Euphoria sepulcralis (F., 1801)

Orozco, Jesús, 2012, Monographic Revision of the American Genus Euphoria Burmeister, 1842 (Coleoptera: Scarabaeidae: Cetoniinae), The Coleopterists Bulletin (mo 11) 66, pp. 1-182 : 34-36

Euphoria sepulcralis (F., 1801)

(Appendix 4: Fig. 13 )

Cetonia sepulcralis Fabricius 1801: 156 . Original combination.

Type not seen. In loan from Zoologisches Museums der Christian- Albrechts- Universität zu Kiel ( Germany) to the Zoological Museum of Copenhagen ( Denmark).

Cetonia reichi Gory and Percheron 1833: 210 . Synonym.

Holotype at MHNG, examined.

Euphoria limatula Janson 1881: 585 . New synonymy.

Holotype at RMNH, examined.

Euphoria nitens Casey 1915: 320 . New synonymy.

Lectotype at USNM designated by Hardy (2001), examined. Two paralectotypes at USNM, one examined.

Euphoria sepulcralis kansana Casey 1915: 321 . Synonym.

Lectotype at USNM designated by Hardy (2001), examined. Fifteen paralectotypes at USNM, one examined.

Euphoria sepulcralis crinitula Casey 1915: 321 . Synonym.

Holotype at USNM, examined.

Euphoria sepulcralis floridana Casey 1915: 321 . Synonym.

Lectotype at USNM designated by Hardy (2001), examined. Fourteen paralectotypes at USNM, one examined.

Euphoria sepulcralis cuprascens Casey 1915: 322 . Synonym.

Holotype at USNM, examined.

Euphoria appalachia Casey 1915: 322 . Synonym.

Lectotype at USNM designated by Hardy (2001), examined. Six paralectotypes at USNM, not examined.

Euphoria scolopacea Casey 1915: 322 . Synonym.

Lectotype at USNM designated by Hardy (2001), examined. Twenty-five paralectotypes at USNM, one examined.

Euphoria oxysternum Casey 1915: 323 . Synonym.

Holotype at USNM, examined.

Description (n = 2,754). Length 9.2–13.5 mm; width 5.5–8.3 mm. Color: Surface shiny, dark brown to black with variable green or violaceous reflections. Pronotum frequently with whitish, cretaceous band on lateral margin. Elytra with small to moderate, reniform, vermiform, or irregular, cretaceous markings; markings whitish to yellowish. Mesepimera, metasternum, metacoxae, and sides of abdominal sternites frequently with whitish, cretaceous markings. Pygidium variably covered by tomentum, cretaceous markings frequently on anterolateral margins. Head: Frons densely punctate; punctures moderate in size, round, deeply impressed, at times coalescent; sparsely to densely setose; setae moderate to long, whitish to yellowish. Clypeal surface densely to moderately densely punctate to punctatostrigate; punctures moderate in size, round, coalescent, glabrous to moderately densely setose; setae yellowish to whitish, short to long; subquadrate, lateral margins weakly to moderately raised, lateral declivity moderately to strongly developed, apex vaguely to weakly reflexed, truncate in dorsal view, vaguely to weakly sinuate in frontal view. Antennal club in males slightly shorter than stem, much shorter in females. Pronotum: Surface moderately densely to densely punctate; punctures round to lunulate, moderate in size, denser and confluent towards apex and sides, sparsely to densely setose; setae short to moderate, yellowish to whitish. Scutellum longer than wide, sparsely to densely punctate. Base in front of scutellum moderately to strongly emarginate. Elytra: Surface densely punctate; punctures lunulate and irregular, moderate in size, striae frequently with 3–4 rows of punctures and grooves, costae well defined, frequently punctate. Posterior half of sutural costa raised in lateral view. Surface glabrous to moderately densely setose; setae minute to short, whitish to yellowish. Legs: Protibial teeth well developed, apical tooth oblique, apical and medial teeth closer to each than to basal tooth. Metatibial carina moderately developed. Metatibiae vaguely to weakly expanded apically, internal metatibial spur slightly thicker in females. Venter: Mesometasternal process extended well beyond mesocoxae, process glabrous on mesosternal lobe, apex rounded, flat, or slightly pointed. Mesepimera, metasternum, and metacoxae setose, setae as on legs. Metasternum rugose, setose laterally, glabrous, impunctate at middle. Median sulcus vague to weakly evident. Abdominal sternites densely punctate; punctures small, bearing setae; sparsely setose laterally, glabrous to sparsely setose medially; setae distributed along entire width of segment, moderate to long, whitish to yellowish. Pygidium: Subconcentrically striate, striae discontinuous, deeply to shallowly impressed; glabrous to moderately densely setose, setae short to moderate, whitish to yellowish; apex frequently rounded in males, moderately pointed in females. Male genitalia: Parameres as in Fig. 13c .

Diagnosis. Euphoria sepulcralis is separated from the other species in the group based on the length of the antennal club being shorter than stem in both sexes, weakly raised apex of the clypeus, pronotum densely punctate, and form of the parameres. This species is highly variable in color (dark brown to black with variable green or violaceous reflections) and the extent of the pronotal cretaceous markings.

Notes. Euphoria sepulcralis is frequently dark violaceous in Texas and northwestern Mexico ( Fig. 13e ). This morphotype was previously known as E. sepulcralis nitens Casey, 1915 . Other than the different color, generally larger body size, and distribution, I cannot find any other character to justify this subspecies. Dark violaceous specimens are rare outside of Texas, but are nevertheless present. This is another example of a North American species that exhibits a darker color form in Texas populations (together with E. fulgida and E. kernii ). The cause of this phenomenon is unknown to me. Specimens from Florida and the Cayman Islands were observed to be more coarsely punctate.

Taxonomic History. Fabricius (1801) described E. sepulcralis in Cetonia . Gory and Percheron (1833) described Cetonia reichi based on specimens from Mexico. Janson (1881) described a dark, mislabeled specimen from Guatemala as E. limatula . Casey described E. nitens , E. appalachia , E. scolopacea , E. oxysternum and four subspecies from the United States based primarily on coloration, body size, and geographic locality. Based on the study of 2,754 specimens, including the types, I place E. limatula and E. nitens in synonymy with E. sepulcralis .

The misspelled version of the name “ E. sepulchralis ” was apparently used for the first time by Gory and Percheron (1833). The name was subsequently used by Blanchard (1850), Lacordaire (1856), Casey (1915), and other authors. The correct version of the name ( E. sepulcralis ) was used by Burmeister (1842), Harold (1869), Horn (1880), and Schenkling (1921).

Natural History. Euphoria sepulcralis is frequently collected in a variety of habitats. Adults have been collected on Acanthaceae : Asystasia gangetica (L.) T. Anders; Aceraceae : Acer rubrum L. foliage; Agavaceae : Yucca sp. ; Amaryllidaceae : Crinum americanum L.; Anacardiaceae : Mangifera indica L., Rhus copallina L., Rhus glabra L., Schinus terebinthifolius Raddi ; Annonaceae : Asimina reticulata Chapm. , Asimina triloba (L.) Dunal; Apiaceae : Anethum sp. , Daucus carota L. flowers, Eryngium yuccifolium Michx. , Foeniculum vulgare Mill. , Heracleum maximum Bartr. , Oxypolis rigidior (L.), Pastinaca sativa (L.), Pimpinella anisum L., Sium suave Walt. ; Apocynaceae : Allamanda sp. , Apocynum floribundum Greene flowers; Araceae : Philodendron sp. ; Araliaceae : Polyscias guilfoylei (Bull) , Schefflera actinophylla (Endl.) Harms , Schefflera arboricola (Hayata) Merr. foliage; Arecaceae : Adonidia sp. , Cocos nucifera L. fermenting wound, Phoenix canariensis Hort. ex Chabaud. , Pritchardia sp. , Sabal palmetto flowers (Walt.), Serenoa repens (Bartr.) , Veitchia merrillii (Becc.) ; Asclepiadaceae : Asclepias incarnata (L.); Asteraceae : Ambrosia artemisiifolia L., Aster lanceolatus (Willd.) , Aster sp. blossom, Baccharis angustifolia Michx. , Baccharis halimifolia L., Bidens bipinnata L., Bidens sp. , Carduus ochrocentrum Greene , Carduus spinosissimus Walter , Cirsium odoratum (Muhl.) , Cirsium vulgare Savi , Dahlia sp. , Dendranthema x grandiflorum Kitam, Erigeron quercifolius Lam. , Erigeron sp. flowers, Eupatorium capillifolium (Lam.) , Eupatorium perfoliatum (L.), Eupatorium serotinum Michx. , Flaveria linearis Lagasca , Helianthus sp. , Pseudogynoxys chenopodioides (Kunth) Cabr. , Solidago canadensis (L.), Solidago sp. flowers, Stokesia sp. , Tagetes erecta L.; Bombacaceae : Bombax sp. ; Brassicaceae : Brassica juncea (L.), Brassica oleracea L., Lobularia maritima (L.); Burseraceae : Bursera simaruba (L.); Cactaceae : Cereus sp. fruit, Opuntia sp. ; Caprifoliaceae : Viburnum odoratissimum Ker. - Gawl.; Clusiaceae : Hypericum fasciculatum Lam. ; Combretaceae : Bucida buceras L., Terminalia catappa L.; Convolvulaceae : Ipomoea batatas (L.); Cornaceae : Cornus sp. ; Cyrillaceae : Cyrilla racemiflora L.; Ebenaceae : Diospyros sp. ; Ericaceae : Ceratiola ericoides Michx. ; Euphorbiaceae : Acalypha sp. , Croton capitatus Michx. , Euphorbia pulcherrima Willd. ; Fabaceae : Colvillea racemosa Crotalaria sp. , Desmodium tortuosum (Sw.) , Phaseolus lunatus L., Vigna unguiculata (L.); Fagaceae : Castanea mollissima Blume , Castanea pumila Mill. , Quercus laevis Walt. , Quercus laurifolia Michx. , Quercus palustris Muenchh. ; Ginkgoaceae : Ginkgo biloba L. stem; Juglandaceae : Carya illinoensis (Wangenh.) ; Lamiaceae : Physostegia sp. ; Lauraceae : Persea americana Mill. ; Lythraceae : Lagerstroemia indica L.; Magnoliaceae : Magnolia sp. ; Malvaceae : Abelmoschus esculentus Moench , Alcea rosea L., Hibiscus elatus Sw. , Hibiscus lasiocarpos Cav. , Hibiscus rosasinensis L., Gossypium sp. blooms; Melanthiaceae : Zigadenus densus (Desr.) ; Moraceae : Ficus benjamina L., Ficus carica L. fruit, Ficus retusa L.; Musaceae : Musa sp. ; Myrtaceae : Callistemon viminalis (Sol. ex Gaertn.) , Psidium guajava L.; Oleaceae : Fraxinus caroliniana Mill. , Ligustrum japonicum Thunb. , Ligustrum lucidum Ait. , Ligustrum sinense Lour. ; Onagraceae : Galpinsia hartwegi (Benth.) , Ludwigia sp. ; Orchidaceae : Cattleya sp. , Dendrobium sp. ; Oxylidaceae: Averrhoa carambola L.; Papaveraceae : Argemone platyceras Link and Otto ; Pinaceae : Pinus clausa (Chapm.) , Pinus elliottii Engelm. ; Poaceae : Cenchrus sp. , Cortaderia selloana (Schutt) , Z. mays ; Polygonaceae : Coccoloba uvifera (L.); Rhamnaceae : Ceanothus americanus L. flowers, Condalia obtusifolia (Hook.) ; Rosaceae : Aronia arbutifolia (L.), Crataegus sp. , Malus domestica Borkh. , Malus sp. decaying fruit, Prunus persica (L.), Pyracantha coccynea Roem , Pyrus communis L., Rosa sp. , Rubus allegheniensis (Porter) ; Rubiaceae : Borreria sp. , Spermacoce verticillata L.; Rutaceae : Citrus aurantifolia (Christm.) , Citrus limon (L.), Citrus reticula Blanco , Citrus sinensis (L.), Citrus x paradisi (L.), Fortunella margarita (Lour.) , Xanthoxylum pterota Kunth , Citrofortunella microcarpa (Bunge) ; Salicaceae : Salix sp. sap flow; Sapindaceae : Litchi chinensis Sonn. ; Scrophulariaceae : Penstemon cobaea Nutt. ; Simaroubaceae : Simarouba glauca DC. ; Solanaceae : Capsicum annuum L., Grossum sp., Lycopersicon esculentum Mill fruit, Solanum melongena L.; Staphyleaceae : Staphylea trifolia (L.); Verbenaceae : Lantana sp. ; and Vitaceae : Vitis sp.

In addition, adults have been found feeding on honey in bee hives, in bromeliads, and in millet ( Thomas 1998). In a few instances, individuals were numerous enough to be considered pests in corn, onions, and other crops by homeowners ( Halbert 1996).

Adults have been attracted to baited and unbaited pitfall traps, sugar bait traps, fruit fly traps, Japanese beetle traps, molasses traps, flight intercept traps, sticky traps, and light traps. Landolt (1990) found adults were attracted to isopropyl alcohol, and Cherry and Klein (1992) found them attracted to various aromatic compounds. Habitats include deciduous forest, prairies, pine-oak forest, and sand flats.

In temperate climates, the overwintering stage is the adult, emerging in the summer or fall and flying again the following spring ( Hayes 1925; Ritcher 1945; Ratcliffe 1991; Ratcliffe and Paulsen 2008). Adults were observed to be thanatotic when disturbed ( Ratcliffe and Paulsen 2008).

Adults have been observed burrowing in cow manure, possibly looking for a place to oviposit. Hayes (1925) studied the life cycle of the species and described the eggs. The larval stage was found to last between 53–79 days and the pupal stage between 9–23 days. The larval stage was described by Ritcher (1945, 1966). Larvae have been found beneath dead sod or manure ( Ritcher 1945), sandy soil containing oak roots and decaying oak leaves, nests of Geomys bursarius (Shaw) , and feeding on roots of Paspalum vaginatum Swartz. (Poaceae) on a golf course ( Buss 2004).

Temporal Distribution. January (2), February (32), March (126), April (346), May (649), June (214) July (98), August (145), September (364), October (110), November (13), December (16) ( Fig. 13f ).

Geographic Distribution. The Bahamas, the Cayman Islands, northeastern Mexico, and eastern United States ( Fig. 13g ).

Specimens Examined (2,754). Type material: Cetonia reichi Gory and Percheron, 1833 ; holotype at MHNG labeled “Gory/ TYPE// sepulcralis/ F/ Reich GP/ Am. bor// Coll. Melly ” and my holotype label. Euphoria appalachia Casey, 1915 ; lectotype male at UNSM labeled “ Southern Pines /NC/ A. H. Manee // CASEY /bequest/1925// appalachia. 2/ PARATYPE USNM/ 48679 // LECTOTYPE / Euphoria / appalachia Csy. /By A.R. Hardy/ 1978” , paralectotype male at UNSM labeled “N.E./U.S// CASEY/bequest/1925// TYPE USNM/ 48679// appalachia/Csy// PARALECTOTYPE / Euphoria / appalachia Csy. /By A.R. Hardy ‘78”. Euphoria cuprascens Casey, 1915 ; holotype male at UNSM labeled “Medora, Ks./ W. Knaus// CASEY/bequest/ 1925// TYPE USNM/ 48676// cuprascens/ Csy”. Euphoria crinitula Casey, 1915 ; holotype male at USNM labeled “ S. Ill// CASEY/bequest/1925// TYPE USNM/ 48678// crinitula/ Csy”. Euphoria sepulcralis floridana Casey, 1915 ; lectotype female at UNSM labeled “Jackson/ ville FLA// CASEY/bequest/1925// TYPE USNM/ 48677// floridana/ Csy// LECTOTYPE / Euphoria /sepulcralis/ floridana/ sy./By A.R. Hardy ‘78”; paralectotype male at USNM labeled “Palm beach/ Fla// CASEY/ bequest/1925// floridana. 2/ PARATYPE USNM/ 48677// PARALECTOTYPE / E. sepulcralis / floridana Csy./ A.R. Hardy ‘78”. Euphoria sepulcralis kansana Casey, 1915 ; lectotype male at USNM labeled “Manhattan/ Ks// CASEY/ bequest/ 1925// kansana. 3/ PARATYPE USNM/ 48675// LECTOTYPE / Euphoria /sepulcralis/ kansana Csy./By A.R. Hardy ‘78”; paralectotype male at USNM labeled “Manhattan/ Ks// CASEY/ bequest/ 1925// kansana. 2/ PARATYPE USNM/ 48675// LECTOTYPE / Euphoria /sepulcralis/ kansana Csy./By A.R. Hardy ‘78”. Euphoria sepulcralis nitens Casey, 1915 ; lectotype male at USNM labeled “melancholica/ Gory// CASEY/bequest/1925// TYPE USNM/ 48683// nitens/ Csy// Euphoria / nitens/ Casey// LECTOTYPE / Euphoria / nitens Csy. / By A. R. Hardy 1978”; paralectotype male at USNM labeled “Tex// CASEY/ bequest/ 1925// CASEY determ/ nitens – II// PARALECTOTYPE / Euphoria / nitens”. Euphoria sepulcralis oxysternum Casey, 1915 ; holotype male at USNM labeled “Ind// CASEY/ bequest/ 1925// TYPE USNM/ 48681// oxysterna/ Csy”. Euphoria sepulcralis scolopacea Casey, 1915 ; lectotype female at USNM labeled “Black/ Mts NC// CASEY/ bequest/ 1925// scolopacea/ Csy// TYPE USNM/ 48680// LECTOTYPE / Euphoria / scolopacea/ CASEY = By/ A.R. Hardy”; paralectotype male at USNM labeled“ Southern Pines/ NC// CASEY/ bequest/ 1925// scolopacea. 2/ PARATYPE USNM/ 48680// PARALECTOTYPE ” . Other material: CAYMAN ISLANDS (5): GRAND CAYMAN. Boatswains Point (5). BAHAMAS (2): ELEUTHERA: Governor’ s Harbor (1); NEW PROVIDENCE: Coral Harbour (1). MEXICO (21): CHIHUAHUA: Anáhuac (1); NUEVO LEÓN: China (9), Monterrey (2); TAMAULIPAS: Ciudad Victoria (1), Gómez Farías (1), Matamoros (1); NO DATA: “ Mexico ” (6). USA (2,712): ALABAMA. Baldwin Co.: Fairhope (4); Clarke Co.: Salt Mountain (1); Cullman Co.: Cullman (1); Dallas Co.: Selma (1); Madison Co.: Huntsville (43); Mobile Co.: Delchamps (2), Grand Bay (1), Mobile (7), Whistler (1); Shelby Co.: Alabaster (1), Helena (1); Talladega Co.: Coleta (1); No data (13). ARKANSAS. Chicot Co.: Lake Village (1), no data (2); Crawford Co.: Lee Creek (1), Mountainburg (1); Hope-Hempstead Co.: Hope (4); Izard Co.: Horseshoe Bend (1); Lawrence Co.: Imboden (1); Miller Co.: Fouke (1), Texarkana (2); Mississippi Co.: Neal (4); Ouachita Co.: Shumaker (1), Pulaski Co.: Little Rock (1); Washington Co.: Fayetteville (7), no data (2); No data: “Ark” (5). DELAWARE. New Castle Co.: Wilmington (1); Sussex Co.: Rehoboth (2). DISTRICT OF COLUMBIA : Rock Creek Park (22), Washington (24), Woodridge (1), no data (13). FLORIDA. Alachua Co. : Arredondo (2), Gainesville (50), no data (2); Broward Co.: Fort Lauderdale (3); Charlotte Co.: Murdock (2), Punta Gorda (6), Pt. Charlotte (6); Clay Co.: Green Cove (1); Collier Co.: Marco Island (7); Columbia Co. : Lake City (1); Dade Co.: Homestead (4); Duval Co.: Jacksonville (2); Dixie Co.: Oldtown (1); Jefferson Co.: Monticello (1); Gadsden Co.: Quincy (20), Gilchrist Co.: No data (1); Hardee Co.: Bowling Green (6), Zolfo Springs (1); Hendry Co.: LaBelle (1); Highlands Co.: Lake Placid (Archbold Biological Station) (38), Sebring (2), no data (1); Hillsborough Co.: Tampa (1); Indian River Co.: Vero Beach (2); Lake Co.: Tavares (1); Lee Co.: Buck Key (1), Fort Myers (27), Lehigh Acres (9), Olga (1), Sanibel Island (3); Liberty Co.: No data (1); Levy Co.: Chiefland (3); Manatee Co.: Bradenton (2), Mariposa Key (1); Miami- Dade Co.: Coconut Grove (3), Coral Gables (13), Cutler (4), Key Biscayne (1), Lemon City (1), Miami (186), Naranja (3), Silver Palm (4), no data (8); Monroe Co.: Big Coppitt Key (6), Big Pine Key (10), Key Largo (2), Key West (2), Islamorada (1), Stock Island (11); Nassau Co. Fernandina Beach (3); Orange Co.: Lake Buena Vista (2), Orlando (13); Osceola Co. : Osceola (1); Palm Beach Co.: Palm Beach (3), Jupiter (9); Pinellas Co.: Fort Desoto State Park (1); Polk Co.: Avon park (1), Babson Park (1), Lake Marion Estates (1), Lakeland (2); Pinellas Co.: Anona (1), Paradise Key (1); Polk Co.: Davenport (1); Putnam Co.: Crescent City (1), Ordway Preserve (2), Palatka (4); St. Johns Co.: St. Augustine (1); St. Lucie Co.: Fort Capron (3); Sarasota Co. : Englewood (5), Laurel (4), North Port (16), Sarasota (1); Sumter Co.: Tarrytown (1); Volusia Co. : Enterprise (2), Holly Hill (1), South Daytona (7); Walton Co.: De Funiak Springs (1); No data: Florida (12); GEORGIA. Berrien Co.: Nashville (1); Charlton Co.: Saint George (1); Chatham Co.: Savannah (4), Tybee Island (1); Cobb Co.: Kennesaw Mt. (3), no data (1); Coweta Co.: Madras (9); Crawford Co.: Gaillard (1); DeKalb Co.: Stone Mountain (8); Emanuel Co.: Swainsboro (2); Fulton Co.: Atlanta (3); Grady Co.: Beachton (1); Glynn Co.: Brunswick (1); Lamar Co.: Barnesville (2); McIntosh Co.: Sapelo Island (1); Murray Co.: Chatsworth (1); Muscogee Co.: Columbus (1); Pike Co.: Zebulon (1); Spalding Co.: Experiment (1), Griffin (1); Thomas Co.: Thomasville (1); Upson Co.: The Rock (3), Yatesville (1); White Co.: Unicoi State Park (1), no data (1). ILLINOIS. Clay Co.: No data (1); Champaign Co.: Champaign (2); Hamilton Co.: Broughton (2), Mcleansboro (5); Hardin Co.: Herod (1); Johnson Co.: Goreville (1); Jackson Co.: Carbondale (1); Kankakee Co.: Momence (1); Mason Co.: Sand Ridge State Forest (1), Topeka (1); Tazewell Co.: Tremont (1); Washington Co.: Du Bois (1), no data (15); Williamson Co.: Crab Orchard (1), no data: “South Illinois ” (1). INDIANA. Clark Co.: Borden (3); Clay Co.: Ashboro (5); Jasper Co.: No data (1); Knox Co.: No data (9); Lake Co.: Hessville (5); Laporte Co.: No data (1); Montgomery Co.: Crawfordsville (2); Parke Co.: No data (1); Pulaski Co.: No data (1); Starke Co.: North Judson (1); Spencer Co.: Hammond (1); Sullivan Co.: Sullivan (1); Tippecanoe Co.: Lafayette (1), no data (3); Vigo Co.: Terre Haute (1), no data: “Ind. Ter.” (1). IOWA. Fremont Co.: Hamburg (1). KANSAS. Barber Co.: No data (1); Butler Co.: Beaumont (1); Clark Co: Englewood (1); Douglas Co.: Lawrence (4), Lone Star Lake (1); Franklin Co.: No data (2); Kiowa Co. Belvidere (1); Labette Co.: Oswego (18); Neosho Co.: No data (1); Osage Co.: Carbondale (1); Pottawatomie Co.: No data (3); Reno Co.: Medora (4); Riley Co.: Manhattan (4), no data (26); Sedgwick Co.: Mount Hope (2), Wichita (1); Shawnee Co.: Topeka (2); Sumner Co.: Wellington (2); Waubaunsee Co.: Eskridge (4); Wilson Co.: Benedict (2). KENTUCKY. Boyd Co.: Ashland (3); Clark Co.: No data (1); Daviess Co.: Owensboro (2); Harlan Co.: Black Mountain (1), Hart Co.: Hardyville (16); Henderson Co.: Henderson (1); Lawrence Co.: Louisa (1); Louisville-Jefferson Co.: Louisville (1); Madison Co.: Berea (1); Rockcastle Co.: Livingston (2); Rowan Co.: Farmers (2); Todd Co.: Guthrie (1), no data: “K.Y.” (4). LOUISIANA. Acadia Pa. : Crowley (1); Avoyelles Pa.: Marksville (4); Caddo Pa. : Shreveport (12); East Baton Rouge Pa.: Baton Rouge (4); Evangeline Pa.: Chataignier (2); Iberville Pa.: No data (4); Landry Pa.: Sunset (1); Orleans Pa.: New Orleans (5); Madison Pa.: Mound (1), Tallulah (2), no data (1); Rapides Pa.: Alexandria (1); Saint Charles Pa.: Saint Rose (1); Saint Martin Pa.: Morgan City (1); Saint Mary Pa.: Berwick (1); Saint Tammany Pa.: Covington (4); Tangipahoa Pa.: Ponchatoula (1), no data (1); No data (1). MARYLAND. Anne Arundel Co.: Annapolis (1), Laurel (1), Odenton (1), no data (2); Baltimore Co.: Catonsville (6); Calvert Co.: Chesapeake Beach (4); Charles Co.: Bryantown (1), Marshall Hall (1), Nanjemoy (1); Kent Co.: Chestertown (2); Montgomery Co.: Cabin John (1), Germantown (1), Glen Echo (1), Great Falls (1), Plummers Island (2); Prince George Co.: Bladensburg (3), Beltsville (8), Bowie (1), Camp Springs (1), College Park (20), Hyattsville (3), Lakeland (9); Talbot Co.: Easton (1); Washington Co.: Hagerstown (1), Hancock (4); Wicomico Co.: Nutters Neck (2); Worcester Co.: Snow Hill (1); No data (13). MISSISSIPPI. Adams Co.: Natchez (3); Choctaw Co.: Tombigbee National Forest (1), Harrison Co.: Handsboro; Hinds Co.: Bolton (1), Jackson (3); George Co.: Lucedale (2); Harrison Co.: Handsboro (2); Jackson Co.: Ocean Springs (1); Jones Co. : Laurel (1); Lamar Co.: Purvis (1); Lee Co.: Saltillo (3); Marshall Co.: Holly Springs (4); Oktibbeha Co.: Starkville (1), Tippah Co: “At Alcorn Co. Line on 72” (1), no data: “Southern Mississippi ” (1). MISSOURI. Barry Co.: Roaring River State Park (1); Boone Co.: Columbia (6); Buchanan Co.: San Antonio (1); Cape Girardeau Co.: Cape Girardeau (7); Christian Co.: Keltner (1); Clair Co.: Osceola (1); Daviess Co.: Gallatin (1); Franklin Co.: Meramet State Park (1), New Haven (1); Greene Co.: Willard (1); Jackson Co.: No data (3); Jasper Co.: Joplin (1); Iron Co. : Graniteville (1); Mississippi Co.: Charleston (1); New Madrid Co.: New Madrid (4); Newton Co.: No data (1); Ozark Co.: Caney Mountain (2); Saint Louis Co.: Gray Summit (6), Kirkwood (1), Overland (1), Ranken (1), Saint Louis (14); Shannon Co.: Birch Tree (1); Stone Co.: Red Rock Creek (1); Texas Co.: Cabool (4), Mountain Grove (1); Wright Co.: Mountain Grove (3), no data: “ Missouri ” (2). NEBRASKA. Adams Co.: No data (1); Cass Co.: Platte River State Park (1); Hall Co.: Wood River (6 mi. S) (1); Jefferson Co.: No data (1); Keya Paha Co. : Jamison (1); Lancaster Co.: Lincoln (2), no data (7); Nemaha Co.: Indian Cave State Park (1); Pawnee Co.: Prairie Knoll (2); Richardson Co.: Camp Cornhusker (2), Indian Cave State Park (8), Rulo (1), no data (5). NEW JERSEY. Burlington Co.: Upper Mill (1); Camden Co.: Clementon (1); Cumberland Co.: Rutgers Experimental Station (2), Vineland (1); Gloucester Co.: Glassboro (1); Ocean Co.: Bamber Lake (1), Lakehurst (7), Ocean City (1), Warren Grove (1); Passaic Co.: Midvale (2) No data: “N.J.” (3). NEW YORK. New York Co.: New York (1). NORTH CAROLINA. Alleghany Co.: Ennice (1); Buncombe Co.: Asheville (8); Carteret Co.: Bogue Island (1); Dare Co.: Frisco (1), Nags Head (1); Franklin Co.: Louisburg (1); Gaston Co.: Crowders Mountain State Park (1); Harnett Co.: Dunn (1); Haywood Co.: Balsam (1), Canton (1), Mount Guyot (1), Retreat (1), Sunburst (3); Hereford Co.: No data (1); Jackson Co.: Balsam (20), Dillsboro (1); Johnston Co.: Clayton (1); Macon Co.: Highlands (2); Mecklenburg Co.: Charlotte (1), Mooresville (1); Moore Co.: Southern Pines (11); Randolph Co.: Asheboro (1); Rowan Co.: Salisbury (3); Tyrrell Co.: Columbia (1); Wake Co.: Raleigh (3); Wilson Co.: Wilson (1), no data: “Church IA” (1). OHIO. Adams Co.: Cedar Falls (2); Athens Co.: Athens (4); Butler Co.: Oxford (2); Carroll Co.: No data (1); Hamilton Co.: Cincinnati (1); Hocking Co.: No data (1); Meigs Co.: No data (1); Muskingum Co.: Roseville (1); No data: “ Ohio ” (1). OKLAHOMA. Blaine Co.: Bridgeport (1); Canadian Co.: El Reno (18); Carter Co.: Ardmore (7); Comanche Co.: Lawton (4), Fort Sill (1), Wichita Mountains (2), no data (2); Craig Co.: Centralia (1); Ellis Co.: Arnett (1), Shattuck (1); Garfield Co.: No data (3); Garvin Co.: Paoli (1); Grady Co.: Tuttle (3); Logan Co.: Guthrie (6), Mulhall (1); Marshall Co.: Willis (3); McIntosh Co.: Eufaula (1); Oklahoma Co.: Oklahoma City (5); Payne Co.: Coyle (1), Ripley (1), Stillwater (4), no data (7); Sequoyah Co.: Sallisaw (1); Texas Co.: Optima (1); Woodward Co.: Woodward (1), no data (1); No data: “Chateau” (1), “Mefrimec” (1). PENNSYLVANNIA. Allegheny Co.: Mount Lebanon (1); Bedford Co.: No data (5); Chester Co.: Downingtown (1); Clyde Co. : No data (1); Cumberland Co.: Newville (12); Delaware Co. Broomall (1); Fayette Co.: Uniontown (1); Franklin Co.: Chambersburg (1); Lancaster Co.: Lime Rock (1); Union Co.: Lewisburg (2); Westmoreland Co.: Jeannette (1); York Co.: York (1). SOUTH CAROLINA. Aiken Co.: Aiken (1), White Pond (1); Beaufort Co.: Bluffton (3), Charleston Co.: Charleston (17), Clarendon Co.: Gable (1), Pocotaligo (1), Saint Paul (2), no data (1); Colleton Co.: Walterboro (2); Florence Co.: Florence (5); Georgetown Co.: Georgetown (1), Sampit (1); Jasper Co.: Ridgeland (2); Lee Co.: Meredith (45), Rosehill (1); Oconee Co.: Seneca (1), Walhalla (314); Pickens Co.: Clemson (13), Six Mile (1); No data: “ South Carolina ” (3). TENNESSEE. Campbell Co.: Stinking Creek (1); Chester Co.: Henderson (4); Cumberland Co.: Black Mountain (1), Grassy Cove (1); Clarksville-Montgomery Co.: Clarksville (6); Davidson Co.: Nashville (8); Grainger Co.: Elm Springs (3); Haywood Co.: Brownsville (49); Hardeman Co.: Bolivar (1); Knox Co.: Fountain City (1); Lawrence Co.: David Crockett State Park (1); Madison Co.: Jackson (8); Morgan Co.: Burrville (5), Deer Lodge (2); Putnam Co: Baxter (6); Roane Co.: Harriman (17); Rutherford Co.: Murfreesboro (7); Shelby Co.: Memphis (1), no data (2); Trousdale Co.: Hartsville (1); Warren Co.: McMinnville (2); Weakley Co.: Dresden (1); Williamson Co.: Spring Hill (1); No data: “ Tennessee ” (2). TEXAS. Archer Co.: No data (1); Anderson Co.: Tennessee Colony (5); Atascosa Co. : Campbellton (65); Bastrop Co.: “Stengall Ranch” (1); Bexar Co.: San Antonio (22), Somerset (13), Stone Oak (2); Brazoria Co. : Alvin (3); Brazos Co.: Bryan Air Force Base (9), College Station (6), no data (1); Bowie Co.: Texarkana (1); Burleson Co.: Caldwell (1); Cameron Co.: Brownsville (23), Port Isabel (1); Carson Co.: Panhandle (1); Brooks Co.: Falfurrias (1); Cass Co.: Atlanta (1); Camp Co.: Pittsburg (2); Kleburg Co.: Ricardo (1); Colorado Co.: Columbus (2), Rock Island (20), no data (46); Comal Co.: Bulverde (2), New Braunfels (7), no data (3); Cooke Co.: Gainesville (3) Lake Ray Roberts (2); Dallas Co.: Dallas (6); DeWitt Co.: Cuero (2), Nordheim (1); Dimmit Co.: Catarina (4), no data (1); Duval Co.: Benavides (1), Freer (3), San Diego (6); Fannin Co.: Ladonia (2); Frio Co.: Dilley (1); Goliad Co.: No data (1); Harris Co.: Hockley (2); Hidalgo Co.: Hidalgo (1), Mercedes (1), no data (7); Jack Co.: Antelope (1); Jackson Co.: Ganado (16); Jasper Co.: Jasper (1); Jefferson Co.: Beaumont (2); Jim Wells Co.: Alice (3); Karnes Co. : Ecleto (11), Runge (7); Kleberg Co.: Kingsville (8), Ricardo (2); La Salle Co.: Cotulla (5); Leon Co.: Oakwood (2); McMullen Co.: Tilden (1); Medina Co. : Castroville (1), Hondo (2); Montague Co.: Forestburg (26); Nacogdoches Co.: Nacogdoches (5); Nueces Co.: Corpus Christi (5), Robstown (2); Rusk Co.: Henderson (1); San Patricio Co.: Lake Corpus Christi (1), Mathis (10), Sinton (3), Welder Wildlife Foundation (2); Shelby Co.: Center (4); Starr Co.: El Sauz (8), Grande City (1), no data (8); Tarrant Co.: Arlington (1), Handley (6); Taylor Co.: Taylor (1); Travis Co.: Austin (13), no data (1); Victoria Co. : Victoria (14); Uvalde Co.: Concan (1), Uvalde (2); Val Verde Co.: Del Rio (2); Webb Co.: Aguilares (2), Laredo (26), no data (1); Wheeler Co.: Shamrock (1); Wichita Co.: Burkburnett (1); Zavala Co.: Crystal City (1); No data: “Shoal Creek” (1), “ Texas ” (64). VIRGINIA. Albemarle Co.: Charlottesville (1), Spring Hill (1); Alex Co. : Barcroft (1), no data (2); Alexandria Co. : Hunting Creek (1); Arlington Co.: Alexandria (1), Arlington (6), Four Mile Run (1), Glencarlyn (2); Chesterfield Co.: Pocahontas State Park (1); Bath Co.: Hot Springs (1); Essex Co.: Tappahannock (2); Fairfax Co.: Ash Grove (1), Black Pond (1), Jefferson (1), Fairfax (1), Great Falls (1), Holmes Run Park (1), Hunter (1), Mount Vernon (1),Vienna (1), no data (1); Fauquier Co.: No data (3); James City Co.: Springhill (2); Loudoun Co.: Leesburg (10), Middleburg (1); Madison Co.: Madison (1); Montgomery Co.: No data (3); Nansemond Co.: Cypress Chapel (1); Nelson Co.: Afton (1), no data (15); Norfolk Co.: Norfolk (10); Prince William Co.: Bristow (3), Occoquan (1); Virginia Co.: Falls Church (22); No data: “ Virginia ” (2). WISCONSIN. Green Co.: Albany (1). WEST VIRGINIA. Greenbrier Co.: White Sulphur Springs (7); Kanawha Co.: Kanawha (2); Lewis Co.: Jackson’ s Mill (1); Logan Co.: Peach Creek (4), Monroe Co.: Slaty Mountain Preserve (1); Racine Co.: Racine (3). NO DATA: “Cherokee” (1), no data (14).

Figures

Fig. 13. Euphoria sepulcralis. a) Habitus, b) Head, c) Parameres, d-e) Color variation, f) Temporal distribution, g) Geographic distribution in the Bahamas, Cayman Islands, Mexico, and the United States.