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Symposium: Insect Behavioral Ecology—2001: Lloyd 99

ON RESEARCH AND ENTOMOLOGICAL EDUCATION VI:FIREFLY SPECIES AND LISTS, OLD AND NOW

J

AMES

E. L

LOYD

Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611

A

BSTRACT

Lists of insect species are useful for insect biologists and students in several fields, includingtaxonomy, behavioral ecology, conservation, and biological control, and they are useful to theteacher of classical entomology for the insight and drama they can provide to the history andbiography of our science. Such lists can be viewed as cooperative projects that have combinedthe efforts and enthusiasms of naturalist/taxonomists along a time line, and as they evolvethey can become ever better guides to observation and identification, and to new and inter-esting biotaxonomic problems. The list of NA fireflies recorded here gives the number andcontinental location of working species now recognized after long study of this taxon at thebench, in the library, and afield by many naturalists and taxonomists across more than twocenturies, and makes status changes in a few species.

Key Words: Lampyridae, fireflies, fauna, checklist, species problems, teaching

R

ESUMEN

Las listas de especies de insectos son útiles para los biólogos y estudiantes de insectos en va-rios campos, incluyendo la taxonomía, la ecologia de comportamiento, la conservación, y elcontrol biológico, y son útiles para los maestros de entomologia clásica para la comprensióny el drama ellas que pueden proveer a la historia y la biografía de nuestra ciencia. Tales lis-tas pueden ser vistas como proyectos cooperativos que han combinados los esfuerzos y el en-tusiasmo de los naturalistas/taxónomos por toda una linea de tiempo, y mientras que ellossean se desarrollados pueden convertirse en mejores guias para la observación e identifica-ción y para los problemas nuevos e interesantes biotaxonómicos. La lista de luciérnagas deNA registrada aqui dá el número y la ubicación continental de especies ahora reconocidasdespués de un largo estudio de este taxón en el laboratorio, en la biblioteca, y en el campopor muchas naturalistas y taxónomos a travéz de más de dos siglos, y hace cambios en el es-

tatus en algunas especies.

Ambiguous Firefly

A lightningbug flashes o’er wet meadows,short-long,

No taxonomist ‘til Barber ever noted its song;Oh Firefly,

Did you before Barber to a species belong,Or, is it really no matter, forLists can . . . not be wrong?

After forty years of pursuing firefly species viabiotaxonomy using a “semiosystematic” approach(Lloyd 1969, 1990a), I ask whether it has been afool’s mission—“an unexamined life is not worthliving.” I knew at the outset that the chase was tobe neither occupation nor career, but preoccupa-tion and life, common among insect naturalist/taxonomists in a Camelot once upon a time. Dis-cussions with teachers and mentors about blackand sugar maples and the intermediates of thesenamed species that range from wet bottoms to dryuplands in Michigan’s hardwood forests; aboutvirtually identical field crickets with differentnames in Bermuda and Florida; and about period-ical cicadas across eastern U.S. with the same

names but separated in time and space, allseemed to the student to reveal unfathomable tax-onomic and genetic mysteries or nomenclaturalanomalies. These examples with those emergingfrom the chase, juxtaposed and contrasted withthe relative simplicity and undeniable genetic im-perative, the

sine qua non

of Biological Speciesfound in taxonomic textbooks, have encourageduncertainty and repeated reevaluation of almostevery taxonomic conclusion I have been temptedto draw about fireflies (Lloyd 2001).

While chasing in the dark I learned signalsand seasonality, interactions and distributions,and watched then hand-collected (only one at atime) thousands of voucher specimens which Ikilled and saved, each co-referenced with verbaland sometimes electronic records of signaling be-havior. This was the best of lives! And the worst ofit—or was this somehow the best too?—was thatthe more I learned from watching fireflies flash-ing, the less confidence I had in my understand-ing of species, of what should be named and puton lists. My appreciation of

Acer

,

Gryllus

, and

Magicidada

taxonomists grew, and textbook dis-cussions of species and speciation, driven mainly

100

Florida Entomologist

86(2) June 2003

by vertebrate mega-models perhaps, seemedlargely irrelevant for their authors mostlydwelled at grander scale.

For students who pursue the nature and originof species while wading through literal and liter-ature sloughs I suggest a stress-relieving addi-tion to the Species Concept list of Mayden (1997),a perspective that I withheld last time (2001).This can help them find more appreciation for oldand seemingly useless lists of species, and placetheir own studies in a time line of uncertain ter-mination and perhaps infinite length. A

Tran-scendental Species Concept

suits my presentcomprehension (apprehension too) of many fire-flies, and those of the genus

Photuris

in particu-lar. It may comfort and reassure thoughtfulneophyte biotaxonomists as they increasinglycome to grips with: (1) the array of diversity foundamong “conspecific” local populations (or should Isay an unexpected number of very localized “spe-cies” in the field?); (2) the seeming unreasonabil-ity and maybe even theoretical improbability ofany connection and genetic cohesion among “con-specific” demes of many of them; and (3) our inad-equacy to ever gain all of the information,especially the imponderables of genetics, neededto understand them and in particular, their demehistories and origins.

From handy desk and forgotten references Iretrieve fragments of thoughts and phrases aboutthings transcendent and transcendental that en-courage this suggestion: . . . ideas beyond therange of experience . . . elements of experiencebut not from sense-perception . . . extending or ly-ing beyond the limits of ordinary experience . . .beyond comprehension . . . we may discover manyfacts and learn many details but there are somethings we can never truly know . . . beyond hu-man experience but not human knowledge . . . re-lating to experience as determined by the mind’smakeup. . . . I can also suggest this axiom, a mne-monic aphorism paraphrased from the late MayorRichard Daley (Sr.) of Chicago, though the Mayorwas speaking of politics—“All firefly biospeciesare local.”

In seeming contradiction, such apparent meta-physical sophistication does not cool my interestin species lists, and serves here as prelude andpredicate for mentioning some old favorites, andmaking a new one, already looking toward nexttime. Entities that historically have been namedand listed as species can be understood as book-marks that taxonomists have worked up to, havereached in their reading of nature. This is becausetaxon recognition and the characters examinedand ultimately valued up and down the hierarchyfrom Species (or should I start with Subspecies?)to Order have changed, intermingled, and syner-gized over time (Wilson & Doner 1937; Table 1),just as taxonomy’s concepts, preconceptions, andpreconditions for taxa have.

Who cannot appreciate the fact that the spe-cies-level taxonomy of North American fireflieshas evolved? It has tried and today retains andcombines useful elements from external anatomy,genitalic structure, and flash pattern form andvariation, accomplishing this under the guidanceof lampyrid luminaries such as J. L. LeConte(>mid 1800s), F. A. McDermott and H. S. Barber(early and mid 1900s), and J. W. Green (mid1900s). Who cannot predict that it will continue toevolve as (1) field-savvy molecular biologiststranslate and understand the texts, both wordsand syntax, and the enigmatic operations of DNAstrands (K. Stanger-Hall, in prog.; M. Branham, inprog.); and (2) behavioral ecologists scrutinize theinfluence of signal-tracking predators (e.g.,

Pho-turis

females) on the signal-codes cum counter-measures of their firefly prey, and sexualselection’s guidance of mating behavior and repro-

TABLE 1. SYSTEMS OF CLASSIFICATION THROUGH HISTORY,AFTER WILSON AND DONER (1937). THE EARLI-EST MENTIONED DATES AND AUTHORS ARESHOWN. SYSTEMS AND/OR CHARACTERS FROMTHEM HAVE BEEN DISCUSSED, WINNOWED, DE-VELOPED, COMBINED, AND REFINED OVER DE-CADES, AND CENTURIES. SOME IDEAS HAVESEEMINGLY DISAPPEARED AS SUCH (CIRCULARCLASSIFICATION), BUT ONE CONCEPT HAS RE-PLACED ALL OTHERS AS THE CENTRAL THEMEAND SCAFFOLD OF BIOLOGICAL CLASSIFICATION,AND CHARACTERS FROM OTHER SYSTEMS ARE IN-TERPRETED ONLY THROUGH IT—PHYLOGENET-ICS—WHICH APPEARED SOON AFTER DARWIN’S“ORIGIN.” NAMES AND DATES IN THIS TABLE ARENOT NECESSARILY THOSE OF THE MOST IMPOR-TANT OR THOUGHTFUL CONTRIBUTIONS.


ductive morphology (Lloyd 1979ab, 1981, 1984,1990a; Lloyd & Wing 1983; Wing 1985, 1991, et al.1983; Lewis & Monchamp 1994)—and in particu-lar, the variations of these in time and space?

Species on lists are beginnings and steps, butnever, in our time or in any foreseeable future, theends of biotaxonomic discernment and discrimi-nation. Taking the remarkable

Photinus

and

Pyractomena

revisions of Green (1956, 1957) asexamples, each provided an insightful and solidmorphological foundation from which to pursuebiospecies. Many of Green’s carefully consideredomnispective species (Blackwelder 1967; Mayden1997; Lloyd 2001) are already known to be focalpoints of biotaxonomy’s species “complexes”—isn’t “fuzzy clans” more accurate and descriptive?I view species lists past and passed as mile posts,even commemorative cornerstones of taxonomicaccomplishment, and a foundation for biotaxono-mists and allies today as they seek and track life’smore cryptic paths to diversity.

I like old species lists that include forgotten lo-calities and other lore. I especially like old liststhat tell things about comrades-in-pins who madeor collected for them, who often chased insects asa way of life, and the times when they lived. Thelabels on two firefly specimens in the Museum ofComparative Zoology at Harvard say “BelfrageTexas” (Lloyd 1968). I couldn’t find Belfrage onany Texas map or in any Gazetteer (though itshould have been). Then, Prof. Irving Cantrall,himself virtually part of the collection at Ann Ar-bor who worked daily to put the archived orthopsin good order, and a walking repository of such in-formation, told me that Gustaf Belfrage collectedand sent specimens to several museums in Amer-ica and Europe, and “Irv” recommended “Natural-ists On the Frontier” by Geiser (1937). Geiser’sbook told the tale of this Swedish nobleman whospent the last 15 years of his life collecting insectsin Texas, beginning about 1867. Though his spec-imens were in museums from Washington to St.Petersburg, Russia, the inventory of his Texas es-tate was “almost indecent in its revelation ofstark poverty” (Table 2). Nevertheless, on one oc-casion he purchased his first two glowworms(lampyrid beetle larvae) for the exorbitant priceof 5 dollars (cf Table 2), out of delight in them, andlater sold them for much less (Geiser 1937:304).

The hardships and hazards some travelers en-dured to make collections went far beyond whatwe encounter today—except for rare individualssuch as the late Joseph Anderson, a prospectivestudent who died not long ago of malaria whilecollecting fireflies in Africa—though I personallyknow of several scientists on one well-endowedtropical expedition who, with considerable suc-cess, “deliberately” exposed themselves to thismalady by sleeping in native huts to sample fullythe flavour of their exotic excursion. Who cannotbe interested and smile when reading the travels

of the 19th century naturalist who feigned de-mentia for protection from scalping abuse by res-idents, who even helped him to the regionaltrading post!; or feel the anguish when reading ofthe sinking of a sailing ship with the long-nur-tured and carefully protected collections of an en-tomological or botanical adventurer’s travels?; orknow the grief of a father whose son died of yellowfever in the New World, mayhaps after sending

TABLE 2. THE INVENTORY AND ESTATE OF GUSTAV BEL-FRAGE FROM HIS RUSTIC CABIN IN CENTRALTEXAS, CA. 1884. HE WAS A 19TH CENTURY IN-SECT COLLECTOR LIVING THE SIMPLE BUT NOTEASY LIFE. HIS LAST NAME APPEARS ON THE LA-BELS OF MANY ARCHIVED SPECIMENS AND CANEASILY BE MISTAKEN FOR A LOCALITY—AND . . .WITH INFORMATION PROVIDED BY HIS BIOGRA-PHER, INDEED IT ALMOST IS.

102 Florida Entomologist 86(2) June 2003

him now-acclaimed specimens, three of whichthat became the syntypes of Fabricius’ Photurisversicolor (Fig. 1)?

In the spirit of these FES firefly Letters (1998-2001), species lists are a passport to take studentsto classical insect taxonomy with the promise ofpersonal adventure, and they can provide rolemodels of persistence and endurance, and lifelongfulfillment—(see Osborn 1937; Peattie 1936; Gei-ser 1937; Mallis 1971; R. F. Smith et. al. 1973;Kastner 1977; Elman 1982; Porter 1986; So-rensen 1995)—and show that there was life be-fore and it will and must be better after NSF, aswell as encourage a spirituality and pride, and ex-pectation that seems lacking in many academicinstitutions and curricula (Bennett 2001-02).

Letter 41Regional and Other Lists Of Fireflies—

More ThanPassenger Lists For Arks

The hours I spent with thee, dear heart,Are as a string of pearls to me;

I count them over, every one apart,My Rosary.

(Robert Cameron Rogers)

Dear Fireflyers, Listing the names of insectspecies that occur in a region is more than an es-oteric ritual of taxonomic entomologists, and suchlists are more than scorecards for life-list hobby-ists or doomsday records for suspicious environ-mentalists. In insect taxonomy species listing isas fundamental as naming species, and listingcan contribute to understanding the biological,geological, and even the cultural history of a re-gion. Species lists are as marked stepping placesinto a murky bayou, at first tentative and inse-curely grounded, but they provide guidance andfooting for further exploration, and welcome in-formation for naturalist/taxonomists who will fol-

low. As species lists evolve they can becomeannotated catalogues, with references to taxo-nomic histories, and eventually provide detailsfor finding and recognizing each entered entity,with notes on their biology, and more.

History and Overview. Over the past centuriesthere have been several lists with fireflies ofNorth America. In the 1800s fireflies were in-cluded in lists of beetles found during scouting ex-peditions to the west by the U.S. Army, and nearlya century later the “Leng catalogue” listed all ofthe described species of beetles known to occur inNorth America north of Mexico. Leng numberedeach species for easy reference (Table 3), an opti-mistic flourish that had been used before. In 1885Samuel Henshaw had used different numbers inhis list for the same domain—unlike scientificnames, older assigned numbers for each speciesdo not have official and compelling seniority, “pri-ority” as it is called in taxonomy. (Leng remem-bered, and dedicated his list to Henshaw.)

Some lists were exclusively of fireflies and oth-ers included fireflies as but one of many families.There were lists of Lampyridae when the familyincluded leatherwings now classified as net-winged beetles (Lycidae), giant glowworm beetles(Phengodidae), soldier beetles (Cantharidae), andOmethidae (Matheteus, Ginglymocladus) (Le-Conte & Horn 1883)—and some we now don’tknow where to classify (Pterotus, Branham &Wentzel 2001). One “list” was an encyclopedic1386 page treatment of the Coleoptera of Indiana,which the author referred to as a “paper” (Blatch-ley 1910; Fig. 2)! This opus is a model to whichany taxonomist/naturalist worthy of brass-headed, stainless-steel, German-made insect pinsshould aspire. It has within-state distributions,habitat notes, keys, and sketches for identifica-tion. As explanation for his monumental workWillis Blatchley wrote (:3):

“Happiest those days in which I havewandered far and wide through field andwoodland, adding here and there somespecimen before unseen, noting now andagain some life habit, some food plant orplace of retreat, before unobserved. Everand always, however, have I felt the need ofsome one work to which I could refer, somemanual or descriptive list by which I couldlocate the name and place [i.e., relation-ships] of the specimens at hand. Since thebeetles or Coleoptera form one of the mostabundant and attractive groups of insectsand are easily collected and preserved,they would furnish a favorite subject forstudy, especially in high schools, could theyonly be named and placed.”

Lists sometimes had an appendix with thenames and “nicknames” of habitual and favoritecollecting localities in the region (Table 4). These

Fig. 1. The three syntypes of Fabricius’ Lampyris(now Photuris) versicolor, perhaps sent to “Dom. Her-schel” in Europe by his son, thence given to Fabricius(but see Madge 1994).


are useful even today when the labels of old but es-pecially valuable archived specimens bear obscurelocalities. A student might wish to determinewhether a listed rare species yet survives, and ifso, say, whether DNA texts or other chemicals inthe dried, detached legs of museum specimensmatch those in legs found hopping and climbingthere today. There is a dictionary of “AmericanPlace-names” (Stewart 1970), and an entomologi-cally-focused taxonomic list with unknown placesfound on specimen labels (Townes & Linna 1963).

Frederich E. Melsheimer’s 1853 list is one ofthe older for North America; in 1806 his father’shad published a catalogue of Pennsylvania in-sects. Their personal collection is preserved atHarvard and has special value for us in our effortsto make lampyrid nomenclature as error-free aspossible. Here’s why: though Thomas Say’s “type”(name-vouchering) specimen of Pyractomena an-gulata (Say) is lost—some say it was recycled intocarpet beetles—Say corresponded with both F. E.Melsheimer and his father Frederich V., and Saymay have examined and compared his P. angulataspecimen with the Melsheimers’. Thus we have

TABLE 3. EXTRACTS FROM THE FIREFLY LISTING IN THEORIGINAL LENG 1920 CATALOGUE OF NORTHAMERICAN BEETLES, A MAJOR AND EXTREMELYAMBITIOUS COMPILATION IN WHICH EACH SPE-CIES WAS GIVEN ITS OWN NUMBER. FORMAT-TING SIMULATES THAT OF LENG AND HIS(PRESUMPTIVE) ERRORS ARE KEPT. THE INDI-CATED HAND-CORRECTIONS WERE PRESUMABLYPENNED BY THE PREVIOUS OWNER OF MY COPY,FRANK MCDERMOTT. NOTE: SPECIES NUMBERSAT THE LEFT, HYPHENATED REFERENCE NUM-BERS (DATE-PAGE), AND GEOGRAPHIC DISTRIBU-TIONS; UNNUMBERED SPECIES EPITHETS ARE(A) PRESUMPTIVE SYNONYMS OF THE NUM-BERED EPITHET IMMEDIATELY ABOVE, OR (B)NAMES PROPOSED BY ONE AUTHOR BUT DIS-PUTED BY ANOTHER. THE DOUBLE VERTICALLINE SYMBOL UNDER 6988 INDICATES THAT“VITTIGER” WAS SYNONOMIZED BECAUSE THENAME WAS “PREOCCUPIED,” AND SINGLE RIGHT-FACING BRACKETS ([) BEFORE LOCALITIES INDI-CATE THAT THEY BELONG TO THE LINE ABOVE;“L. SUP.” IS THE LAKE SUPERIOR REGION (PAGEVII), PROBABLY FROM A MUCH EARLIER ANDWELL-KNOWN EXPEDITION THERE BY LECONTEAND OTHERS (AGASSIZ 1850).

Fig. 2. A portrait of Willis Blatchley at age 45, in1904. There are several extant photographs of this nat-uralist/taxonomist in action, camping, panning for gold,and picking insects out of a beating umbrella, but theone most often seen, unfortunately, is from much later inhis life and lacks the dynamism and strength of his per-sonality. This photo is from his “Blatchleyana” of 1930.


circumstantial reason to believe that the Melshe-imer specimens compared favorably with Say’s, atleast to Say’s taxonomic eye—and after all, Say

has been referred to as the “Father of AmericanDescriptive Entomology” (Mallis 1971:16).

For history and flavor here are some titles andphrases from title pages of lists that include fire-flies: Explorations and surveys for a railroad routefrom the Mississippi River to the Pacific Ocean—War Department; route near the forty-seventh andforty-ninth parallels, explored by I. I. Stevens,Governor of Washington Territory, in 1853-‘55—Report upon the insects collected on the survey—The coleoptera of Kansas and eastern New Mexico(accepted for publication 1859)—Coleoptera ofFort Whipple, Arizona (1866 [Wyatt Earp wasabout 18 and not yet in Arizona]—List of the Co-leoptera of Vancouver’s Island (1869)—Check listof the coleoptera of America, north of Mexico(1873)—Catalog of the Coleoptera of Mount Wash-ington, New Hampshire (1874)—New species ofColeoptera collected by the expedition for geo-graphical surveys west of the 100th meridian, Lt.Geo. M. Wheeler, Corps of Engineers, U.S. Army, incharge (1876)—Coleoptera of the Lake SuperiorRegion (1878)—On the lists of Coleoptera pub-lished by the geological survey of Canada, 1842-1888 (1890);—Insect Fauna of the Mount DesertRegion (1927)—Insects of North Carolina (1938)—List of Beetles of South Dakota (1975)—and fi-nally a pragmatic, zoogeographic tally, PrecinctiveInsect Species In Florida (1995). Government tax-onomists, both federal and state have among theirresponsibilities the maintenance of reference col-lections and lists of all the insect species that occurin their districts. For example, the late Arizona Co-leopterist and Professor Floyd Werner had a com-puter list of all of the beetle species that occur inhis State; he gave me a printout so that I could addhis firefly records to my distribution maps, and aspecimen of a rare species I had not previouslybeen fortunate enough to acquire!

Arguably the most important firefly list gener-ator before the 20th Century was John L. LeConte,a Civil War surgeon and medical inspector withthe rank of lieutenant-colonel, who was connectedwith the Academy of Natural Sciences in Philadel-phia and at one time was the Assistant Director ofthe U.S. Mint in Philadelphia (Mallis 1971:242).He was an energetic field-and-bench man, andjourneyed to the far west in 1843, then collectedaround Lake Superior in 1844, working his wayalong the south shore and on to the source of theMississippi River in Minnesota; in 1845 he trav-eled up the Platte River to Fort Laramie and thento the Rocky Mountains, and so on (Mallis: 245).I’d like to retrace his Lake Superior trip with aclass of dedicated taxonomy students and find oneof his now-puzzling if not gone fireflies. LeConte’sname will be recognized even by beginning firefly-ers as the author of many North American species(Table 5). “‘LeConte was the greatest entomologistthis country has yet produced’ [Scudder 1886] . . .not because he named almost five thousand spe-

TABLE 4. A SELECTION OF QUAINT, PERHAPS FORGOTTENAND OFTEN AGGRAVATING (TO A HURRY-UPWORKER) COLLECTING LOCALITIES THAT AP-PEAR ON ARCHIVED INSECT SPECIMENS, FROMVARIOUS SOURCES. SEVERAL “LOST” ONESLISTED IN TOWNES AND LINNA (1963) HAVEBEEN FOUND BY FIREFLYER STUDENTS FORTHEIR LAB REPORTS VIA INTERNET REFER-ENCES. AN ASTERISK INDICATES THOSE THATTOWNES AND LINNA LOCATED; A ‡ INDICATESCOLLECTING LOCALITIES OF FIREFLYER H. S.BARBER NEAR WASHINGTON DC.—IT MUST BENOTED THAT EVEN MODERN LISTS AND SPECIESDESCRIPTIONS TODAY MUST SOMETIMES BEMADE VAGUE OR ENCRYPTED WITH RESPECT TOLOCALITY BECAUSE SUBSEQUENT COLLECTORSHAVE BEEN KNOWN TO TOTALLY ERADICATE LO-CAL POPULATIONS OF PRIZED AND POPULARGROUPS, FOR COMMERCE.


cies of beetles, but because he showed their sys-tematic relationships and pointed the way to thescientific classification of American insects.” (Mal-lis:242). Today LeConte’s beetle collection remainsso important as a reference and archive for name-vouchering specimens in Coleoptera taxonomythat it is specially housed in steel cabinets in Har-vard’s Museum of Comparative Zoology.

In the 20th Century, after the exchange of spec-imens and literature was improved, two majorfirefly lists were published. These lists representthe most comprehensive and exhausting insect

listing project that one can imagine that a taxon-focused individual might undertake, especially be-fore the age of computers. In 1909 Ernest Olivierand again in 1966 Frank McDermott attempted tolocate all of the formal species descriptions thatwere ever published for fireflies anywhere in theworld. Each, in turn, with McDermott building onOlivier’s work, tried to determine which assignednames had date seniority, had been not used pre-viously for other species, and were based on ade-quate descriptions of archived, name-holdingspecimens of record. Note the synonymies indi-cated and footnotes in the excerpt from the Lengcatalogue in Table 3; imagine attempting this forall firefly names in the literature. The Olivier/Mc-Dermott mission, in short, was to straighten outfirefly book-keeping since the beginning of zool-ogy’s “official” species-naming system (1758).Whew! Such a mission is of course impossible, butfireflyers will keep working on it. In their respec-tive editions of the Coleopterorum Catalogus, Pars9, Lampyridae. Olivier listed 1097 species in 53genera and McDermott 1891 species in 92 genera.Any contemporary, computer-assisted, globe-round project that attempts to list all of the organ-isms in the world must surely rely heavily uponMcDermott’s edition for the Lampyridae.

Value and Problems of Species Listing. Thereare a number of reasons why someone might wantto have a list of insects that occur in a given geo-graphic region. Species lists with key referencesare useful and a necessity for students who wish tostep beyond a field guide in preparing term papersor graduate theses; conservationists want to knowwhat species occur in an area they are trying tosave or restore, or to determine whether there havebeen species changes through local extinctions orthe introduction of exotics; economic entomologistsneed to know what non-target species might beharmed by or could be useful for a prospective pestcontrol measure, or when exotic species were firstreported; Sunday supplement writers may com-pare the number of species that occur in their area,say, Central Park NYC, with the numbers found inPelham Bay Park in the Bronx, or in the naturalarea on Staten Island that is connected with a mu-seum where there was considerable taxonomic ac-tivity early in the 20th century—Charles Leng wasthe director of this museum when he first pub-lished his catalogue of coleoptera in 1920. In “ANatural History of the Chicago Region” Joel Green-berg (2002) included a firefly tally for his region, inconnection with efforts made there for the restora-tion of lost ecologies.

Interestingly, through examination of earlylists and other old literature that has now beencollected in modern libraries and archived mu-seum specimens, an entomology student todayhas at hand better information for the state of hisregion’s taxonomy in 1875 than did taxonomistsof this postwar era themselves. In fact, complete

TABLE 5. FIREFLY SPECIES THAT JOHN L. LECONTE AU-THORED THAT ARE TODAY CONSIDERED VALID,WITH THE YEAR THEY WERE ORIGINALLY DE-SCRIBED. NAMES IN BRACKETS SHOW ORIGINALPLACEMENT. DATA ARE FROM MCDERMOTT’S1966 “COLEOPTERORUM CATALOGUS,” PARS 9.


post facto “contemporary” lists can be made todaythat could never have been available to the taxon-omists in their own time. Species lists from the19th Century seem absurdly useless when com-pared with what we think we know today, but hadthese early prospectors known of this they wouldhave carried on anyway, for fun, the solitude orcompanionship of the chase—still an option andchoice—and perhaps the opportunity for a sciencepresentation at their society’s meeting and thepersonal gratification of seeing their contributionin their society’s journal. They would presumethat their lists would have been of some interestand value to those who followed. To save an oth-erwise wasted summer wouldn’t you enjoy identi-fying and listing the insects emerging from aprostrate white pine log, one dating and rottingsince the great forest mowdown of the northwoods around the end of the 19th century, or liv-ing in one square meter of beach grass along theshore of a kettle (lost ice-block) lake in Minnesota,and then tell someone about it? Today you couldrecord trophic levels with photographs and videosof interactions to illustrate your talk.

Judging from the view at the beginning of the21st Century, early makers of comprehensive in-sect lists can be seen to have had the disadvan-tages of: (1) inadequate sampling, (2) little or nocommunication with each other and limited accessto taxonomic literature, (3) inadequately describedspecies, (4) few and poor keys to described species,and (5) a great underestimation of the number ofinsect species present, which was partly due to ig-norance of the profusion of noninterbreeding, sym-patric (sibling/cryptic) look-alike species. For themany “species problem” could be remedied whenmore “material” was available, that is, more speci-mens were acquired for their collections.

Today’s list makers would perhaps agree thatthere are at least three major species problemsthat taxonomists need to deal with. First, we needdescriptions and workable illustrated keys, with a“conservative reverence” toward maintaining afunctional yet stable nomenclatural system, sothat taxonomists and their client biologists andothers can know with as much precision as possi-ble, what specific organisms other workers are re-ferring to when they present results of theirstudies. Second, taxonomists and biologists need tobe mindful of the fact that sometimes, often, orusually there are many more independent geneticpopulations (biospecies), than can be estimatedfrom contemporary, routine, omnispective meth-ods. A third aspect of today’s species problem iscomparable to one listed for 19th Century taxono-mists, except that we already have “the mate-rial”—millions, perhaps tens of millions of archivedspecimens—but much of it remains unsorted andundescribed because of the shortage of taxonomistsand qualified collaborators. In the “good old days”much insect taxonomy was performed as an avoca-

tion by gentle folk with other occupations, physi-cians, parsons, pharmacists, and “pedagogists”;perhaps we are heading in that direction now,which would be a good thing, but professionals per-haps remain too leisurely in recruitment.

A modern and growing problem is how to dealwith the quantity and complexity of informationthat is relevant to taxonomic practice and essen-tial for improving the quality of lists. Once upon atime a taxonomist made species “available” toothers merely by describing and naming new spe-cies, and identifying specimens for them. The lat-ter especially was understood as their in-servicefunction for biological research—Table 6 is a sam-pler of my institutional “firefly clients.” Today it isnecessary for taxonomists to keep track of andsynthesize much more new information related totheir science, and taxa, and serve more scientistsrepresenting a broader range of disciplines. Fur-ther, today identifications are often much moretime-consuming and difficult because, (1) evermore of the species are “cryptic” and difficult todistinguish from close relatives, and (2) tenderedspecimens must be maintained and handled withmore care, with sterile technique and regard fordamage that can result even from room tempera-

TABLE 6. A PARTIAL LISTING OF THE INSTITUTIONS REP-RESENTED AMONG JEL “CLIENTS.” WHEN SUCHARCHIVES HAVE SERIES OF SPECIMENS THATHAVE BEEN IDENTIFIED BY A CURRENT AUTHOR-ITY OF THE TAXON THEY CAN IN TURN PROVIDETAXONOMIC IDENTIFICATIONS AND SERVICESFOR REGIONAL STUDENTS AND OTHER RE-SEARCHERS. DETERMINATION LABELS OF THELATE JOHN WAGONER GREEN, A MASTER FIRE-FLY TAXONOMIST AT THE CALIFORNIA ACAD-EMY OF SCIENCES, ARE ATTACHED TOSPECIMENS IN MANY OF THESE INSTITUTIONS.


tures, to avoid contaminating or degrading DNAand other chemicals. Of course there is a taxo-nomic motivation for such “routine bench IDs”,because new information can provide insight forunderstanding biospecies, and for further refin-ing species lists. A taxonomist is sometimes ableto provide ID guidance for prospective research—say, suggest which firefly species is the best suitedto study the impact of a flash-seeking predator onspecies-specific signaling behavior, and which ofits “sibling species” in another region would makea good comparison; or tell a DNA seeker studyingthe variation in the code for a particular protein,the location of suitable populations for examina-tion. Some taxonomists will collect critical speci-men samples for clients (but I always declinerequests for fireflies to be released at weddings).

It comes down to this: Every insect taxon needsa fanatical specialist who will make a life of pas-sionate proprietary concern for his “own personal”charges (explain that to a university administra-tor of the business model ilk). Words of Pavlov, asquoted by Blatchley, make a connection here: “Re-member that Science demands from a man all hislife. If you had two lives that would not be enoughfor you. Be passionate in your work and yoursearchings.” The taxon that an individual taxono-mist so nurtures may be a genus, family, or order,depending on its size, complexity, amount of con-temporary research on the organisms, and per-sonal interest. Indeed, such specialists, alongwith field books and other records of archivedspecimen collections, are virtually part of collec-tions themselves, though more poorly archived(underappreciated) and obviously, because theyare the caregivers themselves, shorter lived. Keepin mind that such specialists must see to the pro-fessional education and technical training of theirsuccessors. In my view, every biology departmentin U.S. academies should have an insect taxono-mist who will take professional care of a personaltaxon, teach a section and lab of introductory or-ganismic biology to freshmen, and teach a special-ized course and seminar in taxonomic/evolutionary biology. Realistically and gloomily,though it is a collective responsibility that all ac-ademic institutions have an obligation to share, Iwill bet that it is quite unlikely to happen.

Contemporary Firefly Lists. Some regions ofNorth America have many species of fireflies, andlocal naturalists will spend many years trying toresolve the easy ones and outline and begin to un-ravel the problems presented by others. McDer-mott and Barber spent a half-century watchingthe fireflies around the Chesapeake Bay and lowerPotomac River, but they left much for us to dis-cover (Fig. 3). Florida and Georgia, which are themost firefly-rich States, currently have 56 listedspecies, including those formally named and thosewith informal working nicknames (Lloyd 1997). Atthe other extreme, Alaska apparently has only one

species, a member of the Ellychnia corrusca clan;Hawaii briefly had three borrowed species, all ofwhich seem to have disappeared soon (weeks?) af-ter their introduction in the 1950s (Table 10).Some regions have only a few species, and presentfew if any problems in getting a fairly straight-for-ward general outline. North Dakota, for example,apparently has about 20 species in 6 genera, andeven the species of Photuris present few problems.On the other hand, the list for Bay County, Floridahas 34 species in 9 genera, including 5 unnamedspecies (Table 7, Fig. 4A).

The species list for a county in North Dakotamakes an interesting contrast with the BayCounty list (cf Tables 7 and 8). The species inthese two as now understood and except for Pho-turis, can be identified though morphological de-scriptions and keys in the literature. The threePhoturis species listed for Stutsman County (Ta-ble 8, Fig. 4B) can easily be identified in the fieldfrom Barber’s monograph (1951), and the mor-phological, behavioral, and ecological notes heprovided—given that one knows which three Pho-turis are present. However, the situation in BayCounty is much more difficult because 9 or morePhoturis species are present. The mere listing ofthe Photuris fireflies of Bay County will not alloweven a persistent and dedicated user to recognizewhat a list maker referred to. A supplementaryflash pattern chart is needed, or better yet, a keyto flash patterns such as the one for east-centralAlabama (Table 9; Lloyd 1990b). A student natu-ralist guided by an informed teacher could makesuch a chart and key for his region in two or threesummers, but resolving the name problems wouldtake much longer.

Fig. 3. The domain of fireflyer pioneers F. A. McDer-mott and H. S. Barber, around the Chesapeake Bay andthe lower Potomac, with dots marking observation/col-lecting spots mentioned in their literature, and by J. W.Green in his generic revisions, from insect labels.


The difficulty a fireflyer has in actually findingpopulations in the field will depend upon a vari-ety of circumstances. Some species are common,some are rare; some appear in disturbed areas,such as creek washouts, hurricane blowdowns,under power lines, and over oldfields. Some aresecretive and occur only in isolated pockets in un-disturbed areas, and others may be nearly extinctbecause their habitat is virtually gone, withoutour knowledge that its special circumstanceseven existed. Sometimes species from extincthabitats have taken up residence in special man-made situations. For example, the long-unknown,

early-destroyed oak savannah that extendedsouthwest from Chicago (Greenberg 2002), mayhave been the original habitat of some firefliesthat are now gone too, but maybe some may still

TABLE 7. SPECIES LIST FOR BAY COUNTY, FLORIDA, IN-CLUDING ADDITIONAL DATA FOR COLLECTORSAND OBSERVERS: F-FACTORS ARE PREDICTIONSFOR THE COUNTY (AND IMMEDIATELY ADJA-CENT AREAS), WITH 1 BEING THE EASIEST TOFIND AND 5 BEING THE MOST DIFFICULT (SEETEXT); IN INDICATES THE SPECIES IS KNOWN TOOCCUR OR IS SUSPECTED TO OCCUR IN BAYCOUNTY; EDGE INDICATES THAT THE SPECIES ISKNOWN TO OCCUR NEAR BAY COUNTY.

Fig. 4. Maps concerning distributions of fireflies dis-cussed in text: (A) Location of Bay County FL, see Table7; (B) Location of Stutsman County ND, see Table 8; (C)asymmetry of distribution rings at Cleveland OH be-cause of Lake Erie; (D) loops of dotted lines In New YorkState showing the eastern or northeastern limits ofknown distributions of 10 EDGE fireflies “approaching”New England (i.e., VT, NH, CT, MA, RI, ME).


survive along highway and railroad swards andberms—and in old, untended graveyards, a neatbit of ecology passed along to this student by aninsect curator at Ann Arbor.

The distinctiveness of flashing patterns willalso influence how quickly fireflies are found orrecognized as different entities, but once a localpopulation has been found others may be moreeasily located near by. I once discovered an unde-scribed species of Photuris in a roadside marsh ata culvert in northwestern South Carolina, butfour days later their ugly, cattail-infested puddlewas bulldozed to become a pretty reflecting pooland waterfall for a golf course. I will look for fire-flies flashing the same pattern, upstream anddown along the valley and the montane creek thatnow feeds Golfers’ Pool.

The lists for Stutsman and Bay Counties (Ta-bles 7 and 8) also suggest how difficult it may be tofind the initial population of each species. F-Fac-tors abstract a subjective judgement of whether aspecies is rare (local), common, or abundant. Itsuggests the difficulty I expect a new fireflyerwould have in finding a population—were one toambitiously set out to find it, and stick with thechase. Translations are: 1, easy, find during firstyear; 2, relatively easy, find in two years; 3, con-siderable effort required, find in five years; 4,great effort required, much difficulty, find in 10years, if at all. Fives (5’s) are species presently notknown. They may occur elsewhere but there is noreason to presume they are in the region. Or, theymay be new species, undescribed by taxonomists,

TABLE 8. SPECIES LIST FOR STUTSMAN COUNTY, NORTHDAKOTA. F-FACTORS, IN AND EDGE AS IN TABLE7. IN ALL CASES THE KNOWN RANGES OF EDGESPECIES END A FEW OR SEVERAL MILES EAST OFSTUTSMAN COUNTY.

TABLE 9. SEVERAL COUPLETS OF A PICTURED KEY TO DIS-TINCTIVE FLASH PATTERNS OF FIREFLIES INEAST-CENTRAL ALABAMA. ACTUALLY, TO BEUSEFUL FUTURE REGIONAL KEYS TO SPECIES OFPHOTURIS FIREFLIES WILL REQUIRE A COMBI-NATION OF FLASH PATTERNS AND MORPHOLOGY.


TABLE 10. A PRELIMINARY/WORKING CHECKLIST OF NORTH AMERICAN FIREFLIES. NO INFORMATION IS PROVIDED HERETO AID IN IDENTIFICATION OR FOR DETERMINING SEASONAL OR GEOGRAPHIC DISTRIBUTIONS; THE LISTMERELY INDICATES WHICH NOMINAL SPECIES IN THE LITERATURE APPEAR TO BE LEGITIMATE, HOW MANYOTHERS HAVE BEEN DISCOVERED AFTER FOUR DECADES OF FOCUSED SEARCH, AND WITH RESPECT TO BOOK-KEEPING, MAKES CHANGES IN TAXONOMIC STATUS AND INDICATES THOSE CONTEMPLATED. THE LIST ALSOREPRESENTS AN INDEX TO FILES OF DATA AND OBSERVATION SUMMARIES ON THE SPECIES, WITH ASSOCIATEDFIELD BOOKS AND SPECIMEN-IDENTIFICATION RECORDS, PHOTO-MULTIPLIER CHARTS, AND SEVERAL THOU-SAND VOUCHER SPECIMENS THAT WILL BE ARCHIVED FOR FUTURE REFERENCE. WORLD TOTALS ARE APPROX-IMATIONS USING MCDERMOTT (1966) AS A BASE; ADDITIONAL SPECIES HAVE SINCE BEEN NAMED IN SOMEGENERA BUT ARE NOT INCLUDED HERE. FOOTNOTES: (1) IF AN ESTABLISHED POPULATION IS FOUND IN NA IRECOMMEND IT BE GIVEN SPECIES RANK; (2) SPECIES HERE REMOVED FROM SYNONOMY; (3) FORMERLYVIEWED AS CONSPECIFIC WITH A CUBAN SPECIES, WILL BE DESCRIBED AS A NEW SPECIES; (4) SPECIES HEREELEVATED FROM SUBSPECIES TO FULL SPECIES.


and after one finds them it may take two or threeyears to be convinced that they are indeed new.Nor are 6’s listed—these are species that will onlybe found after new ideas and methods of lookingand analyzing are used for searching and under-standing genetic diversity and population trajec-tories through evolutionary time.

Species listed as occurring at the EDGE (Ta-bles 7 and 8) can sometimes be presumed to actu-ally occur IN, but be too rare or too habitat-restricted to have yet been found. This is relatedto the question, how regional are regional lists?How far out from the specified region does the listapply? To begin to answer this one can draw a se-ries of concentric circles around the center of aconsidered focus region. The further one travelsfrom the center the less applicable the regionaltreatment should be. Each species will drop out atsome distance from the center, but the drop-offwill certainly not be gradual nor symmetrical. Asan extreme condition in both respects, if Cleve-land, Ohio were a used as a center, the drop off tothe north, until Lake Erie someday fills and be-comes marshland, would be precipitous and con-centric rings highly asymmetrical (Fig. 4C). Ringsaround Phoenix, Arizona, for the few fireflies thatoccur there, would mask the patterns of distribu-tion that actually exist because probably only gal-lery habitats along streams will harbor mostfireflies of the region.

When listing the firefly species of New Englandone finds 22 species IN and 10 at the EDGE. Of the10 near the edge, can one presume that some actu-ally are IN but yet unfound? The 10 EDGE speciesapproach New England from the west and south-west, and based on current distribution maps theyreach their eastward limit at the Hudson River/Taconic and Green Mountains, paralleling theeastern border of New York State (Fig. 4D). Inspite of highways and their bordering grasslands(swards and berms) and vehicular traffic with mil-lions of opportunities for fast range-extendingrides in sod and plant pots, and prevailing westwinds blowing at this latitude, and considerablehabitat modification in New England that wouldseem to make at least some places livable for someof these 10 species, none have yet been seen amongthe examined collections made there for more thana century. Note that a firefly taxonomist of nomean reputation, Henry Clinton Fall, a retired sci-ence teacher and author of Photinus ignitus a spe-cies of the region, lived in Massachusetts for sometime—he also authored “A list of the Coleoptera ofthe Southern California Islands with Notes andDescriptions of New Species”! Thus, not all EDGEspecies are promising candidates for inclusion in aregional listing—but such species would seem ex-cellent candidates for ecological studies to find thelimiting factors responsible. Surely there is moreto the apparent exclusion of these EDGE speciesthan meets the eye.

I would like to incorporate many of the featuresof Willis Blatchley’s Indiana list and Rev. Henry S.Gorham’s “Biologia-Centrali Americana” (1880-1886) in the firefly list I am aiming for, but for thepresent Table 10 gives a bare bones beginning. Ithas codens for many unnamed species, and FindFactors would be useless at such a scale and withso many ecological unknowns. The next edition ofthis list promises to have distribution maps andseasonal and flash charts to aid in identification.In the meantime the centuries-long chase by fireflynaturalist/taxonomists creeps resolutely onward.

ACKNOWLEDGMENTS

I thank John Sivinski and Scotty Long for readingand commenting on the manuscript, Mike Sanford andPamela Howell for providing technical assistance withthe preparation of figures and manuscript; ProfessorsW. F. Stanley, W. H. Wagner, R. D. Alexander, T. E. Moore,and W. L. Brown for key and provoking statements anddiscussions during student days; D. Furth and H. Evansfor guidance through the beetle archives at the Museumof Comparative Zoology; and innumerable firefly stu-dents whose presence and interest over the years havebeen inspirational for viewing lists and alpha taxonomyin new and optimistic ways, and to write this Letter.Florida Agricultural Experiment Station Journal SeriesNumber R-08966.

REFERENCES(Including notes, fragmentary citations,

and a list sampler)

ADAMS, C. C. JR. 1913. An ecological study of prairie andforest invertebrates. Bulletin of the Illinois StateLaboratory of Natural History. Vol. 11, article 2[Lampyridae pp. 176 & 221].

AGASSIZ, L. 1850. Lake Superior: physical character,vegetation, and animals, with a narrative of the tourby J. E. Cabot, and contributions by other scientificgentlemen, elegantly illustrated. Gould, Kendalland Lincoln, Boston.

ANON. 1893. The classification of insects and their rela-tion to agriculture. Bulletin No. 28. University ofMinnesota Agricultural Experiment Station, Minne-sota [Lampyridae s.l. pp. 148-150].

BENNETT, JOHN B. 2001-02 (winter). On institutionalfaithfulness. Academic Questions 15(1): 77-81.

BLACKWELDER, R. E. 1967. Taxonomy; a text and refer-ence book. John Wiley & Sons, Inc. New York.

BLATCHLEY, W. 1910. An illustrated descriptive cata-logue of the Coleoptera or beetles [exclusive of theRhynchophora] known to occur in Indiana, with bib-liography and description of new species. Indianapo-lis [Lampyrinae pp. 816-824].

BLATCHLEY, W. 1930. Blatchleyana. The Nature Pub-lishing Co., Indianapolis.

BRANHAM, M. A. AND J. W. WENTZEL. 2001. The evolu-tion of bioluminescence in cantharoids (Coleoptera:Elateroidea). Florida Entomol. 84: 565-579.

BRIMLEY, C. S. 1938. Insects of North Carolina; being alist of the insects of North Carolina and their closerelatives. North Carolina Department of Agricul-ture, Division of Entomology [supplementary lists;Lampyridae pp. 153-154].


CROTCH, G. R. 1873. Check list of the Coleoptera ofAmerica, north of Mexico. Naturalists’ Agency, Sa-lem Press (F. W. Putnam & Co. proprietor), Salem,Mass. [Lampyridae s.l. pp. 74-75].

DAILEY, P. J., R. C. GRAVES, AND J. M. KINGSOLVER.1978. Survey of the Coleoptera collected on the com-mon milkweed, Asclepias syriaca, at one site in Ohio.Coleopterists Bulletin 32: 223-229.

DILLON, E. S. AND L. S. DILLON. 1961. A manual of com-mon beetles of eastern North America. Row, Peter-son and Co., Evanston, Illinois [with State andCanadian lists of beetles, pp. 852-865].

DURY, D. 1884. List of coleoptera around Cincinnati, Ohio[cited in Hamilton 894-21; earlier lists, 1879, 1882].

ELMAN, R. 1982. American pioneering naturalists; theirlives and times, exploits and adventures. WinchesterPress, Tulsa.

EMMONS, E. 1854. Agriculture of New-York: comprisingan account of the classification, composition and dis-tribution of the soils and rocks, and of the climateand agricultural production of the state; togetherwith descriptions of the more common and injuriousspecies of insects. C. Van Benthuysen, Albany. [vol-ume 5?; Chapter III, Remarks on the classification ofinsects; Lampyridae s.l. pp. 89-90].

EPSTEIN, M. E. ca 1979. The Colorado firefly project.Colorado Department of Natural Resources, Colo-rado Natural Areas Program, Contract NumberC100063. 10 p.

FAUVEL, A. 1889. Liste des coleopteres communs a L’Eu-rope et a L’Amerique du nord—apres le catalogue deM. J. Hamilton, avec remarques et additions. 8:[Lampyridae s.l. p. 145].

FENDER, K. 1961. Lampyroid Section. 3:35-43. [in] Bee-tles of the Pacific Northwest, [ed] M. H. Hatch, Univ.Pub. Biol. 16: 1-503.

FORBES, S. A. 1915. An ecological study of prairie andforest invertebrates. Bulletin of the Illinois StateLaboratory of Natural History, Vol. 11, Article 2: Ur-bana, Illinois [Lampyridae pp. 176, 221].

FROST, S. W. 1964. Insects taken in light traps at theArchbold Biological Station, Highlands County, Flor-ida. Florida Entomol. 47(2): [Lampyridae p 140].

GEISER, S. W. 1937. Naturalists of the frontier. Pub-lished by the author.

GORHAM, H. S. 1880-1886. Biologia Centrali-Americana,Insecta, Coleoptera, Vol. 3, Part 2, Malacodermata.

GREEN, J. W. 1956. Revision of the nearctic species ofPhotinus. (Lampyridae: Coleoptera). Proceedings ofthe California Academy of Sciences 28: 561-613.

GREEN, J. W. 1957. Revision of the nearctic species ofPyractomena (Coleoptera: Lampyridae). The Was-mann Journal of Biology 15: 237-284.

GREENBERG, J. 2002. A natural history of the Chicagoregion. The University of Chicago Press, Chicago.

HAMILTON, J. 1894. Catalogue of the Coleoptera com-mon to North America, Northern Asia and Europe,with distribution and bibliography. Transactions ofthe American Entomological Society. 2nd edition. 21:[Lampyridae s.l. page 361?].

HAMILTON, J. 1895. Catalogue of the Coleoptera ofSouthwestern Pennsylvania, with Notes and De-scriptions. Trans. Am. Entomol. Soc. 22: [Lampy-ridae s.l. p. 334].

HARRINGTON, W. H. 1890. On the lists of coleoptera pub-lished by the geological survey of Canada, 1842-1888. Canadian Entomologist 22: 153-160 [Lampy-ridae s.l. det. by J. L. LeConte].

HAYWORTH AND SAVAGE. Catalogue of coleoptera ofGreen Mountain, Vermont. Quart. J. Boston. Zool.Soc. II. (cited in Hamilton 1894: 21).

HENSHAW, SAMUEL. 1885. List of the coleoptera of Amer-ica, North of Mexico. American Entomological Society,Philadelphia. [Lampyridae s.l. pages 75-77+].

HUBBARD, H. G. AND E. A. SCHWARZ. 1878. List of Co-leoptera found in the Lake Superior Region [Lampy-ridae s.l. pp. 637-638].

JOHNSON, C. W. 1927. The insect fauna with reference tothe flora and other biological features. [in] Biologicalsurvey of the Mount Desert Region: 1. Wistar Insti-tute of Anatomy and Biology, Philadelphia [Lampy-ridae pp. 97-98].

KASTNER, J. 1977. A species of eternity. Alfred A. Knopf,New York.

KIRK, V. M. 1969. A list of Beetles of South Carolina:Part 1—Northern Coastal Plain. Technical Bulletin1033 [Lampyridae pp. 55-56].

KIRK, V. M. 1970. A list of Beetles of South Carolina: Part2—Mountain, Piedmont, and Southern Coastal Plain.Technical Bulletin 1038 [Lampyridae pp. 55-56].

KIRK, V. M. 1975. A list of the Beetles of South Dakota.Agricultural Experiment Station, South Dakota StateUniversity, Brookings [Lampyridae pp. 71-72].

LAVIGNE, R., R. KUMAR, AND J. A. SCOTT. 1991. Addi-tions to the Pawnee National Grasslands InsectChecklist. Entomological News 102(3): 150-164[Lampyridae p. 153].

LECONTE, J. L. 1860. Report upon the insects collectedon the survey. Report No. 1. In Zoology Report—Ex-plorations and surveys for a railroad route from theMississippi River to the Pacific Ocean.—War De-partment: Route near the forty-seventh and forty-ninth parallels, explored by I. I. Stevens, Governor ofWashington Territory, in 1853-‘55. Washington, D.C.[Lampyridae p. 18].

LECONTE, J. L. 1866. List of coleoptera collected in themountains of Lycoming County, Pa. by John L. Le-Conte, M.D. Proceedings of the Academy of NaturalSciences of Philadelphia 19: [Lampyridae p. 347].

LECONTE, J. L. 1866. List of coleoptera collected nearFort Whipple, Arizona, by Dr. Elliott Coues, U.S.A.,in 1864-65. Proceedings of the Academy of NaturalSciences of Philadelphia 19: [Lampyridae p. 348].

LECONTE, J. L. 1869. List of coleoptera collected in Van-couver’s Island by Henry and Joseph Matthews, withdescriptions of some new species. The Annals andMagazine of Natural History 4: [Lampyridae p. 371].

LECONTE, J. L. 1876. Report upon new species of co-leoptera collected by the expeditions for geographicalsurveys west of the 100th meridian, Lieut. Geo. M.Wheeler, Corps of Engineers, U.S. Army, in charge.Being extract from Appendix JJ of the Annual Reportof the Chief of Engineers for 1876. [Lampyridae s.l. p.481] Government Printing Office, Washington.

LECONTE, J. L. 1883. Lists of coleoptera collected in 1881by Dr. Bell and others, in the Lake Superior districtand in the north-west territories, east of the 112thMeridian and south of the 60th parallel [Lampyridaepages 31, 33-37]. Rept. Progress Canad. GeologicalSurvey, 1880-81-82. Appendix II to part c.

LECONTE, J. L. AND G. H. HORN. 1883. Classification ofthe coleoptera of North America. Smithsonian Misc.Coll. No. 507. Washington.

LENG, C. W. 1920. Catalogue of the Coleoptera of Amer-ica, North of Mexico. Mount Vernon, New York[Lampyridae 141-142].


LEONARD, D. L. [ed.] 1928. A list of the insects of NewYork, with a list of the spiders and certain other al-lied groups. Agricultural Experiment Station, Cor-nell University, Ithaca [Lampyridae 320-322].

LEWIS, S. M., AND J. D. MONCHAMP. 1994. Sexual andtemporal differences in phorid parasitism of Photi-nus marginellus fireflies (Coleoptera: Lampyridae).Ecology and Population Biology 87: 572-575.

LINNAEUS, C. 1758. Systema Naturae. Regnum Animale(10th ed. tomus I; L. Salvii, Holminae).

LLOYD, J. E. 1969. Flashes of Photuris fireflies: theirvalue and use in recognizing species. Florida Ento-mol. 52(1): 29-35.

LLOYD, J. E. 1979a. Mating behavior and natural selec-tion. Florida Entomol. 62(1): 17-34.

LLOYD, J. E. 1979b. Sexual selection in luminescent bee-tles. Pages 293-342 In M. Blum and A. Blum, eds.Sexual selection and reproductive competition in in-sects. Academic Press, New York, N.Y.

LLOYD, J. E. 1980. Male Photuris fireflies mimic sexualsignals of their females’ prey. Science 210(4470):669-671.

LLOYD, J. E. 1981. Firefly mate rivals mimic their pred-ators and vice versa. Nature 290: 498-500.

LLOYD, J. E. 1984. Evolution of a firefly flash code. Flor-ida Entomologist 67: 228-239.

LLOYD, J. E. 1990a. Firefly semiosystematics and preda-tion: a history. Florida Entomologist 73(1): 51-66.

LLOYD, J. E. 1990b. Checklist and keys to fireflies ofeast-central Alabama. Stridulator (F. S. Arant Ento-mology Club, Department of Entomology, AuburnUniversity 4(3): 9-21.

LLOYD, J. E. 1995. Fireflies of Arrowwood NationalWildlife Refuge and Stutsman County, North Da-kota. (prepared for the refuge).

LLOYD, J. E. 1997. Signaling with glows, flashes, andpheromones. Fireflyer Companion 1(3): 33-38.

LLOYD, J. E. 2001. On research and entomological edu-cation V: a species (c)oncept for fireflyers, at thebench and in old fields, and back to the Wisconsianglacier. Florida Entomol. 84: 587-601.

LLOYD, J. E. 2002. Fireflies of the Connecticut Valley inVermont, prepared for the Montshire Museum, Nor-wich.

LLOYD, J. E., AND S. R. WING. 1983. Nocturnal aerialpredation of fireflies by light-seeking fireflies. Sci-ence 222: 634-635.

LÖDING, HENRY PETER. 1945. Catalogue of the Beetlesof Alabama. Monograph 11. Geological Survey of Al-abama, University, Alabama [Lampyridae pp. 46-47,154, 158].

MCDERMOTT, FRANK A. 1964. Coleopterorum Catalo-gus, Pars 9, Lampyridae. Junk, Berlin.

MADGE, R. B. 1994. Who was Herschel? The CanadianEntomologist 126: 543-548.

MALLIS, A. 1971. American Entomologists. Rutgers Uni-versity Press, New Brunswick.

MARVIN, D. E., JR. 1965. Fireflies (Lampyridae) of Ohio.Ann. Entomol. Soc. Am. 65(1): 37-42.

MAYDEN, R. L. 1997. A hierarchy of species concepts: thedenouement in the saga of the species problem. Pp.381-424. In M. F. Claridge, H. A. Dawah, and M. R.Wilson [eds.], Species: the units of biodiversityChapman and Hall, London.

MELSHEIMER, F. E. 1853. Catalogue of the described co-leoptera of the United States Lampyridae. Smithso-nian Institution, Washington [Lampyridae s.l. pp.76-80].

NISHIDA, G. M. [ed.]. 1994. Hawaiian terrestrial arthro-pod checklist, 2nd edition. Hawaii Biological Survey,Contribution No. 94-04. Bishop Museum TechnicalReport No. 4 [no Lampyridae, only one Cantharoi-dea, an adventive Cantharidae; three species of1950s lampyrid biocontrol imports not listed].

OLIVIER, E. 1909. Coleopterorum Catalogus, Pars 9,Lampyridae. Junk, Berlin.

OSBORN, H. 1937. Fragments of entomological history;including some personal recollections of men andevents. Published by the author, Columbus.

PACKARD, 1871. Coleoptera in Labrador. Ann. Rep. Pea-body Acad. Sci. (cited in Hamilton 1894, 21).

PEATTIE, D. C. 1936. Green laurels. Simon and Schuster,New York.

PORTER, C. M. 1986. The eagle’s nest; natural historyand American ideas, 1812-1842. Univ. AlabamaPress, USA.

REINECKE, 1880. List of Coleoptera of Buffalo, New York.Zesch. and Reinecke (cited in Hamilton 1894, 21).

SCHAEFFER, C. 1908. List of the Lampyridae [s.l.] fromthe Huachuca Mountains, Arizona, and descriptionsof new species. Journal of the New York Entomolog-ical Society 16: 61-67.

SCHWARZ, E. A. 1878. The Coleoptera of Florida. Proc.Am. Philosophical Soc. 17: 35-36.

SMITH, J. B. 1899. Insects of New Jersey: A list of thespecies occurring in New Jersey, with notes on thoseof economic importance. 27th Annual Report of theState Board of Agriculture (prefatory statementdated March 19th, 1900). Trenton, MacCrellish &Quigley, State Printers.

SMITH, R. F., T. E. MITTLER, AND C. N. SMITH. 1973. His-tory of entomology. Annual Reviews Inc., Palo Alto.

SMITH, R. C., E. G. KELLY, G. A. DEAN, H. R. BRYSON,AND R. L. PARKER. 1943. Common Insects of Kansas.Kansas State Board of Agriculture. Vol. 62, No. 255:440 p. [Lampyridae pp. 280-281].

SORENSEN, W. C. 1995. Brethren of the net; Americanentomology, 1840-1880. Univ. of Alabama Press, Tus-caloosa.

STEWART, G. R. 1970. American Place-names; a conciseand selective dictionary for the continental UnitedStates of America. Oxford Univ. Press, New York.

TOWNES, M., AND E. LINNA. 1963. The location of someobscure entomological collecting localities in theUnited States and Canada. Proc. Entomological So-ciety of Washington 65: 233-246.

ULKE, H. 1902. List of the beetles of the District of Co-lumbia. Proc. United States National Museum, Vol.25 (1275): [Lampyridae s.l. pp. 22, 47].

WERNER, FLOYD G. 1988. Arizona Insects in Universityof Arizona Collection. Computer file [Lampyridaelist to jel dated 19 October 1988].

WILSON, H. F., AND M. H. DONER. 1937. The historicaldevelopment of insect classification. (published bythe authors)

WING, S. R. 1985. Prolonged copulation in P. macder-motti with comparative notes on Photinus collus-trans (Coleoptera: Lampyridae). Florida Entomol.68: 627-634.

WING, S. R. 1991. Timing of Photinus collustrans repro-ductive activity: finding a mate in time (Coleoptera:Lampyridae). The Coleopterists Bulletin 45: 57-74.

WING, S. R., J. E. LLOYD, AND T. HONGTRAKUL. 1983.Male competition in Pteroptyx fireflies: wing-coverclamps, female anatomy, and mating plugs. FloridaEntomol. 66: 86-91.

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