YPM ENT429625 and YPM ENT799979) (Figs. 22, 23, 25, 26). These specimens are from the collection of C. P. Whitney, who received them from Mead in early 1872 (6.iii.1872, RC). Both bear Whitney's labels that attribute them to Mead (Figs. 24, 27). In its original condition, the discolored female at CMNH, identified by Mead as M. eurytion, was phenotypically analogous to the female at PMNH dated 4 July (Figs. 25, 26). Mead's specimens from the vicinity of Kenosha House were likely collected at elevations between 2941 and 3200 m (9650 and 10,500 ft.). Also included among Mead's material at CMNH are three specimens from near Fairplay (Park Co.) and one from Twin Lakes (Lake Co.). At the request of John E. Rawlins, I provided locality information for all of Mead's dated specimens of E. anicia from Colorado at CMNH.
Discussion. Reflecting his frustration with identifying specimens, Mead declared, “Melitaea is I think about the most difficult genus we have …” (15.xii.1872, RC). His idea of M. eurytion applied to a particular color variation of the butterfly currently represented by this name and likely involved a second taxon. A review of the history of Melitaea eurytion is helpful in understanding Mead's concept of this butterfly.
On 9 July 1871, William H. Edwards (WHE) sent a batch of Mead's Colorado specimens to Henry Edwards (HE) for his opinion about their identities. In the associated letter, WHE asked if the species identified as “Melitaea 1” was “anicia or what,” to which HE scrawled “not anicia” (AMNH). WHE informed Mead on 25 August 1871 (MGCL) that HE had determined the species as Melitaea nubigena Behr (=Euphydryas editha nubigena; a California taxon). In the collection catalog of HE (AMNH) is an entry for “Melitaea nubigena” from “Rocky Mts, Colorado” from “T. Mead,” but these may refer to four other specimens, identified as “nubigena”, which Mead sent during 1872 (18.i.1872, RC; 19.vi.1872, AMNH). The two female specimens at PMNH from the Whitney collection are labeled as Melitaea nubigena, as originally identified by Mead on the advice of HE (Figs. 24, 27). Still confused, WHE wrote on 21 March 1873, “There are some remarkable forms among Mead's Colo [Colorado] Melitaeas that I had labelled as Nubigena & I think there are 2 species mix'd“ (AMNH). WHE was clearly flummoxed by the array of melitaeine phenotypes found by Mead and even proposed to call several specimens Melitaea idas, but he never published this name.
In late 1873, WHE proposed the name Melitaea eurytion (14.xii.1873, AMNH), but it was Mead () who actually published it. Crediting the name to WHE, Mead wrote only two sentences about eurytion: “This species is found associated with Nubigena in Colorado, but is much rarer, and does not seem to range to quite so great elevations. The most obvious point of distinction from Nubigena is that the yellow spots of the latter are largely obscured in Eurytion by fulvous.” Despite Mead's publication of the name, Melitaea eurytion was subsequently abandoned. Strecker (1878) was the first to argue that Mead's () comments were not sufficient to serve as an original description. Even WHE disregarded eurytion and instead maintained that all these butterflies from Colorado were either Melitaea anicia E. Doubleday or M. nubigena, and he did not list eurytion as a synonym of either (Edwards , 1877, ). Melitaea eurytion was not included in any checklists until Barnes and Benjamin (1926) considered it to be a synonym of anicia. Holland (1931) resurrected the name to define specimens “from comparatively low altitudes” in Colorado. He figured two of Mead's specimens as “types,” thus establishing the popular concept of this taxon. Brown et al. (1955a) also regarded eurytion as a full species. Miller and Brown (1981) considered eurytion to be a subspecies of anicia, yet acknowledged the name's tenuous nomenclatural availability. This treatment is generally accepted today.
Mead () considered the butterfly that he called Melitaea nubigena to be “quite common throughout the mountain district of Colorado.” He described several larvae that he believed to be this species, one of which suspended itself on 19 June and pupated the following day. He mentioned these same larvae in a letter to W. H. Edwards on 22 June, stating that he had found them in South Park (Brown & Brown 1996). From 8 to 20 June Mead explored around Fairplay and Kenosha House, where he collected many adults of this butterfly. His concept of nubigena was clearly synonymous with the taxon now known as E. a. eurytion and probably included populations from above timberline, which are currently recognized as the subspecies E. a brucei (W. H. Edwards). Mead () also listed Melitaea anicia from Colorado, but stated, “I did not meet with it in the Territory.” Like WHE, Mead apparently associated the darkest high-elevation phenotypes (i.e. extreme brucei) with nominotypical anicia. Lastly, Mead () tentatively identified a single specimen from Bradford Junction as “Melitaea chalcedon” (=Euphydryas chalcedona E. Doubleday), but this was presumably based on an unusually dusky or aberrant individual.
In late 1873, WHE informed Mead of his concept of eurytion. Mead responded, “I found I could separate what I take to be Eurytion quite readily from the Nubigenas by the general appearance but in the details of markings I was unable to satisfactorily find the precise manner of difference except indeed in the greater proportion of yellow in Nubigena.” By introducing the name Melitaea eurytion, Mead () clearly attempted to segregate the more fulvous butterflies within montane populations of “nubigena” (=anicia) in Colorado. The undated female specimen in CMNH, which Mead personally identified as M. eurytion (Figs. 19–21), is consistent with this concept in having its pale spots “largely obscured with fulvous.” Likewise, the pale spots of the analogous female from PMNH, collected on 4 July (Figs. 26, 27), are broadly infused with fulvous scales. Brown (1966b) supposed that Mead used the name eurytion to denote the “red form” of E. a. brucei, but Mead's named female does not agree with brucei, whose alpine populations contradict his account that eurytion “does not seem to range to quite so great elevations.” Adults currently found near the former site of Kenosha House are extremely variable, with some showing characteristics of E. a. capella and a few resemble E. a. brucei. Populations in the vicinity of Kenosha Pass embody the popular notion that eurytion is a highly variable taxon (see Ferris 1981).
Mead's concept of eurytion probably extended to lower-elevation populations of E. a. capella. Although capella is variable, its yellow (or yellowish-white) pattern elements are typically fulvous, corresponding with Mead's original description of eurytion. On 21 June, while exploring along the road just south of Bradford Junction (“Turkey Creek Junction”), Mead mentioned finding “another species of Melitaea of the same size as No. 1, but without any yellow spots” (Brown & Brown 1996). At CMNH is a dated male capella which Mead collected along the same road two days later. In addition, a male capella was figured by Wright (1905, Pl. 19, fig. 165) with the statement “no data, from Colorado; T. L. Mead.” This specimen (Figs. 28, 29) is deposited among the Wright “plesiotypes” at CAS and was listed by Tilden (1975). Unfortunately, it bears only one small label reading “165,” which corresponds to the species/figure number used by Wright (1905) (Fig. 30). This specimen was most likely also captured in the vicinity of Bradford Junction. Because Mead () did not separately mention fulvous lower-elevation populations (i.e. capella), it is reasonable to assume that he considered all ruddy phenotypes to represent eurytion.
Lectotype designation. To preserve nomenclatural stability and fix the type locality, I hereby designate the male syntype at CMNH (Figs. 16, 17) as the lectotype of Melitaea eurytion, Mead,  in accordance with ICZN (1999, Art. 74.7). This specimen was selected because 1) it is from Mead's personal collection, 2) it is dated, 3) it is from an extant population where Mead collected many of these butterflies, 4) its illustration as a “type” by Holland (1931) helped forge the modern concept of eurytion, and 5) Scott et al. (2006) referred to this specimen as “typical” and nearly designated it as the lectotype. The specimen bears 1) a white handwritten rectangular label [Eurytion /♂] in the hand of W. J. Holland, 2) a small rectangular printed and handwritten label [Butterfly Book / Pl. 57 Fig. 15], 3) a small rectangular printed label [Collection / T. L. Mead], and 4) a portion of T. L. Mead's original field envelope, reading “Melitaea 1 / 7/4” in faded purple ink (Fig. 18). A red printed label [LECTOTYPE / Melitaea eurytion / Mead  / Designated by / John V. Calhoun, 2014] (Fig. 18) has been affixed to the specimen. All other specimens of eurytion collected by Mead at CMNH, as well as those at PMNH, are paralectotypes. The specimens sent to Henry Edwards were not located by staff at AMNH. Additional paralectotypes of eurytion likely exist, as Mead's letter copybooks (RC) indicate that he also sent specimens of “nubigena” to J. Behrens, H. K. Morrison and others. Mead collected 297 specimens of “Melitaea and Phyciodes” in Colorado (Mead 1877).
Euphydryas anicia carolae T. Emmel & L. N. Harris is treated by most authors (e.g. Fisher 2006, Scott et al. 2006, Pelham 2008, 2014) as synonymous with M. eurytion. The type locality of carolae is 2.1 km (1.3 mi) south of Fairplay at an elevation of 3005 m (9860 ft.) (Emmel & Harris 1998). This location is about 32 km (20 mi) southwest of Kenosha House, where Mead collected the lectotype of eurytion. Fisher (2006) suggested that Emmel and Harris (1998) disregarded eurytion in the tradition of prior authors, but Thomas C. Emmel (pers. comm.) confirmed that he recognizes eurytion as a valid subspecies and carolae was described solely to differentiate populations of brightly colored anicia that occur in dry-meadows within open valley floors of northern South Park. However, Scott et al. (2006) observed that most adults from the carolae type locality “look like the eurytion types (some quite like the types).” Moreover, some of the adults found around Kenosha Pass are very similar to carolae (Andrew D. Warren, pers. comm.). The various phenotypes of anicia in Colorado need further study to better understand the extreme variability between, and within, populations.
Type locality. As defined by the lectotype, the type locality is restricted to the vicinity of Kenosha House, 2.16 km (1.34 mi) north of the summit of Kenosha Pass, Park County, Colorado.
Phyciodes nycteis var. drusius W. H. Edwards, 1884
(=Chlosyne nycteis drusius; Nymphalidae)
Described from “Colorado and Arizona,” Brown (1966) designated a lectotype from one of Mead's specimens and defined the type locality as “Turkey Creek Junction, Jefferson County, Colorado.” The lectotype (CMNH) bears Mead's small identification label dated “June 26,” indicating that it was probably mounted prior to being purchased by Holland. On 26 June 1871 Mead collected butterflies along the Mt. Vernon Wagon Road and “up a small brook” north of Bradford Junction (Table 1). Specimens collected that day were mailed to Edwards from Fairplay on 7 July 1871.
Type locality. As defined by the lectotype, the type locality is restricted to within several kilometers north of Bradford Junction (“Turkey Creek Junction,” within the present-day town of Conifer), Jefferson County, Colorado.
Melitaea calydon W. Holland, 1931
(=Chlosyne palla calydon; Nymphalidae)
Edwards first proposed the name calydon in 1873 (14.xii.1873, AMNH), but never published it. Citing Edwards, Mead () listed M. calydon without description in association with specimens that he collected at “Turkey Creek Junction.” Holland (1931) resurrected the name calydon to define Colorado specimens that resembled Melitaea (=Chlosyne) palla Boisduval. He figured a pair of “types” and, following Mead (), suggested a type locality of “Turkey Creek Junction, Colorado.” Mead () reported that he collected specimens at that locality during 20–30 June 1871. Although he arrived at that locality on 20 June, he actually departed on 28 June (Table 1). Mead mailed the lectotype specimen to Edwards from Fairplay on 7 July 1871.
Type locality. The type locality is more accurately defined as the vicinity of Bradford Junction (“Turkey Creek Junction,” within the present-day town of Conifer), Jefferson County, Colorado.
Phyciodes camillus W. H. Edwards, 1871
(=Phyciodes pulchella camillus; Nymphalidae)
Described from specimens “Taken in Colorado by Mr. Mead.” Brown (1966) designated a lectotype and defined the type locality as “Fairplay, Park County, Colorado.” Affixed to the lectotype (CMNH) is a portion of Mead's original field envelope, dated “7–4,” written in the same purple ink as Mead's journal entry for that date. On 4 July 1871 Mead was lodging at Kenosha House (Table 1). He mailed the lectotype specimen to Edwards from Fairplay on 7 July 1871.
Type locality. As defined by the lectotype, the type locality is restricted to the vicinity of Kenosha House, about 2.16 km (1.34 mi) north of the summit of Kenosha Pass, Park County, Colorado.
Phyciodes emissa W. H. Edwards, 1871
(=Phyciodes pulchella camillus; Nymphalidae)
Described from “several specimens taken in Colorado by Mr. Mead.” Brown (1966) designated a lectotype and defined the type locality as “Denver, Denver County, Colorado.” The lectotype (CMNH) bears a portion of Mead's original field envelope with the date “6/3.” On 3 June 1871 Mead was lodging in Denver, where he “walked out on the prairie and found plenty butterflies” (Table 1). He followed Cherry Creek, a tributary of the South Platte River that runs southeast from the center of the city. The lectotype specimen was mailed to Edwards from Denver the same day on which it was collected.
Type locality. Restricted to Denver, along Cherry Creek, Denver County, Colorado.
Erebia rhodia W. H. Edwards, 1871
(=Erebia epipsodea brucei Elwes; Nymphalidae)
Described from “Colorado; from several specimens taken by Mr. Mead.” Brown (1934) suggested the type locality of “Fairplay [Park Co.].” Brown (1964) designated a lectotype (CMNH), which bears a portion of Mead's original field envelope dated “7–6.” On 6 July 1871 Mead collected “up” Beaver Creek near Fairplay (Table 1). This specimen was mailed to Edwards from Fairplay the following day.
Type locality. As defined by the lectotype, the type locality is restricted to along Beaver Creek, east/northeast of Fairplay, Park County, Colorado.
Chrysophanus sirius W. H. Edwards, 1871
(=Lycaena rubidus sirius; Lycaenidae)
Described from specimens “taken in Colorado by Mr. Mead.” Brown (1934) suggested a type locality of “Twin Lakes,” as well as “Mt. Lincoln, South and Middle Park.” Brown (1969) designated a lectotype and defined the type locality as “vicinity of Twin Lakes, Lake Co., Colorado.” The lectotype (CMNH) bears a portion of Mead's original field envelope, which is dated “7–13.” On 13 July 1871 Mead was indeed at Twin Lakes (Table 1). He walked “up to the head of the upper lake,” where he found “lots of Chrysophanus” He collected at least 40 males and two females in a “grassy spot” (Mead , Brown & Brown 1996). Additional specimens of sirius which Mead collected during his stay at Twin Lakes are deposited at CMNH (ex. Mead colln.), MCZ (ex. Scudder colln. via Morrison), and PMNH (ex. Whitney colln.). These specimens were mailed to Edwards from Oro City on 20 July 1871.
Type locality. As defined by the lectotype, the type locality is restricted to Twin Lakes, along the northern shore of Upper Twin Lake (the western lake), Lake County, Colorado.
Thecla ninus W. H. Edwards, 1871
(=Callophrys spinetorum spinetorum (Hewitson); Lycaenidae)
Described from three specimens “Taken in Colorado by Mr. Mead.” Brown (1934) cited Mead (), who stated that specimens were taken “on the South Park road four miles from the park.” Shields (1966) defined this as “ca. 1 mi. E Kenosha Pass summit, U.S. Highway 285,” which Brown (1970a) cited as “about 1 mile east of Kenosha Pass, Park County, Colorado.” Although the lectotype is not dated, Mead () reported that he caught all of his specimens at the same locality on 17 June 1871. On this date, he was collecting along the Denver & South Park Road near Kenosha House (Table 1). His comment about taking the specimens “four miles from the park” denoted road miles from the South Park basin. Although the suggested type locality of Shields (1966) is essentially correct, the old wagon road (and existing U.S. Hwy 285) actually crossed the pass in a more north/south direction, not east/west. One mile east of the pass would place the collection site far off the road among the nearby peaks. Mead's specimens of ninus were mailed to Edwards from Bradford Junction on 20 June 1871.
Type locality. Restricted to the vicinity of Kenosha House, about 2.16 km (1.34 mi) north of the summit of Kenosha Pass, Park County, Colorado.
Lycaena daunia W. H. Edwards, 1871
(Glaucopsyche piasus daunia; Lycaenidae)
Described from specimens “Taken in Colorado by Mr. Mead.” Based on Mead (), Brown (1934, 1970b) defined the type locality as “Turkey Creek, Jefferson Co., Colorado.” Mead collected only three specimens during the last week in June, two of which are deposited at CMNH (ex. Edwards colln.). Neither is dated, but Mead spent 20–28 June at “Turkey Creek Junction” (Table 1), where on several occasions he walked along South Turkey Creek (Turkey Creek). His specimens from this period were mailed to Edwards from Fairplay on 7 July 1871.
Type locality. Based on the location of the settlement and Mead's documented activities, the type locality is defined as along South Turkey Creek, within 9.7 km (6 mi) northeast of Bradford Junction (“Turkey Creek Junction,” within the present-day town of Conifer), Jefferson County, Colorado.
Lycaena melissa W. H. Edwards, 1873
(=Plebejus melissa; Lycaenidae)
Described from “many specimens taken in Colorado by Mr. Mead, in the summer of 1871.” Although Edwards (1873) mentioned that he had also received this species from Nevada and Arizona, it was implied that the description was based solely upon Mead's specimens from Colorado. Brown (1934) suggested a type locality of “Fairplay [Park Co.],” which Nabokov (1949) expanded to “Park Co., between Fairplay and Californian Gulch, on the way to Mosquito Pass, 9,500–13,188 ft. alt.” Brown (1970b) designated a lectotype (Fig. 31) and defined the type locality as “vicinity of Twin Lakes, Lake Co., Colorado … probably taken on La Plata Peak,” which Miller and Brown (1981) reinterpreted as “La Plata Peak, vic. Twin Lakes, Lake Co., Colorado.” Scott (2006a) suggested “Tinytown [Jefferson Co.].” Andrew D. Warren agreed with Brown (1934) that Fairplay is a more likely type locality (Pelham 2008).
The lectotype (CMNH) designated by Brown (1970b) bears a portion of Mead's original field envelope dated “7–11.” Mead was at Twin Lakes on 11 July 1871, but he did not climb any nearby peaks (Table 1). He was stiff after ascending a peak the previous day and instead collected in the vicinity of the lodging house (east of the lakes) and walked 4 km (2.5 mi) “above” the house, presumably northwestward along the road (now State Hwy 82). Mead remarked that this species was “very plentiful” around Twin Lakes, where it was “abundant by the first week in July” (Mead , Brown & Brown 1996). In addition to the lectotype, at least two males and five females of melissa which Mead collected around Twin Lakes are deposited at CMNH (ex. Mead colln.). These specimens, including the lectotype, were mailed to Edwards from Oro City on 20 July 1871.
Type locality. As defined by the lectotype, the type locality is restricted to Twin Lakes, along the eastern side of Lower Twin Lake (the eastern lake), Lake County, Colorado.
Discussion. The type locality of Lycaena melissa (Fig. 31) is located about 16 km (10 mi) east of the type locality of Lycaeides melissa pseudosamuelis Nabokov (Fig. 32). The proximity of the type localities and similarity of the primary types prompted Scott (2006a) to speculate on the reasons why a drab high-elevation phenotype was chosen as the lectotype of melissa, even suggesting that F. Martin Brown was “peeved at Nabokov” and deliberately intended to force pseudosamuelis into synonymy. Although Brown (1950a, 1950b) disagreed with Nabokov's statistical methods, I have found no evidence that he had any such deliberate intentions regarding melissa. Appreciating Brown's motivation requires a review of the complex conceptual histories of the nominal taxa Lycaena melissa and Lycaeides melissa pseudosamuelis.
On 4 August 1871, W. H. Edwards informed Mead that three of his Lycaena species appeared to be new, but cautioned that he was not yet certain (RC). In early 1873, Edwards was unsure of the identity of Mead's “Lycaenas,” particularly one “like Scudderii, Edw., that has passed for Anna”(16.i.1873, AMNH). Edwards was referring to Lycaena scudderii (=Plebejus idas scudderii) and Lycaena anna (=Plebejus anna), both of which he had previously described. Seeking another opinion, he sent several specimens to Henry Edwards, referring to them as “all of Mead's collecting in Colorado.” After comparing his type of anna with Mead's specimens, W. H. Edwards wrote, “certainly the two seem distinct and I think the Coloradian is undescribed” (23.i.1873, AMNH). Within two months he had chosen a name: “The Lycaena like Anna I call Melissa” (9.iii.1873, AMNH). One of the male specimens that W. H. Edwards sent to H. Edwards eventually made its way back to Mead. It is deposited at CMNH (Brown 1970b) and was figured by Holland (1898, 1931; Pl. 31, fig. 25). Another specimen, a female, is deposited at AMNH and labeled “4880” in H. Edwards' hand. It corresponds to an entry in his collection catalog (AMNH) for an unidentified species of “Lycaena,” which was collected by Mead during June in “Rocky Mts. Colorado.”
In his original description of Lycaena melissa, Edwards (1873) defined a butterfly with a “complete sub-marginal row of large orange spots” on the ventral hindwings, as well as a row of orange spots on the ventral forewings. He characterized the female as having a “complete orange band” on the dorsal forewings and hindwings. However, Edwards typically chose one specimen of each gender from which to derive his original descriptions (Brown 1965). To define melissa, he presumably selected a pair that significantly differed from his concepts of scudderii and anna (Edwards 1873). Lower-elevation populations of melissa tend to be more brightly colored. Mead's specimens of melissa at CMNH suggest that he collected more from high elevations than from low, supporting the idea that Edwards cherry-picked the specimens he described.
Edwards' correspondence reveals that he intended to illustrate melissa with his description, but did not do so. Instead, he figured dorsal and ventral aspects of the male and female in Mead (, Pl. 36, figs. 5–8) (Fig. 34). Edwards prepared the plates for that publication and undoubtedly selected the figured specimens (Calhoun 2013b). Because Edwards had exchanged much of his share of Mead's 1871 material by that time, the melissa specimens he figured were probably not those characterized in the original description. None of the melissa at CMNH were identified by Edwards as type specimens (Brown 1970b).
Barnes and McDunnough (1916) examined specimens at CMNH, including “presumable types from Colorado” and defined typical melissa as having “heavy red submarginal markings on the underside.” They considered specimens in their possession from Colorado, Utah, Arizona, and New Mexico as “typical.” Holland (1931, Plate 66, fig. 17) figured a brightly colored female from Colorado and identified it as a “type.” Today, nominotypical melissa is generally recognized as a low-elevation taxon with well-developed ventral orange spots that occurs over a large portion of western North America (Downey 1975, Fisher 1981, 2009, Scott 1986, Lane & Weller 1994, Layberry et al. 1998, Gompert et al. 2008) (Fig. 35). Fisher (1981) considered this taxon to be most common in the Rocky Mountains at elevations of 1525–2135 m (5000–7000 ft.). Despite our modern interpretation of nominotypical melissa, this taxon was treated differently during the four decades between the publication of Holland (1931) and Downey (1975).
Around 1932, F. Martin Brown began researching Mead's itinerary in an effort to clarify the type localities of taxa attributed merely to “Colorado” by W. H. Edwards. Regarding melissa, Brown (1934) stated, “Mead gives Fairplay as the probable locality for the 1871 series.” This assertion was derived from Mead's () remark that specimens “were brought in by the expedition of 1871, probably from the vicinity of Fairplay.” Brown, however, mistakenly assumed that Mead was a member of a larger survey party and likely interpreted Mead's comment to mean that another party member had collected the type series, but the locality was unknown. Rather, Mead's uncertainty about the locality implies that he was referring to a different expedition than his own. In several instances Mead () cited the “expedition of 1871” in reference to other exploring parties. Mead was probably referring to the Allen Expedition, who was in the vicinity of Fairplay in late July 1871 (Brown et al. 1956). It is also possible that Mead intended to cite the Wheeler Expedition of 1873, whose party collected in South Park, including Fairplay, during July and August (Yarrow [1876b]). Mead also listed melissa from Denver, and “In the South Park and about Twin Lakes.” Based on what we now know about his itinerary, he was referring to his own specimens from those areas. Nonetheless, Brown's concept of nominotypical melissa was forever biased by Mead's reference to Fairplay, which is roughly 3048 m (10,000 ft.) in elevation.
While Brown (1934) began to shift the concept of nominotypical melissa, it was Nabokov (1949) who formally advocated the change. Among the melissa specimens that Nabokov examined at MCZ was a female collected by Mead on 8 July 1871. Nabokov stated that “Mead's label” read “Divide between the Arkansas and S. Platte Valleys.” This specimen supported Brown's (1934) assertion that the type material was from the vicinity of Fairplay. In addition, Brown (1934) placed Mead on 8 July 1871 “on the divide between Fairplay and California Gulch” (i.e. Mosquito Pass), which corresponds with the data on the specimen. Nabokov therefore considered the type locality of melissa to be “Park Co., between Fairplay and Californian Gulch, on the way to Mosquito Pass, 9,500–13,188 ft. alt.” Accordingly, Nabokov (1949) regarded Mead's female specimen as a topotype and figured it on his Plates 6 and 9. I examined images of this specimen and found that its label was not written by Mead, but by Herbert K. Morrison, who received the specimen from Mead on 20 February 1873 (Mead referred to it in his letter to Morrison as “L. anna?”) (RC). Morrison probably sent this specimen to S. H. Scudder, whose collection is deposited at MCZ. Morrison's label, based on data provided by Mead, reads, “Divide between the / Arkansas & Platte / valleys July. 8.1871.” In an effort to reconcile the locality label with Brown's (1934) allusion to Fairplay, Nabokov affixed a label to this specimen which reads, “vic. Fairplay, / Mosquito Pass / (see Brown 1934 J. N.Y. / Ent. Soc.) leg T. L. Mead.” It is no secret that Nabokov greatly admired Brown's studies of Mead (Boyd & Pyle 2000).
Influenced by Brown's assumptions and encouraged by Mead's specimen, Nabokov (1949) concluded, “typical melissa is not the low level (sage belt, oak brush, alfalfa, prairie, etc.) form or forms, with richly ornamented underside and female upperside, but an altitudinal, comparatively drab race, little known to collectors.” To conserve the “general concept of melissa,” he considered the “richly ornamented” low-elevation populations as an extreme form. A male melissa, collected by Mead on 22 July 1871 between Fairplay and Montgomery City, Park Co., Colorado (PMNH, ex. C. P. Whitney colln.) (Fig. 33), is consistent with nominotypical melissa as interpreted by Brown (1934) and Nabokov (1949).
Nabokov (1944) was the first to mention “a curious Colorado form” of melissa with a reduced ventral orange band. He later described this form as the subspecies L. m. pseudosamuelis to “delimit the geographically adjacent type form of melissa melissa on the negative side of its pattern” (Nabokov 1949). He admitted that pseudosamuelis (Fig. 32) was the “weakest of melissa races in Colorado,” with the “Fairplay typical form coming next, and the S. E. Colo. form bringing ssp. melissa to its maximum expansion in Colorado” (Figs. 33, 35). In essence, Nabokov described pseudosamuelis to narrow his definition of the nominotypical subspecies. Nabokov (1955) later characterized a population of melissa from above timberline in the Sierra Madre Range of Wyoming as a “colony of typical (alpine) L. melissa melissa as described by Edwards,” thus confirming his belief that typical melissa is a high-elevation taxon separate from pseudosamuelis, which he limited to Pitkin and Lake Counties of Colorado.
Ten of the 17 type specimens of pseudosamuelis, including the holotype (Fig. 32), were collected by the Kansas lepidopterist John R. (J. E. Rice ) Turner (1910–2000) along US Hwy 82 in the vicinity of the Red Mountain Inn, Lake Co., Colorado. The former Red Mountain Inn was a late 19th century stage stop along the Twin Lakes toll road, 9.7 km (6 mi) west of the town of Twin Lakes, at an elevation of about 3048 m (10,000 ft.) (Scott 2004) (this is about where the red arrow points to Lake Creek on Fig. 8). Turner's specimens are undated, but they were most likely collected during the 1940s (that section of the old toll road was designated as part of US Hwy 82 in 1927; it was paved in 1967 (CDOT 2012)). Possibly in an attempt to further segregate pseudosamuelis, Brown et al. (1955b) limited this taxon to populations “generally above 11,500 feet,” ignoring the fact that the elevation of its type locality is 10,000 ft. Echoing Nabokov's analysis, Brown observed that specimens of melissa from the plains seem to “differ more from typical material caught at Fairplay than the Fairplay specimens differ from pseudosamuelis.”
Brown's notion of typical melissa was reinforced during the 1960s when he examined Mead's specimens at CMNH and discovered that they incorporated both high-elevation and low-elevation phenotypes. Edwards received and examined all of Mead's material from Colorado as it was collected, thus Brown considered Mead's entire catch to represent the type series (Brown 1970b). Agreeing with Nabokov (1949), Brown defined high-elevation populations as “true melissa” and low-elevation populations as the “better known atypical form.” That Edwards himself was aware of the variation in melissa is suggested by the figures in Mead (), which Edwards prepared under his personal direction. The ventral male is especially evocative of a high-elevation phenotype (Figs. 33, 34). In fact, Nabokov (1949) considered all the specimens figured by Mead () to represent “typical” (i.e. high-elevation) melissa.
In selecting a lectotype of melissa, Brown worked with CMNH curator Harry K. Clench, a specialist of Lycaenidae. Brown (1970b) recalled that when reviewing lycaeinid material at CMNH, he and Clench “spent days upon days at the Carnegie Museum studying, discussing and arguing about this and that related to the selection of the Edwardsian specimens to be designated the type.” In November 1968 they selected one of three dated male specimens of melissa at CMNH which could be attributed to a high-elevation locality. Presumably because none of the available specimens at CMNH were from Park County, they chose one from another locality mentioned by Mead (): Twin Lakes (Lake County), which is the only high-elevation locality definitely represented among Mead's specimens in that collection (see Brown 1970b). Brown affixed a label to the specimen indicating that it was chosen as the lectotype by “F. M. Brown & H. K. Clench/XI 1968.”
The lectotype designation for melissa was based on a high-elevation concept as defined by Nabokov and supported by Clench, both of whom Brown recognized as authorities on this group of butterflies. This action, however, created a potential conflict with pseudosamuelis, which Brown (1970b) did not address. This is all the more surprising given that Brown believed that the lectotype was collected on La Plata Peak, a mountain over 4267 m (14,000 ft.) in elevation —well within the domain of pseudosamuelis as previously defined by Brown himself—and located only 4.8 km (3 mi) from the type locality of pseudosamuelis. Moreover, Nabokov (1949) had previously questioned if populations from Twin Lakes represented nominotypical melissa or another subspecies (i.e. pseudosamuelis). Undoubtedly because of the proximity of the type localities, Brown subsequently treated pseudosamuelis as a junior subjective synonym of melissa in Miller and Brown (1981). Perhaps bowing to criticism from those who preferred conventional usage, pseudosamuelis was again listed as a subspecies of melissa by Miller and Brown (1983).
Despite the interpretation of nominotypical melissa by Brown and Nabokov, Downey (1975) reintroduced the concept of this taxon as a “lowland, brightly colored” sagebrush-prairie inhabitant. He recognized the subspecies pseudosamuelis as “the extreme of an altitudinal cline.” Perhaps in protest, Downey made no mention of Brown's lectotype of melissa and instead reiterated the type locality of Nabokov (1949): “Park county, Colorado, between Fairplay and California Gulch.” Ironically, Downey's acceptance of a high-elevation type locality contradicted his premise and contributed to the confusion.
Lane and Weller (1994) mapped the distribution of pseudosamuelis to include a very small area surrounding its type locality in Lake County. Scott (2008a) collected additional examples of pseudosamuelis at its type locality and defined its range as “several of the cold valleys of the northern Sawatch Range.” Fisher (2009) observed that phenotypes resembling pseudosamuelis also occur at higher elevations in the southwestern mountains of Colorado. Scott (2006a) mentioned similar butterflies in the Bighorn Mountains of Wyoming, which he satirically referred to as “bighornimuelis.” Scott (2008a) later associated those populations with the taxon now recognized as Plebejus idas longinus (Nabokov). Although pseudosamuelis was reported from New Mexico by Ferris (1976) and Holland (1984), these records were not listed by Toliver et al. (2001). Further complicating matters is the extreme similarity between pseudosamuelis and the taxon currently recognized as Plebejus idas sublivens (Nabokov). Nabokov (1949, 1950) mentioned their resemblance and subsequent authors (e.g. Brown et al. 1955b, Scott 2006a, 2008a, Fisher 2009) postulated on their potential relationship. Gompert et al. (2010) found that idas and melissa have hybridized extensively in portions of the central Rocky Mountains, resulting in substantial admixture and nuclear introgression. Although authors continue to associate pseudosamuelis with melissa (Pelham 2008, 2014, Fisher 2009, Scott 2008a, 2014), it may be more closely allied with idas (A. D. Warren, pers. comm.). Scott (2008c) included all melissa subspecies within his own concept of an idas superspecies, which he sardonically termed a “stenchospecies.”
In an effort to fix the name melissa to a brightly colored lowland taxon, Scott (2006a, 2008c) asserted that Holland (1931) had designated a lectotype using a specimen from a low-elevation population. Holland referred to three figured specimens as “types:” a pair on Plate 31 and a female on Plate 66. Only the female on Plate 66 was identified as a type on the accompanying plate legend, which, Scott argued, demonstrates Holland's intent to select that specimen as the sole name-bearing type (i.e. lectotype) per ICZN (1999, Art. 74.5), thereby invalidating the lectotype designation by Brown (1970b). Scott (2006a) suggested a type locality of “Tinytown” (Jefferson County), which is about 2073 m (6800 ft.) in elevation. I disagree with this reasoning, as Holland did not unambiguously select the female to serve as the unique type as required by the Code (ICZN 1999, Art. 74.5), nor was there any visible intent to do so. What Scott interpreted as a conscious act by Holland was likely nothing more than an editorial artifact.
The revised edition of The Butterfly Book by Holland (1931) was based upon a first edition of 1898, which included 48 plates. Only two specimens figured in Holland (1898) were identified as types on the accompanying plate legends (Pl. 30, figs. 9, 33). For the revised edition of 1931, Holland added 29 additional plates, many of which included specimens that were identified as types on the plate legends. In only a few instances did he go back and add “type” captions to the original 48 plates. Simply stated, Holland (1931) did not identify the two specimens as types on the legend for Plate 31 (figs. 25, 26) because he had not done so in Holland (1898). Finally, there is no explicit statement on the legend for Plate 66 of Holland (1931) to suggest any intent to select that female as the sole name-bearing type. In the text of this book, Holland referred to three figured specimens as “types” (plural) without further restriction. The plate legend does not supersede the textual reference and instead merely denotes one of the three “types” mentioned in the text. I therefore agree with Pelham (2014) that the lectotype designation of Brown (1970b) is valid.
James W. Tillery found Mead's 1871 journal and recognized its importance. Wenxian Zhang and Darla Moore (RC) granted access to the manuscripts of T. L. Mead. Jacqueline Y. Miller and Andrew D. Warren (MGCL) hosted my visits to examine specimens and manuscripts under their care. For patiently responding to my many inquiries and providing useful images, I thank Rodney Eastwood and Rachel Hawkins (MCZ), Lawrence F. Gall (PMNH), Suzanne Rab Green (AMNH), Vincent F. Lee and Mike Narahara (CAS), John E. Rawlins and Jane Hyland (CMNH), Andrew D. Warren (MGCL), and Weiping Xei (LACM). Sarah Gilmor (History Colorado, Denver), Susan Morris (Conifer Historical Society, Conifer, Colorado), Hanah Q. Parris (Denver Public Library, Denver, Colorado), Gregory A. Rami and Mia Qaraman Reitmeyer (Archives of the Library, AMNH), and Christie Wright (Park County Local History Archives, Fairplay, Colorado) supplied information, as well as scans of articles, manuscripts, and historical photos. For bestowing additional helpful information and support, I thank Thomas Bentley, Michael M. Collins, Thomas C. Emmel, Nick V. Grishin, James L. Monroe, Chris C. Nice, and Jonathan P. Pelham. Paul Butler generously shared some of his research and publications about Mead's life. Michael E. Toliver kindly reviewed the manuscript and offered helpful comments. Finally, I thank Andy Warren, a respected authority on Colorado butterflies, for thoroughly reviewing drafts of the manuscript and providing much valuable input.