Euphoria Burmeister, 1842

Orozco, Jesús, 2012, Monographic Revision of the American Genus Euphoria Burmeister, 1842 (Coleoptera: Scarabaeidae: Cetoniinae), The Coleopterists Bulletin (mo 11) 66, pp. 1-182 : 10-16

Euphoria Burmeister, 1842

Euphoria Burmeister 1842: 370 .

Type species: Cetonia sepulcralis F., designated by Casey (1915).

Erirhipi s Burmeister 1842: 385 . Synonym.

Type species: Cetonia fulgida F., designated by Casey (1915).

Stephanucha Burmeister 1842: 385 . Synonym.

Type species: Cetonia areata F., by monotypy.

Goraqua Péringuey 1907: 358 . Synonym.

Type species: Goraqua smithsana Péringuey , by monotypy.

Anatropis Casey 1915: 298 . Synonym.

Type species: Euphoria verticalis Horn , by monotypy.

Euphoriaspis Casey 1915: 298 . Synonym.

Type species: Euphoria hirtipes Horn , by monotypy.

Euphoriopsis Casey 1915: 298 .

Type species: Euphoria hera Burmeister , by monotypy.

Erirhipidia Casey 1915: 308 . Synonym.

Type species: Scarabaeus indus L., by original designation.

Haplophoria Casey 1915: 310 . Synonym.

Type species: Euphoria kernii Haldeman , by original designation.

Euphorhipis Casey 1915: 314 . Synonym.

Type species: Cetonia subtomentosa Gory and Percheron , by original designation.

Rhipiphoria Casey 1915: 316 . Synonym.

Type species: Cetonia geminata Chevrolat , by original designation.

Isorhipina Casey 1915: 326 . Synonym.

Type species: Cetonia biguttata Gory and Percheron , by original designation.

Description. Length 7.6–21.5 mm; width 4.5–12.2 mm. Color: Dorsal surface highly variable, shiny or dull, frequently tomentous. Pronotum unicolored to medially or laterally vittate; vittae black, dark green, or dark brown; sides occasionally with cretaceous band. Elytra frequently with pattern of color or cretaceous markings. Pygidium frequently with cretaceous markings or entirely cretaceous. Melanistic forms observed. Head: Surface unarmed. Frons medially depressed or flat, occasionally with weak, median, longitudinal ridge, rarely with central protuberance, moderately to densely punctate, rugopunctate, or rugose; punctures round, deep to moderately impressed, small to moderate in size, occasionally confluent, glabrous to densely setose. Clypeus subquadrate, subrectangular, or acuminate, sides subparallel to strongly anteriorly convergent, flat to strongly raised, lateral declivity weakly to strongly expanded; apex vaguely to strongly reflexed, occasionally with 2–4 small denticles, truncate to moderately sinuate in dorsal view; surface moderately to densely punctate; punctures round, deep to moderately impressed, small to moderate in size, occasionally confluent, glabrous to densely setose. Antenna with 10 antennomeres, antennal club frequently sexually dimorphic in length (as long as or longer than stem in males, shorter in females). Pronotum: Surface sparsely to densely punctate; punctures round to lunulate, minute to moderate, frequently denser and larger toward sides, sparsely to densely setose. Widest at base, sides strongly angulate to evenly rounded, subparallel to strongly convergent anteriorly. Base in front of scutellum evenly rounded to strongly emarginate. Scutellum longer than wide, sides straight, impunctate to densely punctate; punctures minute to moderate, glabrous to densely setose. Elytra: Sides subparallel, apex truncate. Surface sparsely to densely punctate, punctures small to moderate, 2 discal costae weakly to strongly raised. Subhumeral emargination moderately to strongly developed. Pygidium: Surface concentrically to subconcentrically striate, occasionally polished at middle, sparsely to densely setose. Legs: Surface moderately densely to densely setose, setae moderate to long. Protibiae tridentate, teeth equidistant or apical and medial teeth closer to each other than to basal tooth; teeth oblique or transverse to tibia, basal tooth frequently obsolete to subobsolete. Meso- and metatibiae with variably developed carinae. Mesotarsal length frequently sexually dimorphic (as long as or longer than mesotibia in males, shorter in females). Metafemora with long suture on internal angle. Metatibiae apically expanded at times, with 2 apical spurs; coxae flattened in posterolateral area. Venter: Prosternal process laterally compressed, short. Mesometasternal process weakly to strongly compressed laterally, variably extended anteriorly, apex variably rounded. Mesepimera, metasternum, and metacoxae setose. Metasternum flattened at middle, rugose, moderately densely to densely setose laterally, weakly to moderately densely punctate and setose at middle, median sulcus variably impressed. Sides of abdominal sternites rounded to ridged. Male genitalia: Aedeagus without spines or hooks, parameres fused at times. Some diagnostic for species.

Diagnosis. Euphoria is separated from the other American Cetoniini , Chlorixanthe , by the short prosternal process, elevated elytral costae, rounded scutellar apex, and body not strongly deplanate. Separating Euphoria from some Old World genera is difficult using a single character but can be done by the following combination of characters in Euphoria : unarmed head; presence of 2 discal costae on elytra; straight sides of scutellum; flattened metasternum; long posterior striae on metafemora; metacoxae flattened in posterolateral area; pygidium concentrically to subconcentrically striate; and parameres without hooks or spines.

Composition. Euphoria is comprised of 59 species, ten of which are described here as new. More than half of the 134 names in the group are synonyms (Appendix 1). Based on morphological characters of adults, all Euphoria species can, for the most part, be assigned to one of 14 species-groups ( Table 5 ).

Species-Groups. The amount of interspecific variation in the clypeal apex, clypeal sides, shape and length of mesometasternal process, legs, genitalia, body shape, and body vestiture seen in Euphoria is only found in other groups or other geographic regions when several genera are studied. Therefore, a further breakdown of this genus appears inevitable. Based on my morphological analysis, I have enough information to propose a new classification that would result in the creation of several genera. However, given the difficulties in ascertaining a classification based on phylogenetically meaningful characters ( Orozco and Philips 2010), I am reluctant to propose new genera. Therefore, I resort to species-groups instead. Additional evidence on the phylogenetic relationships of these groups might provide the support necessary to consider them genera.

The division of this heterogeneous genus into informal species-groups is based on the character examination of almost 20,000 specimens including most of the types. The diagnoses of the speciesgroups are products of the aforementioned character analysis and are based on hypothesized synapomorphies. All groups are supported by a guild of characters rather than unique characters. Characters from the head (i. e., shape of the clypeus and presence or absence of sexual dimorphism in the antennal lamellae) frequently used to support genera in scarab beetles are here used to support some of the species-groups. Other characters used to support genera in different groups of scarabs are also used here to support species-groups: pronotal shape, shape of the mesometasternal process, tarsal length, and general shape of the parameres ( Table 6 and each species-group diagnosis). The groups I consider basal, based on the characters they exhibit, are presented first. Table 5 shows the species composition of each group.

Natural History. Life history for most of the species is unknown. What information is known is listed under each species treatment. Unless indicated by a reference citation, the information presented was recorded from label data.

In general, the life cycle is estimated to be one year, but it is known for only a few species. In temperate climates, the overwintering stage is the adult ( Blatchley 1910; Hayes 1925; Ritcher 1945; Ritcher 1966; Lago et al. 1979; Ratcliffe 1991; Ratcliffe and Paulsen 2008). The larvae are known to occur in highly organic soil, sandy soil, ant mounds, ant debris piles, rodent nests, and under dry dung. Most larvae feed directly on the substrate where they live, but some have been recorded feeding on the roots of grasses (e. g., Euphoria sepulcralis (F.); Buss 2004). The third instar is known for ten species: Euphoria abreona Janson (described as Euphoria precaria Janson ) ( Orozco and Pardo-Locarno 2004), Euphoria areata (F.) (described as Stephanucha thoracica Casey ) ( Skelley 1991), Euphoria basalis (Gory and Percheron) ( Ramírez-Salinas et al. 2001) , Euphoria devulsa Horn ( Micó et al. 2000) , Euphoria fulgida (F.) ( Hayes 1925, 1929; Ritcher 1945, 1966), Euphoria herbacea (Olivier) ( Hayes 1925, 1929; Ritcher 1945, 1966), Euphoria hirtipes Horn (described as Euphoriaspis hirtipes (Horn)) ( Ratcliffe 1976) , Euphoria inda (L.) ( Hayes 1925, 1929; Ritcher 1945, 1966), Euphoria lurida (F.) ( Micó et al. 2000), and E. sepulcralis ( Hayes 1925, 1929; Ritcher 1945, 1966). The pupae have only been described for E. abreona ( Orozco and Pardo-Locarno 2004) and E. hirtipes ( Ratcliffe 1976) .

Adults have coriaceous mandibles and are known to feed on liquids from plant exudates, soft parts from plants, fruits, and dung. Pollen-feeding is suspected, but there is still no direct evidence of it. Some species have been reported to cause damage to flowers and crops ( Halbert 1996; Cunha et al. 2007), but this is mostly due to mechanical damage caused by the grasping legs of numerous individuals when feeding en masse.

Some species are frequently found associated as adults and larvae with mounds or debris piles of ants of the genera Acromyrmex Mayr , Atta F., Formica L., and Pogonomyrmex Mayr ( Hymenoptera : Formicidae ): E. areata , Euphoria biguttata (Gory and Percheron) , Euphoria canescens (Gory and Percheron) , Euphoria dimidiata (Gory and Percheron) , Euphoria discicollis (Thomson) , E. hirtipes , E. inda , Euphoria leucographa (Gory and Percheron) , Euphoria levinotata Orozco new species, Euphoria pilipennis (Kraatz) , Euphoria pulchella (Gory and Percheron) , and Euphoria subtomentosa (Gory and Percheron) ( Wheeler 1910; Hinton and Ancona 1935; Windsor 1964; Ratcliffe 1976; Deloya 1988; Rojas 1989; Deloya and Morón 1994; Deloya et al. 1995; Navarrete-Heredia 2001; Paulsen 2002). Adults of some species are also frequently found associated with nests of rodents of the genera Cynomys Rafinesque , Geomys Rafinesque , and Neotoma Say and Ord ( Rodentia : Sciuridae , Geomyidae , and Cricetidae , respectively): E. areata , E. devulsa , E. discicollis , Euphoria fascifera (LeConte) , E. histrionica Thomson , E. levinotata , and Euphoria verticalis Horn. These associations appear to be opportunistic, since the species are also known to occur in other habitats.

Geographic Distribution. Euphoria species are distributed from southern Canada to northern Argentina . At both extremes the diversity is low, having only two and one species, respectively (Appendix 2). Species are known from every continental country in the Americas except Chile and Suriname. The only species known from the Antilles ( E. sepulcralis from Eleuthera and New Providence Island in the Bahamas) was probably recently introduced to the islands by shipments of fruits or flowers as it has been the case with other cetoniines (i. e., Protaetia fusca (Herbst)) . The highest species diversity is found between the southern United States and Guatemala, decreasing south of the Nicaraguan depression and north of the states of Arizona and Texas in the United States. This distribution resembles the Paleo- American pattern described by Halffter (1987). The Paleo-American pattern includes organisms that had an early immigration to the continent from lineages that originated in the Old World ( Halffter 1987). As implied by this pattern, species of Cetoniini could have diversified and flourished in the Mexican mountain ranges north of the Isthmus of Tehuantepec and slowly extended their ranges and speciated both to the north and south.

Phylogenetic Relationships. Previous phylogenetic analyses of the tribe ( Orozco and Philips 2010) have illustrated the difficulties in studying this genus morphologically. Molecular characters are being used in the phylogeny of the Cetoniinae (JO, unpublished data; D. Hawks and M. Paulsen, personal communication). Preliminary results from these analyses support the monophyly of Euphoria when the species formerly considered in Stephanucha are included, but the analyses are far from definitive in the study of the relationships among the species. Larval characters have not been exhaustively explored, but appear to be only of diagnostic use. The larvae of only 10 of 59 of the species are known to date.

Chlorixanthe was hypothesized to be the sistergroup of Euphoria by Orozco and Philips (2010). The flattened body, the unusually long and rhomboidal mesometasternal process, and the unusually compressed tarsi set Chlorixanthe species apart from all other Cetoniini . Two species, C. flavoviridis and C. propinqua , are currently known in the genus.

Previous Groupings. Burmeister (1842) and Casey (1915) provided the only two classifications supported by morphologic characters. The speciesgroups of Bates (1889) and Hardy (2001) are taxonomically untestable because the basis for them is unknown.

Regarding the American Cetoniini , Burmeister’ s (1842) classification is plagued with imprecisions. Due to the fact that he had access to few (in some cases, apparently just one) specimens per species and at times only one sex, his classification and character discussion are of limited use. For example, at the generic level, he separated Euphoria from Erirhipis and Stephanucha by it having the antennal club equally long in both sexes and the clypeus anteriorly reduced. Nevertheless, of 12 species he included in Euphoria , only four actually have this character: E. lurida , E. basalis , E. canescens , and E. biguttata .

Casey (1915), while successful in showing the diversity in the group, failed in providing a predictable classification. Most, if not all, of Casey’ s genera or subgenera are so generally defined and filled with exceptions that almost any given species could be included in any of them without much trouble. His classification is also, like that of Burmeister (1842), plagued with imprecisions (for examples, see taxonomic history of Euphoria hera Burmeister and E. hirtipes ). At the species level, Casey was able to observe numerous minor differences in the characters. Unfortunately, for a group with such cornucopian intraspecific variation, this was not a good attribute. All of the 17 species and eight subspecies described by Casey (1915) are synonyms. Figure 1 shows the total number of species described by year when synonyms are included and excluded.

KEY TO THE SPECIES- GROUPS OF EUPHORIA

1. Clypeal apex strongly to moderately sinuate in dorsal view (e. g., Figs. 41b , 48b ). Base of pronotum in front of scutellum weakly to moderately emarginate (e. g., Figs. 44a , 47a ) ................. geminata species-group (p. 72)

1′. Clypeal apex truncate or weakly sinuate in dorsal view. Base of pronotum variable................................................................2

2(1′). Pronotal base evenly rounded in front of scutellum (e. g., Figs. 58a , 60a ). Mesometasternal process strongly compressed and setose. Clypeus frequently quadridentate or bidentate (e. g., Figs. 56b , 59b )..............................................................3

2′. Pronotal base sinuate to variably emarginate in front of scutellum. Mesometasternal process variable. Clypeus without denticles ..................................................................... 5

3(2). Clypeal apex without denticles ( Fig. 58b ) ........... discicollis species-group (p. 102)

3′. Clypeal apex with 2–4 denticles...........4

4(3′). Apex of clypeus with 2 denticles ( Figs. 59b , 60b ) ........ verticalis species-group (p. 103)

4′. Apex of clypeus with 4 denticles ( Figs. 55b , 56b , 57b ) ...... areata species-group (p. 98)

5(2′). Apex of clypeus strongly emarginate ( Fig. 25b )...... hera species-group (p. 57)

5′. Apex of clypeus not strongly emarginate..............................................................6

6(5′). Clypeus quadrate to trapezoidal with apex and sides not raised (slightly raised at most) (e. g., Figs. 5b , 9b ). Frons flattened and strigose. Dorsum variably covered by cretaceous markings..................................... ............ histrionica species-group (p. 21)

6′. Clypeus variable in form, never quadrate or trapezoidal; apex and sides variable. Frons never flattened and strigose. Dorsum with or without cretaceous markings............................................7

7(6′). Elytral striae densely setose, composed almost exclusively of long grooves (e. g., Figs. 49a , 52a ). Clypeal apex and sides not raised. Pygidium surface strongly concentric...... pulchella species-group (p. 85)

7′. Elytral striae never densely setose nor composed almost exclusively of long grooves. Clypeal apex variable. Pygidium variable...................................................8

8(7′). Pronotum densely setose, with regular, glabrous patches on surface ( Figs. 53a , 54a ). Parameres as in Figs. 53c and 54c .... ......................... inda species-group (p. 92)

8′. Pronotum variably setose, if densely setose then never with regular glabrous patches on surface. Parameres not as in Figs. 53c and 54c ................................... 9

9(8′). Clypeus moderately to strongly attenuate, sides and apex not raised (e. g., Figs. 2b , 3b ). Humeral area frequently with red markings (e. g., Figs. 3a , 4a ). Cretaceous markings on elytra small to large when present.... biguttata species-group (p. 16)

9′. Clypeus never attenuate, sides frequently raised. Humeral area without red markings. Cretaceous markings on elytra small when present.........................................10

10(9′). Pronotum with vittae (except Euphoria submaculosa (Gory and Percheron)) (e. g., Figs. 30a , 32a ). Dorsum generally yellowish brown with black or brown markings, rarely green with yellowish markings. Without cretaceous spots on dorsum......... ....................... avita species-group (p. 58)

10′. Pronotum without vittae. Dorsum never yellowish brown, or with brown or black markings. Frequently with cretaceous spots on dorsum ................................................ 11

11(10′). Clypeal apex strongly reflexed in both sexes, sinuate in frontal view (e. g., Figs. 22b , 24b ). Female metatarsi strongly compressed. Male parameres widely expanded apically as in Figs. 22c , 23c , 24c ........ candezei species-group (p. 52)

11′. Clypeal apex reflexed only in males. Female metatarsi not strongly compressed. Male parameres not widely expanded apically ............................................... 12

12(11′). Dorsum tomentous (e. g., Figs. 18a , 20a ) ............ herbacea species-group (p. 47)

12′. Dorsum shiny (e. g., Figs. 11a , 16a )....13

13(12′). Black, brown, or cupreous species (e. g., Figs. 13a , 14a ). Elytra with abundant, vermiform or reniform, cretaceous markings. Pronotum frequently with cretaceous line on lateral margin ............................. .......... sepulcralis species-group (p. 28)

13′. Greenish or violaceous species (e. g., Figs.16a , 17a ). Cretaceous markings on elytra irregular, sparse, minute at times. Pronotum without cretaceous line on pronotal sides...... ................ fulgida species-group (p. 41)

CLAVE PARA LOS GRUPOS DE ESPECIES DE EUPHORIA

1. Ápice clipeal moderada o fuertemente sinuado en vista dorsal (e. g., Figs. 41b , 48b ). Base del pronoto en frente del escutelo débil a moderadamente emarginada (e. g., Figs. 44a , 47a ).... grupo geminata (p. 72)

1′. Ápice clipeal truncado o ligeramente sinuado en vista dorsal. Base del pronoto variable...................................................2

2(1′). Base del pronoto redondeada en frente del escutelo (e. g., Figs. 58a , 60a ). Proceso mesometasternal fuertemente comprimido y setoso. Clípeo frecuentemente cuadridentado o bidentado (e. g., Figs. 56b , 59b ) .................................................................. 3

2′. Base del pronoto en frente del escutelo variablemente emarginada. Proceso mesometasternal variable. Clípeo sin dentículos ........................................... 5

3(2). Ápice clipeal sin dentículos ( Fig. 58b ).... ........................ grupo discicollis (p. 102)

3′. Ápice clipeal con 2–4 dentículos..........4

4(3′). Ápice clipeal con 2 dentículos ( Figs. 59b , 60b ).................. grupo verticalis (p. 103)

4′. Ápice clipeal con 4 dentículos ( Figs. 55b , 56b , 57b ) ............... grupo areata (p. 98)

5(2′). Ápice clipeal fuertemente emarginado ( Fig. 25b ).................. grupo hera (p. 57)

5′. Ápice clipeal no fuertemente emarginado..............................................................6

6(5′). Clípeo cuadrado o trapezoidal, ápice y lados no fuertemente elevados (ligeramente elevados cuando mucho) (e. g., Figs. 5b , 9b ). Frente aplanada y estrigosa. Dorso variablemente cubierto por máculas cretáceas ....... grupo histrionica (p. 21)

6′. Clípeo variable en forma, nunca cuadrado o trapezoidal, ápice y lados variables. Frente nunca aplanada y estrigosa. Dorso con o sin máculas cretáceas...................7

7(6′). Estrías elitrales densamente setosas, compuestas en su mayoría por surcos largos (e. g., Figs. 49a , 52a ). Ápice y lados del clípeo nunca elevados. Superficie pigidial fuertemente concéntrica ............ ......................... grupo pulchella (p. 85)

7′. Estrías elitrales nunca densamente setosas o compuestas en su mayoría por surcos largos. Ápice clipeal variable. Pigídio variable ..................................................... 8

8(7′). Pronoto densamente setoso, con parches glabros e irregulares en su superficie ( Figs. 53a , 54a ). Parámeros como en las Figs. 53c y 54c .......... grupo inda (p. 92)

8′. Pronoto variablemente setoso, nunca con parches glabros e irregulares en su superficie. Parámeros no como en las Figs. 53c y 54c ................................... 9

9(8′). Clípeo moderada a fuertemente atenuado, ápice y lados no elevados (e. g., Figs. 2b , 3b ). Área humeral frecuentemente con máculas rojas (e. g., Figs. 3a , 4a ). Máculas cretáceas elitrales pequeñas o grandes si presentes ............ grupo biguttata (p. 16)

9′. Clípeo nunca atenuado, lados frecuentemente elevados. Área humeral sin máculas rojas. Máculas cretáceas elitrales pequeñas si presentes ................................................. 10

10(9′). Pronoto con vittae (excepto Euphoria submaculosa (Gory y Percheron)) (e. g., Figs. 30a , 32a ). Dorso generalmente caféamarillento con máculas negras o cafés, raramente verde con máculas amarillentas. Dorso sin máculas cretáceas....................... .................................... grupo avita (p. 58)

10′. Pronoto sin vittae. Dorso nunca caféamarillento o con máculas negras o cafés. Dorso frecuentemente con máculas cretáceas.................................................. 11

11(10′). Ápice clipeal fuertemente elevado en ambos sexos, sinuado en vista frontal (e. g., Figs. 22b , 24b ). Metatarsos de las hembras fuertemente comprimidos. Parámeros ampliamente expandidos apicalmente como en las Figs. 22c , 23c , 24c ........................ grupo candezei (p. 52)

11′. Ápice clipeal elevado solo en machos. Metatarsos de las hembras no fuertemente comprimidos. Parámeros no fuertemente expandidos apicalmente.......................12

12(11′). Dorso tomentoso (e. g., Figs. 18a , 20a ) ... ........................... grupo herbacea (p. 47)

12′. Dorso brillante (e. g., Figs. 11a , 16a )...13

13(12′). Especies negras, cafés o cobrizas (e. g., Figs. 13a , 14a ). Élitros con abundantes máculas cretáceas vermiformes o reniformes. Pronoto frecuentemente con línea cretácea sobre el margen lateral ............................... .......................... grupo sepulcralis (p. 28)

13′. Especies verdosas o violáceas (e. g., Figs. 16a , 17a ). Máculas cretáceas elitrales irregulares, escasas, en ocasiones diminutas. Pronoto sin línea cretácea sobre el margen lateral...... grupo fulgida (p. 41)

Figures

Fig. 1. Number of Euphoria species described by year when synonyms are included and excluded.

Fig. 41. Euphoria geminata. a) Habitus, b) Head, c) Parameres, d-g) Color variation, h) Temporal distribution, i) Geographic distribution in El Salvador, Guatemala, Honduras, Mexico, and Nicaragua.

Fig. 48. Euphoria vittata. a) Habitus of paratype, b) Head of paratype, c) Parameres of paratype, d) Temporal distribution, e) Geographic distribution in Mexico.

Fig. 44. Euphoria kernii. a) Habitus, b) Head, c) Parameres, d-e) Color variation, f) Temporal distribution, g) Geographic distribution in Mexico and the United States.

Fig. 47. Euphoria quadricollis. a) Habitus of male, b) Head, c) Parameres, d) Female, e) Temporal distribution, f) Geographic distribution in Mexico and the United States.

Fig. 58. Euphoria discicollis. a) Habitus, b) Head, c) Parameres, d, e) Color variation, f) Temporal distribution, g) Geographic distribution in the United States.

Fig. 60. Euphoria verticalis. a) Habitus, b) Head, c) Parameres, d) Temporal distribution, e) Geographic distribution in Mexico and the United States.

Fig. 56. Euphoria areata. a) Habitus, b) Head, c) Parameres, d-e) Color variation, f) Temporal distribution, g) Geographic distribution in the United States.

Fig. 59. Euphoria bispinis. a) Habitus, b) Head, c) Parameres, d) Temporal distribution, e) Geographic distribution in Mexico.

Fig. 55. Euphoria anneae. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Temporal distribution, f) Geographic distribution in the United States.

Fig. 57. Euphoria pilipennis. a) Habitus, b) Head, c) Parameres, d) Temporal distribution, e) Geographic distribution in the United States.

Fig. 25. Euphoria hera. a) Habitus, b) Head, c) Parameres, d-f) Color variation, g) Temporal distribution, h) Geographic distribution in Colombia and Venezuela.

Fig. 5. Euphoria devulsa. a) Habitus, b) Head, c) Parameres, d) Temporal distribution, e) Geographic distribution in the United States.

Fig. 9. Euphoria subtomentosa. a) Habitus, b) Head, c) Parameres, d) Temporal distribution, e) Geographic distribution in Mexico.

Fig. 49. Euphoria basalis. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Temporal distribution, f) Geographic distribution in Mexico and the United States.

Fig. 52. Euphoria pulchella. a) Habitus, b) Head, c) Parameres, d-e) Color variation, f) Temporal distribution, g) Geographic distribution in Belize, El Salvador, Guatemala, Honduras, and Mexico; open square indicates state record only.

Fig. 53. Euphoria hirtipes. a) Habitus, b) Head, c) Parameres, d) Temporal distribution, e) Geographic distribution in the United States.

Fig. 54. Euphoria inda. a) Habitus, b) Head, c) Parameres, d-e) Color variation, f) Temporal distribution, g) Geographic distribution in Canada, Mexico, and the United States; open square indicates state or province record only.

Fig. 2. Euphoria biguttata. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Temporal distribution, f) Geographic distribution in El Salvador, Guatemala, Honduras, and Mexico.

Fig. 3. Euphoria canescens. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Temporal distribution, f) Geographic distribution in Belize, El Salvador, Guatemala, Honduras, Mexico, and the United States; open square indicates country record only.

Fig. 4. Euphoria dimidiata. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Temporal distribution, f) Geographic distribution in El Salvador, Guatemala, Honduras, and Mexico; open square indicates state record only.

Fig. 30. Euphoria fulveola. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Temporal distribution, f) Geographic distribution in Mexico.

Fig. 32. Euphoria lacandona. a) Habitus of holotype, b) Head of holotype, c) Parameres of holotype, d) Color variation, e) Temporal distribution, f) Geographic distribution in Mexico.

Fig. 22. Euphoria candezei. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Temporal distribution, f) Geographic distribution in Costa Rica, Guatemala, Honduras, Mexico, Nicaragua, and Panama; open square indicates state record only.

Fig. 24. Euphoria westermanni. a) Habitus, b) Head, c) Parameres, d) Color variation, e-f) Females, g) Temporal distribution, h) Geographic distribution in Mexico.

Fig. 23. Euphoria lesueuri. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Female, f) Temporal distribution, g) Geographic distribution in Belize, El Salvador, Guatemala, Honduras, Mexico, and Nicaragua; open square indicates country record only.

Fig. 18. Euphoria herbacea. a) Habitus, b) Head, c) Parameres, d-e) Color variation, f) Temporal distribution, g) Geographic distribution in the United States.

Fig. 20. Euphoria steinheili. a) Habitus, b) Head, c) Parameres, d-f) Color variation, g) Temporal distribution, h) Geographic distribution in Colombia, Ecuador, Panama, Peru, and Venezuela; open square indicates country record only.

Fig. 11. Euphoria leucographa. a) Habitus, b) Head, c) Parameres, d-e) Color variation, f) Temporal distribution, g) Geographic distribution in Guatemala, Mexico, and the United States.

Fig. 16. Euphoria limbalis. a) Habitus, b) Head, c) Parameres, d-e) Color variation, f) Temporal distribution, g) Geographic distribution in the United States.

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.

Fig. 14. Euphoria yucateca. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Temporal distribution, f) Geographic distribution in Belize, Colombia, Costa Rica, El Salvador, Honduras, Guatemala, Nicaragua, Mexico, Panama, and Venezuela.

Fig. 17. Euphoria monticola. a) Habitus, b) Head, c) Parameres, d) Color variation, e) Temporal distribution, f) Geographic distribution in Mexico and the United States.

Tables

Table 6. Diagnostic characters for proposed species-groups of Euphoria. abs = absent; prs = present; tmts = tomentous.

						Antennal Pronotal		Emargination
		Cretaceous Clypeus Clypeus			Frons	club longer sides		of the base
	Dorsal surface	markings reflexed	convergent Longitudinal flattened in males than sexually					of the
Species-group	vestiture	on dorsum in males Clypeal apex anteriorly ridge on frons and strigose in females dimorphic						pronotum
biguttata group	shiny	prs; abs absent	entire present	absent	absent	absent	absent	weak; strong
histrionica group	shiny	prs; abs absent	entire absent	absent	present absent		absent	weak; strong
sepulcralis group	shiny	present prs; abs	entire absent	absent	absent	prs; abs	absent	weak; strong
fulgida group	shiny	prs; abs prs; abs	entire absent	absent	absent	present	absent	weak; strong
herbacea group	tomentous	present present	entire absent	absent	absent	present	prs; abs	weak; strong
candezei group	tmts; shiny	prs; abs absent	entire absent	prs; abs absent		present	prs; abs	strong
hera group	tomentous	absent absent	emarginate absent	present	absent	present	prs; abs	strong
avita group	tmts; shiny	absent present	entire absent	absent	absent	present	prs; abs	weak; strong
geminata group	tmts; shiny	prs; abs prs; abs	sinuate absent	absent	absent	present	prs; abs	absent; weak
pulchella group	tmts; shiny	prs; abs absent	entire present	absent	absent	prs; abs	absent	weak; strong
inda group	tmts; shiny	absent absent	entire absent	absent	absent	prs; abs	absent	weak; strong
areata group	shiny	prs; abs absent	quadridentate absent	absent	absent	present	absent	absent
discicollis group	shiny	absent absent	entire absent	absent	absent	present	absent	absent
verticalis group	shiny	absent absent	bidentate absent	absent	absent	absent	absent	absent
					Metatarsi Tarsal claws as
	Elytral striae Mesometasternal Protibiae Protibial teeth Mesotarsal				strongly long or longer Abdomen Sides of abdomen
	composed mostly process small		sexually short and length sexually compressed than last strongly convex					weakly to
Species-group	of deep grooves and compressed dimorphic perpendicular dimorphic				in females tarsomere in females strongly ridged
biguttata group	absent	absent	absent absent	absent	absent	absent	absent	absent
histrionica group	absent	absent	absent absent	absent	absent	absent	absent	absent
sepulcralis group	absent	absent	absent absent	absent	absent	absent	absent	absent
fulgida group	absent	absent	absent absent	absent	absent	absent	prs; abs	present
herbacea group	absent	absent	absent absent	prs; abs	absent	absent	present	prs; abs
candezei group	absent	absent	absent absent	present	present	absent	present	absent
hera group	absent	absent	absent absent	absent	absent	absent	present	absent
avita group	absent	absent	present absent	present	absent	absent	present	absent
geminata group	absent	absent	present absent	prs; abs	absent	absent	present	absent
pulchella group	present	absent	absent absent	absent	absent	absent	present	absent
inda group	absent	prs; abs	absent absent	absent	absent	absent	prs; abs	absent
areata group	absent	present	absent absent	absent	absent	absent	present	absent
discicollis group	absent	present	present absent	absent	absent	present	present	absent
verticalis group	absent	present	absent present	absent	absent	absent	present	absent