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INTRODUCTION
Scolia Fabricius, 1775 is a cosmopolitan genus of fossorial solitary wasps in the subfamily Scoliinae presently containing 20 described species for America (Hanson, 2006). Scolia guttataBurmeister 1853 was the only described species recorded for Central America (Cameron, 1893).
Scoliid larvae are ectophagous idiobiont parasitoids of large Coleoptera larvae, including Sacarabaeidae, Passalidae, Lucanidae, and Curculionidae (Fernández, 2006; Kumar, 2009), for which they have been used as biological control agents (Clausen, 1978).
In this paper, we review the genus Scolia from Central America. Also, we present seasonal flight records for Scolia guttata on Barro Colorado Island, Panama.
MATERIALS AND METHODS
Flight seasonality data come from the field station of the Smithsonian Tropical Research Institute (STRI) on Barro Colorado Island (BCI), located in Gatun Lake of the Panama Canal, 9° 09'17" N; 79° 50' 53" W. The seasonal flight activity of Scolia guttata was studied by sampling specimens from ten Malaise traps (Townes modified) installed at ground level in the old forest by the Smithsonian Environmental Studies Program (see Richards and Windsor, 2007). The samples were collected weekly, from January 2005 to December 2006. For other information of the study area, see Windsor (1990) and Cambra et al. (2018).
The following acronyms are used for the collections where type specimens and other materials are deposited:
BMNH The Natural History Museum, London, United Kingdom.
INBio Instituto Nacional de Biodiversidad (currently Museo Nacional de Costa Rica).
MIUP Museo de Invertebrados G.B. Fairchild, Universidad de Panamá.
MLUH Zoological Institute of the University of Halle, Germany.
ZMUC Natural History Museum of Denmark, University of Copenhagen.
RESULTS AND DISCUSSION
Scolia (Discolia) guttataBurmeister, 1853 (Fig.1a-1c, Fig. 2a-2c)
Figure 1 a-c. Scolia guttata, a, b. Habitus, male a. lateral view, b. Habitus, Dorsal view, c. Head, frontal view, female, d-f. Scolia rufiventris, female, d. Habitus, lateral view, e. Habitus, dorsal view and lectotype labels, f. Head, frontal view.
Figure 2 Genitalia, males. a-c. Scolia guttata, a. Dorsal view, b. Ventral view, c. Penis valve, lateral view. d-f. Scolia rufiventris, d. Dorsal view, e. Ventral view, f. Penis valve, lateral view.
Scolia guttataBurmeister, 1853: 36, lectotype female (Designation by Bradley and Brethem, 1966), MLUH.
Discolia hecate Kirby, 1889: Bradley and Brethem (1966).
Diagnosis. Integument mostly black (Fig. 1a-1c), terga two-three with lateral yellow spots (Fig. 1a); female with the anterior margin of clypeus convex (Fig. 1c); paramere apex angulate (Fig. 2a-2b); penis valve with small spines on dorsal margin (Fig. 2c).
Material examined. GUATEMALA (no other data), one male (MIUP). COSTA RICA (98 males, 56 females, INBio, see table 1): Alajuela: Est. San Ramón, 3-19.iv.1994, F. Quesada, one female; same data except: 13-30.viii.1996, D. Briceño, one male; P.N. Arenal, San Carlos, R.F. Arenal, Sen. Mena, 600 m, 16.vii.2001, G. Cabarllo, one female; same data except: Send. Pilon, 650 m, 23.vii.2001, two females; 14-25.viii.2001, one male, two females. Cartago: Grano de Oro, 1120 m, 8-30.viii.1992, P. Campos, two females; same data except: Monumento Nacional Guayabo, 1100 m, vii.1994, G. Fonseca, one female. Guanacaste: P.N. Guanacaste, Est. Biol. Cacao, 1000-1400 m, SW side Vol. Cacao, ix.1989, R. Banco and C. Chaves, one male; same data except: 1100 m, xii.1993-ix.1994, G. Chavarría, one male; 1200 m, vi.1997, R. Moraga, four females; Est. Los Almendros, 13 Oct-3.xi.1993, K. Martínez, nine males; Est. Maritza, 600 m, viii.1990, four males, eight females; same data except: 27.ii-10.iii.1994, F. A. Quesada, one female; Est. Murciélago, 100 m, 3-12.xii.1993, F. A. Quesada, one male; Est. Pitilla, 700 m, nine km S Sta. Cecilia, ix.1988, two males; same data except: x.1988, 16 males, one female; xi. 1988, 23 males, 11 females; i.1989, four females; 21 .iii-21. iv.1989, one male; x.1989, C. Moraga and P. Ríos, three males; i.1991, one female; 2-19.iii.1992, P. Ríos, one male; 18 iv-9.v.1993, C. Moraga, one male; iii.1994, C. Moraga, one male; Est. Sta. Rosa, 300 m, vii.1992, R. Moraga, two females; same data except: 1.v.1995, F. A. Quesada, one male; Fca. Jenny, 30 km N Liberia, ix.1988, one male, one female; same data except: x.1988, two males, three females; 5-20.ix.1993, E. Araya, one male; 6-15.xi.1993, one male; ix.1998, three males; Fca. Loaiciga, 500 m, 6-28.i.1992, C. Moraga, one female; La Cruz, Cerro El Hacha, 26-30. vii.1986, I. Chacón, two males; P.N. Rincón de la Vieja, Est. Las Pailas, 800 m, 15.vii-14.ix.1992, J. Sihezar, one female; same data except: 27.vii-15.viii.1992, C. Cano, eight males, five females; 24.viii-14.ix.1992, one male, one female; 5-24.viii.1994, D. García, one male. Limón: R.N.F.S. Barra Del Colorado, 50 m, vi.1994, F. Araya, one female. Puntarenas: P.N. Corcovado, Est. Agujas, Send. Zamia, 300 m, 10.ii.1998, A. Azofeifa, one male; same data except: La Bonanza, 495 m, 4-22.v.1999, J. Azofeifa, one female; Est. Sirena, 100 m, ii.1990, G. Fonseca, one male, one female; Rancho Quemado, Península de Osa, 200 m, viii.1991, F. Quesada, one male; same data except: 7-27.i.1992, A. Gutiérrez, one male; ii.1992, F. Quesada, three males; 1-20. xii. 1993, A. Marin, one male. San José: Reserva Privada, Universidad para La Paz, Fila Diamante, 1021 m, 8.vii.2001, M. Solís, one male. PANAMA: Panamá Province: Barro Colorado Island, ten Malaise traps, 2005-2006, col. D. Windsor, 137 males, one females (MIUP); Parque Nacional Soberanía, camino Plantación, 3.viii.2001, D. Quintero, R. Cambra, one female (MIUP).
Distribution: U.S.A. (Arizona, Texas) to Venezuela (Cameron, 1893; MacKay, 1987; Maes, 1989) (Fig. 3).
Comments: From Costa Rica (years 1986 to 2001) we examined 154 specimens (98 males and 56 females) of S, guttata; the specimens were collected mostly with hand nets during November (33 males /11 females), August (19 / 17) and October (18/ 4), the three months with greatest numbers of specimens captured (102 specimens = 66.2 %). These data suggest that adults of S, guttata are present in all months of the year. However, this information gives an inadequate indication of flight seasonality because the data were not collected with equal effort across the year.
Scolia (Hesperoscolia) rufiventris Fabricius, 18G4 (Fig. 1d-1f, Fig. 2d-2f)
Scolia rufiventris Fabricius, 1804. Lectotype female (designated by J. C. Bradley, 1929), Essequibo [Guyana]. Catalogue number 00241358 ZMUC.
Scolia (Lacosi) ancepsSaussure 1858, type female, Mexico: Osten (2005).
Scolia (Lacosi) drewseniSaussure 1858, type female, Brazil, Minas Gerais: Osten (2005).
Ycasbraia Argaman 1996. Type species: Scolia rufiventris Fabricius 1804. = Hesperoscolia Bradley, 1974: objective synonym (Kimsey and Brothers, 2016).
Diagnosis. Head and mesosoma black, metasoma and legs mostly orange-red (Fig. 1d-1e); female with the anterior margin of clypeus with a triangular projection medially (Fig. 1f); paramere apex rounded (Fig. 2d-2e); penis valve with large and small projections on dorsal margin (Fig. 2f).
Description. MALE. Body length 17.5 mm, forewing 14 mm. Color: integument of head, mesosoma and antenna black except eye inner emargination with small yellow spot; basal halfofmandible black, apex mostly orange; metasoma, legs and tibial spurs orange-red except coxa black; wings infuscate, with feeble violaceous reflection; vertex and gena with sparse, semi-erect, black setae; frons, clypeus and scape with fulvous setae; pronotum, mesoscutum and scutellum mostly with black setae, propodeum, meso and metapleuron mostly with short white pubescence and few semierect black setae; legs and metasoma with semi-erect fulvous setae. Head: mostly with dense, medium-sized, close punctures except gena with small sparse punctures and discal area of clypeus impunctate; head in dorsal view, including eyes, 0.69 x maximum wide of mesosoma; ocelli small, distance between eye margin and lateral ocellus 0.59 x as long as maximum diameter of ocellus; clypeus convex mesally, its anterior margin straight; scape not carinate, 1.66 x as long as wide; first flagellomere 1.10 x as long as second. Mesosoma: with dense, medium-sized, close punctures; antero-dorsal area of propodeum extends backward 1.0 x length of scutellum then sharply bends downward forming nearly right angle; posterior area of propodeum strongly concave; meso and metatibiae with a row of spines in dorsal area, their apices with four spines, not including tibial spurs. Metasoma: First metasomal segment, in dorsal view, 0.67 x as long as the second; paramere and penis valve as fig. 2d-2f.
Material examined. COSTA RICA (28 males, 22 females, INBio): Guanacaste: P. N. Guanacaste, Est. Agua Buena, 220 m, 8.ii-1.iii.1993, E. López, one female; Est. Cacao, 1000-1400 m, Lado SO Vol. Cacao, iii.1991, C. Chaves, one female; same data except: Sector Cacao, 1200 m, vi.1997, R. Moraga, one male, one female; Est. Los Almendros, 0 m, E. López, 15-27.viii.1994, one female; same data except: 300 m, 3-23.xii.1994, E. López, one female; Est. Pitilla, 700 m, nine km S Sta. Cecilia, 18.iv-9.v.1993, C. Moraga, two males; same data except: iv.1994, P. Ríos, one female; Fca. Jenny, 30 km N Liberia, 1.x.1988, two males; same data except: x.1988, two males, nine females; vii.1991, R. Espinoza, one female; P.N. Rincón de la Vieja, Est. Las Pailas, 800 m, 9-27.ii.1993, K. Taylor, one male; same data except: 16-24.viii.1993, D. García, one male. Limón: P. N. Tortuguero, Cerro Tortuguero, 100 m, iv.1989, R. Aguilar and J. Solano, two females; same data except: iii.1993, R. Delgado, one male; R.B. Hitoy Cerere, 100 m, vii.1992, G. Carballo, one male; same data except: Talamanca, Cerro Bitarkara, 1025 m, 15-17 x.2004, Y. Cárdenas, one female; Veraguas Rainforest, Restaurant, 400-440 m, ix.2008, R. Villalobos, one male. Puntarenas: Buenos Aires, P. Int. La Amistad, 1766 m, 5-9.iii.1996, R. Villalobos, one male; same data except: Sendero a Casa Coca, 1700 m, 7-25.xi.2002, D. Rubi, two males, one female; Est. Agujas, Sendero Zamia, 300 m, 1-7.xii.1997, A. Azofeifa, two males; Est. Biol. Las Alturas, 1500 m, x.1991, M. Zumbado, one male, one female; same data except: Sendero Cerro Echandi, Coto Brus, xii.1997, B. Gamboa, one male; Sendero R. Bella Vista, 1400 m, 3-5.iii.1992, M. Zumbado, one male; Est. Cerro de Oro, Sendero Guapinol, 345 m, 10.i.1996, L. Angulo, two males; Est. Pittier, 1670 m, 5-18.i.1995, L. Angulo, one male; Est. Sirena, one-100 m, i.1995, G. Fonseca, one female; Cerro Biolley, Sector Altamira, La Amistad, 1800 m, 4.iii.1995, M. Segura, one male; Pta. Llorona, Sendero a San Pedrillo, 100 m, 2-3.iii.1997, J. Silva and E. Fletes, one male; Rancho Quemado, 200 m, Península de Osa, ix.1991, F. Quesada, two males. San José: Est. Santa Elena, Sendero La Bota, 1690 m, 25.iv.1996, E. Alfaro, one male. PANAMA: Panamá Province: Cerro Azul, Bosque en Residencial Las Nubes, 15.iii.1999, one male (MIUP); ciudad de Panamá, El Cangrejo, 6.x.2015, G. De La Guardia, one female (MIUP).
Distribution: Mexico (without other data) (Saussure, 1858), Costa Rica, Panama, Colombia (First records), Guyana, Peru, Brazil (Mato Grosso and Minas Gerais) (Fabricius, 1804; Saussure, 1858; Fox, 1896; Dos Santos et al., 2015) (Fig. 3).
Comments: The two species of Scolia present in Central America, S. guttata and S. rufiventris, are differentiated by the diagnostic morphological characters presented above. Fernández and Cubillos, (1999) examined specimens of Scolia sp. 1 from Costa Rica (INBio) and Colombia (Antioquia, Cali, Valle, Dagua) and gave a brief description of Scolia sp. 1 that agrees with the description of Scolia rufiventris. Considerable effort was made to locate specimens collected from Colombia without positive results. Existing evidence (morphology and distribution) indicates that the specimens mentioned by Fernández and Cubillos (1999) are S. rufiventris.
Seasonal flight activity for Scolia guttata from Barro Colorado Island, Panama
During the two continuous sampling years (2005-2006) in BCI, 144 specimens of Scolia guttata (137 males and seven females) were collected, 106 individuals during 2005, 38 individuals in 2006. Collections peaked during June with 127 specimens (88.2 %), the remainder captured during May, July, and December (Fig. 4). All seven females from BCI were captured in June. No S, rufventris were recovered from the same BCI trap samples. The much smaller catch of S, guttata females relative to that of males in BCI could be the result of sampling methodology. Female Scoliidae typically spends a long time in the ground digging for larval hosts. For this reason, they are seldom collected with Malaise traps. However, males spend more time flying, mainly in search of females, thereby increasing chances of capture by Malaise traps. The under-representation of females in Malaise traps is not seen in the hand net collections from Costa Rica (INBio) where we had 56 females and 98 males of S, guttata.
Figure 4 Total of Scolia guttata specimens captured by month in two years (2005-2006) with ten Malaise traps in BCI.
The hosts of S, guttata and other Scolia species in the Neotropical region remain unknown. However, the hosts of some Nearctic Scolia species have been reported. Rau and Rau (1918) recorded Cotinisnitida? (L., 1758) (Scarabaeidae: Cetoninae) as a host of S. dubia dubia Say 1837, and Ryckman (1956) reported a larva of Scarabaeidae as the host of S. nobilitata otomitaSaussure, 1858. The greater abundance of S. guttata in BCI during June may be related to a greater abundance of scarabaeid larvae present in the soil and available to be parasitized during the early wet season. Ratcliffe, (2003) notes the abundance of some Dynastinae scarab beetles on BCI is greater in May, June and July (E.g. Cyclocephala species). Further biological and abiotic information is necessary to better correlate flight seasonality and annual abundance of S. guttata in BCI.
