Hoplopyga Thomson, 1880

Shaughney, Jennifer Marie & Ratcliffe, Brett C., 2015, A Monographic Revision of the Genus Hoplopyga Thomson, 1880 (Coleoptera: Scarabaeidae: Cetoniinae: Gymnetini), The Coleopterists Bulletin 69 (4), pp. 579-638 : 582-586

Hoplopyga Thomson, 1880

Hoplopyga Thomson 1880: 268 .

Type species:

Gymnetis marginesignata Gory and Percheron 1833: 72 and 366, by original designation.

Description. Scarabaeidae , Cetoniinae, Gymnetini. Form: Length 10.1–21.1 mm. Body shape elongate, rhomboidal, robust. Dorsum velutinous, opaque, or shiny. Color and pattern highly variable, ranging from golden yellow to brownish yellow to various shades of brown, green, gray, maroon, or black; dorsum with small to large, reddish brown to black spots or with fuscous to piceous clouding at base of head, on pronotum, and on elytra; venter opaque, shiny, or weakly metallic, ground color similar to dorsum, with portions of metasternum and sternites reddish brown to piceous, shiny; punctures reddish brown to black. Head: Subrectangular. Surface longitudinally convex, slightly depressed either side of middle, with lateral margins weakly to moderately raised. Anterior margin of clypeus weakly to strongly reflexed, weakly to strongly emarginate, rounded or angulate either side of emargination. Eye canthus with dense, minute setae on posterior margin and at apex. Punctures each with a minute to short seta. Interocular width variable intraspecifically. Antennae 10-segmented. Pronotum: Subtrapezoidal, lateral margins gradually widening from apex to base or obtusely angulate. Surface punctate, punctures each with a minute seta. Basomedian lobe obscuring all but tip of scutellum. Lateral margins with complete or interrupted, narrow, reddish brown to black bead or not. Elytra: Widest at base, with sides weakly tapering towards apex. Surface with costae as follows: 2 weakly elevated, complete discal costae or 2 distinctly elevated discal costae and with lateral costa on each elytron depressed on mediodiscal area. Lateral margins with narrow, reddish brown bead. Apices at suture produced into weak to distinct spines. Pygidium: Surface weakly to strongly convex, with large, dense, vermiform or n-shaped punctures, punctures and apical margin with minute to short setae. Venter: Mesometasternal process, in lateral view, subparallel to horizonal axis of body or at a slight, oblique angle to horizontal axis of body, rounded, weakly to moderately protuberant beyond mesocoxae; in ventral view, apex subacuminate or rounded. Metasternum and sternites with setigerous punctures. Males with abdomen slightly concave in lateral view, females with abdomen flat or weakly convex. Legs: Protibia with 1–3 teeth. Meso- and metatibiae with row of long, dense setae on inner surface. Metatibia at apex with 2 spurs; females with longer spur broad, rounded at apex, males with longer spur slender, acute. Parameres: In caudal view, form usually subrectangular, shaft divergent between midpoint and apex or not, apices each with weak to distinct lateral spur.

Diagnosis. Hoplopyga species can be distinguished from other genera of Neotropical Gymnetini by a combination of the following characters: lack of armature on the head; clypeal apex reflexed and never deeply bilobed; dorsum glabrous or with sparse, minute or short setae; and sutural apices of the elytra usually prolonged into spines. Most Hoplopyga species have a mesometasternal process that is short, subparallel to the horizontal axis of the body and rounded in ventral view, but there are a few exceptions. Hoplopyga species can be characterized in one of two ways. “Typical” Hoplopyga species have a complete, medial, discal costa on each elytron and a lateral, discal costa that is depressed on the mediodiscal area. These species also have large, n-shaped punctures on the elytra arranged longitudinally in striae in the depressions between the elytral costae. The typical Hoplopyga species resemble other New World genera that lack armature on the head, namely Gymnetis , Hologymnetis Martínez , Hoplopygothrix Schürhoff , and Neocorvicoana Ratcliffe and Micó. The “atypical” Hoplopyga species have indistinct elytral costae or have two weakly raised, complete costae. These species also have large, n-shaped punctures on the elytra, but each puncture is enclosed in a reddish brown to black spot, giving these species a spotted appearance.

“Typical” Hoplopyga species can be distinguished from Gymnetis (sensu lato) species by the elytral costae. In Hoplopyga species , the sutural costae are as described above. In Gymnetis species , the discal costae are complete and weakly raised or arise on the posterior half of the elytra, and the lateral costa of each elytron is not interrupted. The form of the mesometasternal process can usually be used to distinguish the two genera, but this character state should only be relied upon in combination with other characters because there are species in both genera that show variation in the angle and protuberance of the mesometasternal process (e.g., the lectotype of Hoplopyga foeda (Schaum) has a mesometasternal process that is deflexed downward and subquadrate at the apex in lateral view, while most other H. foeda specimens have a mesometasternal process, in lateral view, that has a rounded apex and is subparallel or at an oblique angle to the horizontal axis of the body). Generally, Hoplopyga species have a mesometasternal process that is weakly or moderately protuberant beyond the mesocoxae, subparallel or at an oblique angle to the horizontal axis of the body, and rounded at the apex in lateral view. The mesometasternal process in Gymnetis species is usually strongly protuberant beyond the mesocoxae and, in lateral view, deflexed or at an oblique angle to the horizontal axis of the body with the apex subquadrate or bulbous.

Hoplopyga can be easily separated from Hologymnetis species by the presence of a vertical, prosternal throat spur that is absent in Hologymnetis species.

Hoplopyga is similar to Hoplopygothrix and Neocorvicoana , but Hoplopyga is usually glabrous dorsally or with sparse, minute or short setae. Hoplopygothrix and Neocorvicoana species have a dorsum that is covered with short, dense setae. In addition, Hoplopygothrix and Neocorvicoana each have a unique, sexually dimorphic character that is not present in Hoplopyga . Hoplopygothrix species males have the abdominal sternites deeply, longitudinally sulcate at the middle, and Hoplopyga species males do not. Neocorvicoana species females have the tarsomeres shortened and compact, and Hoplopyga species females do not.

Several of the “atypical”, spotted Hoplopyga species resemble species in the genera Marmarina Kirby and Macrocranius Schürhoff. Spotted Hoplopyga species can be distinguished from Marmarina maculosa (Olivier) by having a tridentate protibia in both sexes and/or beaded lateral margins of the pronotum. In addition, the form of the mesometasternal process can be used to differentiate the genera. Spotted Hoplopyga species have a mesometasternal process that is weakly protuberant beyond the mesocoxae, subparallel to the horizontal axis of the body, and with a rounded apex in lateral view. Marmarina maculosa has a mesometasternal process that is moderately protuberant beyond the mesocoxae, at an oblique angle to the horiztonal axis of the body, and with the apex rounded or subquadrate in lateral view. Hoplopyga species can be distinguished from Macrocranius similis Schürhoff by the shape of the head. Hoplopyga species have a subrectangular or subquadrate head, while M. similis has the apicolateral margins of the clypeus rounded. In addition, the apices of the elytra at the suture are not spinose in M. similis . The male parameres of spotted Hoplopyga species are also distinct and distinguish these species from Marmarina or Macrocranius species.

When delineating species limits, we considered transferring the spotted species to either Macrocranius or Marmarina . We determined that these species do not belong in Macrocranius because of the rounded head of M. similis , whereas the spotted Hoplopyga species all have subrectangular heads, which is a character state also exhibited by the typical Hoplopyga species . These species do not belong in Marmarina because of the characters listed above, as well as the lack of distinct, elytral costae that are present in some Marmarina species. We leave these species in the genus Hoplopyga in the interest of parsimony and, more importantly, because of the elytral apices of H. multipunctata . This is one of the spotted species, and yet it has the elytral apices distinctly prolonged into spines at the suture. This is the principal character state for which Thomson originally described the genus, making it logical to leave H. multipunctata within Hoplopyga . If H. multipunctata remains in the genus, it follows that the other spotted species should remain as well.

Etymology. The name for this genus is Greek in origin and derives from the prefix hoplo, meaning armed, and the suffix pyga, meaning posterior. We believe this refers to the spinose sutural apices of the elytra that Thomson thought to be a defining character for this genus.

Distribution. Hoplopyga species occur from central Mexico to northern Argentina. One species is known from the West Indies.

Natural History. Adults have been found from elevations ranging from sea level to 3,600 m and are generally diurnal, although several species are frequently caught at lights in the early hours of the morning. Adults are attracted to rotting fruit and can be found feeding on fruits, foliage, flowers, and sap flows. Traps baited with old plantains or bananas are particularly effective in attracting specimens. The immature stages of three species have been described ( Vanin and Costa 1984; Micó et al. 2001; Morón and Arce 2002), and the immature stages of several species are known to live as inquilines in termite nests ( Luederwaldt 1911; Micó et al. 2001; Puker et al. 2012; label data) or in decomposing tree trunks.

Nomenclature. Gaston Ruter designated a holotype, allotype, and paratypes at the Museum für Naturkunde in Berlin, Germany for a new species, Hoplopyga hiekei , but never published the species name. The specimens were an assortment of Hoplopyga albiventris ( Gory and Percheron, 1833) , Hoplopyga singularis ( Gory and Percheron, 1833) , and Hoplopyga gosseti Antoine, 2008 from Brazil and Paraguay.

Two species have been removed from Hoplopyga . Gymnetis lugubris Thomson, 1878 was included in Hoplopyga in two catalogs ( Blackwelder 1944; Krajcík 1998) after Schürhoff (1937) noted that G. lugubris is “probably a large, black Hoplopyga ”, but this species was determined to be a junior synonym of Allorrhina nigerrima ( Burmeister, 1842) based on examination of the G. lugubris holotype at MNHN ( Ratcliffe 2015). In addition, we have examined photographs of the holotype of Hoplopyga ruteri Antoine, 2008 at MNHN and determined this species to be conspecific with Macrocranius similis Schürhoff, 1935 due to the similarity in gestalt and the form of the male parameres. Checklists by Schoch (1895a) and Schenkling (1921) included the species Cetonia reticulata Kirby, 1818 in Hoplopyga , but Ratcliffe and Micó (2001) established the genus Neocorvicoana for this species and two others.

CHECKLIST OF HOPLOPYGA SPECIES

Hoplopyga aequatorialis ( Moser, 1918)

Hoplopyga albiventris ( Gory and Percheron, 1833) Gymnetis fuscorubra Gory and Percheron, 1833 (synonym)

Hoplopyga antilliana Ratcliffe, 2012

Hoplopyga boliviensis ( Moser, 1918)

Hoplopyga brasiliensis ( Gory and Percheron, 1833)

Gymnetis prothoracica Thomson, 1878 (synonym)

Hoplopyga cerdani Antoine, 1998

Hoplopyga foeda ( Schaum, 1848)

Hoplopyga lucidiventris ( Thomson, 1878) , new synonymy

Hoplopyga gosseti Antoine, 2008

Hoplopyga liturata ( Olivier, 1789)

Gymnetis spinosa Fischer von Waldheim, 1823 (synonym)

Gymnetis hamata Fauvel, 1861 (synonym)

Hoplopyga spurca ( Janson, 1880) , new synonymy

Hoplopyga marginesignata ( Gory and Percheron, 1833)

Gymnetis fumata Janson, 1880 (synonym)

Hoplopyga miliaris ( Gory and Percheron, 1833)

Gymnetis fodina Gory and Percheron, 1833 (synonym)

Gymnetis suasa Gory and Percheron, 1833 (synonym)

Hoplopyga miniata ( Blanchard, 1846)

Hoplopyga multipunctata ( Gory and Percheron, 1833)

Holopyga [sic] multiguttata Schoch, 1895 (synonym)

Hoplopyga ocellata ( Gory and Percheron, 1833)

Hoplopyga peruana ( Moser, 1912)

Hoplopyga pseudomiliaris Shaughney and Ratcliffe , new species

Hoplopyga ravida ( Janson, 1881)

Hoplopyga riparia Shaughney and Ratcliffe , new species

Hoplopyga singularis ( Gory and Percheron, 1833)

Hoplopyga monacha ( Gory and Percheron, 1833) , new synonymy

Hoplopyga rubida ( Gory and Percheron, 1833) , new synonymy

Hoplopyga corumbana Schürhoff, 1942 (synonym)

Hoplopyga suilla ( Janson, 1881)

KEY TO THE SPECIES OF ADULT HOPLOPYGA THOMSON, 1880

1. Elytra with 2 complete, weakly raised costae or costae indistinct. Dorsum spotted........... 2

1′. Each elytron with 2 distinctly raised, discal costae; lateral costa on each elytron interrupted and depressed on mediodiscal area .............................................................. 6

2. Spots on dorsum coalescing on base of pronotum, base of elytra, and on apical umbone of each elytron ( Figs. 77–78 ). Elytral apices at suture strongly spinose. Argentina, Brazil, Paraguay....................................... ............... Hoplopyga multipunctata (Gory and Percheron)

2′. Spots on dorsum not coalescing as above. Elytral apices at suture subquadrate..........3

3. Puncture enclosed in each colored spot on dorsum moderate in size or large, horseshoeshaped, easily seen with the naked eye....... 4

3′. Puncture enclosed in each colored spot on dorsum minute or small, n-shaped, not easily seen with the naked eye, punctures on dorsum too numerous to count. French Guiana.......... ........................... Hoplopyga cerdani Antoine

4. Dorsum reddish orange ( Fig. 73 ) and venter entirely reddish brown, shiny. Bolivia....... .............. Hoplopyga miniata (Blanchard)

4′. Dorsum yellow or brownish yellow, venter similar to dorsum but enamel-like and weakly metallic...................................... 5

5. Pygidium with n-shaped punctures clustered at base and in 2 spots either side of middle. Brazil, Paraguay........................................... .... Hoplopyga miliaris (Gory and Percheron)

5′. Pygidium with n-shaped punctures irregularly spaced, covering entire surface of pygidium. Guatemala............................... ........ Hoplopyga pseudomiliaris Shaughney and Ratcliffe

6. Protibia with 2 prominent distal ( Fig. 8 ); remaining teeth may be reduced ...............7

6′. Protibia each with 1 prominent apical tooth ( Fig. 45 ); remaining teeth may be reduced.................................................................8

7. Pronotum with lateral margins subparallel on posterior half. Mesometasternal process entirely reddish brown, shiny, or with 1 spot at base. Anteromedial margin on penultimate abdominal sternite with reddish brown, shiny band not reaching posterior margin of sternite, or with band narrowing significantly before posterior margin. Paraguay............. ...................... Hoplopyga gosseti Antoine

7′. Pronotum with lateral margins obtusely angulate. Base of mesometasternal process with 2 yellow spots at base (spots rarely absent). Anteromedial margin on penultimate abdominal sternite with reddish brown, shiny spot or middle of penultimate sternite reddish brown, shiny. Argentina, Brazil, Paraguay.................... Hoplopyga albiventris (Gory and Percheron)

8. Elytra with yellow or orangish yellow scalloping on lateral and apical margins ( Figs. 61–62 ). Northern South America................................ ............... Hoplopyga marginesignata (Gory and Percheron)

8′. Elytra without yellow scalloping...............9

9. Mesometasternal process, in ventral view, with lateral margins expanding just before apex (e.g., Figs. 26 and 111 )...................10

9′. Mesometasternal process, in ventral view, with lateral margins not expanding before apex (e.g., Fig. 2 )..................................... 11

10. Elytra with discal area mostly monocolored, and each elytron with oblique, fuscous line extending from humeral umbone to mediodiscal area ( Figs. 24–25 ). South America ...... ....................... Hoplopyga brasiliensis (Gory and Percheron)

10′. Elytra with discal area mottled in appearance, with at least 2 distinct colors, and lacking oblique, fuscous line ( Fig. 109 ). Brazil, Peru.................... Hoplopyga riparia Shaughney and Ratcliffe

11. Mesometasternal process with numerous minute, round punctures at base, punctures sometimes with a short or long seta........12

11′. Mesometasternal process without numerous minute, round punctures at base..............15

12. Pronotum with most punctures enclosed in a large, round, black spot. Females with abdominal sternites entirely black, shiny, and with distinct, brownish yellow, posterolateral spots on each sternite. Bolivia.......... .................. Hoplopyga boliviensis (Moser)

12′. Pronotum with punctures not enclosed in a spot. Sternites on females lacking distinct, brownish yellow, posterolateral spots......13

13. Punctures between apical umbone and suture on each elytron greatly reduced in size and density or absent. Male protibia with only 1 distinct, apical tooth and female protibia tridentate. Females with apical declivity of elytra sulfur yellow. Western South America, from Colombia to Bolivia................... Hoplopyga peruana (Moser)

13′. Punctures between apical umbone and suture on each elytron not reduced in size and density. Males and females with tridentate protibia. Females with apical declivity of elytra colored as on rest of dorsum.......... 14

14. Body robust or elongate, with 1or a combination of the following: head with short setae arising from punctures; base of mesometasternal process with long, dense setae; pygidium with “fuzzy” appearance in lateral view due to numerous short setae arising from punctures. Andean mountains in northern and western South America ....................... ..................... Hoplopyga foeda (Schaum)

14′. Body elongate, with punctures on head lacking setae, base of mesometasternal process without long, dense setae, and pygidium with minute setae arising from punctures. Ecuador, Peru............................................. ............ Hoplopyga aequatorialis (Moser)

15. Color of dorsum orangish yellow. Pronotum with narrow, longitudinal, orangish yellow band on midline. Abdominal sternites mostly reddish brown, shiny, each with distinct, orangish yellow spots on posterolateral corners of each sternite. Northern Brazil, Trinidad, Venezuela................................... ..................... Hoplopyga ravida (Janson)

15′. Color of dorsum brownish yellow, olivebrown, cinereous, greenish gray, or a combination of earth-colored hues. Pronotum with narrow, longitudinal band on midline, on midline posterior to M-shaped mark, or with band absent. Abdominal sternites not as above....................................................16

16. Dorsum entirely cinerous, piceous, or dark, greenish gray. Venter entirely reddish brown or black, shiny. Pronotum with sparse to moderately dense punctures. Abdominal sternites with elongate, weakly n-shaped, shallow punctures on lateral thirds, punctures each with a short seta. Trinidad, Venezuela....................... Hoplopyga suilla (Janson)

16′. Dorsum not colored as above, or colored as above with the following characteristics: venter with portions of metasternum and/or abdominal sternites greenish gray, brownish yellow, cream-colored, or caesious. Pronotum with moderately dense to dense punctures. Abdominal sternites with distinctly n-shaped, deep punctures, punctures on lateral thirds or continuing across middle of each sternite, punctures each with a minute seta ............. 17

17. Punctures between medial, discal costa and sutural costa on each elytron becoming smaller and extending to base near scutellum in 2 columns. Pygidium with punctures not usually reaching apex. Metasternum with impunctate spot on posterolateral corners. Argentina, southern Brazil, Paraguay, rarely in northern South America................ ....................... Hoplopyga singularis (Gory and Percheron)

17′. Punctures between medial, discal costa and sutural costa on each elytron ending on mediodiscal area or just past mediodiscal area on each elytron, not extending to base near suture................................................18

18. Head with 2 clearly defined, pitchy spots at base. Brownish yellow color of dorsum mostly obscured by black to piceous clouding and flecks on middle of pronotum and elytra ( Fig. 13 ). West Indies........................ ................... Hoplopyga antilliana Ratcliffe

18′. Head with fuscous to pitchy clouding at base either side of middle or not.............19

19. Fuscous clouding on middle of pronotum with straight, clearly defined edges, even on dark specimens ( Figs. 51 , 57–58 ). Mexico to Argentina..... Hoplopyga liturata (Olivier)

19′. Fuscous clouding on middle of pronotum with wavy or indistinct edges. Elytra each with fuscous or piceous spot or band between apical umbone and suture, giving appearance of hourglass shape on elytra ( Figs. 83–84 ). Dark specimens with mediodiscal area on each elytron cream-colored with distinct black spot. Mexico to Paraguay.......................... ........................ Hoplopyga ocellata (Gory and Percheron)

Figures

Figs. 77–82. Hoplopyga multipunctata. 77–78) Habitus; 79) Mesometasternal process, ventral view; 80–81) Parameres; 82) Distribution.

Figs. 73–76. Hoplopyga miniata, holotype (Photographs courtesy of A. Mantilleri, MNHN). 73) Dorsal view; 74) Ventral view; 75) Labels; 76) Distribution.

Figs. 7–12. Hoplopyga albiventris. 7–8) Habitus; 9) Mesometasternal process, ventral view; 10–11) Parameres; 12) Distribution.

Figs. 45–50. Hoplopyga gosseti. 45–46) Habitus; 47) Mesometasternal process, ventral view; 48–49) Parameres; 50) Distribution.

Figs. 61–66. Hoplopyga marginesignata. 61–62) Habitus; 63) Mesometasternal process, ventral view; 64–65) Parameres; 66) Distribution.

Figs. 24–29. Hoplopyga brasiliensis. 24–25) Habitus; 26) Mesometasternal process, ventral view; 27–28) Parameres; 29) Distribution.

Figs. 109–114. Hoplopyga riparia. 109–110) Habitus of allotype female; 111) Mesometasternal process, ventral view; 112–113) Parameres; 114) Distribution.

Figs. 1–5. Hoplopyga aequatorialis. 1) Habitus; 2) Mesometasternal process, ventral view; 3–4) Parameres; 5) Distribution.

Figs. 13–17. Hoplopyga antilliana. 13) Habitus; 14) Mesometasternal process, ventral view; 15–16) Parameres; 17) Distribution.

Figs. 51–56. Hoplopyga liturata. 51) Habitus; 52–53) Variation in mesometasternal process, ventral view; 54–55) Parameres; 56) Distribution.

Figs. 57–59. Hoplopyga liturata. 57–58) Dorsal color variation; 59) Adult H. liturata in French Guiana (photograph courtesy of Julien Touroult).

Figs. 83–88. Hoplopyga ocellata. 83–84) Habitus; 85) Mesometasternal process, ventral view; 86–87) Parameres; 88) Distribution.