TRABAJOS CIENTÍFICOS

Status of the forensically important genus Ophyra (Diptera: Muscidae) in Argentina

Estado del género de importancia forense Ophyra (Diptera: Muscidae) en Argentina

Patitucci, Luciano D.^*,**, Pablo R. Mulieri^*,**, Adriana Oliva^*,*** and Juan C. Mariluis^*,**

* CONICET, Buenos Aires, Argentina.
** Departamento Vectores-CeNDIE, ANLIS "Dr. Carlos G. Malbrán", Av. Vélez Sarsfield 563, C1281AFR, Buenos Aires, Argentina; e-mail: lpatitu@yahoo.com.ar, mulierii@yahoo.com, jcmariluis@yahoo.com.ar
*** Laboratorio de Entomología Forense, Museo Argentino de Ciencias Naturales, Av. A. Gallardo 470, C1405DJR, Buenos Aires, Argentina; e-mail: aoliva@macn.gov.ar

RESUMEN: El género Ophyra Robineau-Desvoidy es un grupo de múscidos necrófagos distribuidos en los climas cálidos de todo el mundo. La información aquí presentada se basa en la recopilación de datos de distribución, obtenida a partir del material de diferentes colecciones y bibliografía para la Argentina. Ophyra albuquerquei Lopes, Ophyra capensis (Wiedemann), Ophyra chalcogaster (Wiedemann) y Ophyra solitaria Albuquerque se registraron por primera vez para el país. Se presenta una clave para las especies argentinas. Se discuten los datos biológicos y forenses de las distintas especies.

PALABRAS CLAVE: Ophyra; Muscidae; Entomología forense; Argentina.

ABSTRACT: The genus Ophyra Robineau-Desvoidy is a necrophagous group of Muscidae distributed in warm climates worldwide. The information here presented is based on the compilation of distributional data obtained from material of different collections and bibliography for Argentina. Ophyra albuquerquei Lopes, Ophyra capensis (Wiedemann), Ophyra chalcogaster (Wiedemann) and Ophyra solitaria Albuquerque were recorded for the first time for the country. A key for the Argentinean species is presented. Biological and forensic data of species are discussed.

KEY WORDS: Ophyra; Muscidae; Forensic entomology; Argentina.

Recibido: 3-III-2010;
Aceptado: 14-V-2010

INTRODUCTION

The genus Ophyra Robineau-Desvoidy (Diptera: Muscidae) comprises approximately 20 species of small flies distributed in warm climates worldwide. The species of this genus are frequently associated with decaying matter, especially on corpses (Centeno et al., 2002; Couri et al., 2009). Ophyra larvae act as predators on other dipteran larvae, and may be recovered on human cadavers during the late or active decay stage (Skidmore, 1985; Byrd & Castner, 2001). The species of Ophyra are frequently found in domestic situations that contribute to both medical and forensic importance (Oliva, 1997, 2001; Aballay et al., 2008; Carvalho & Mello-Patiu, 2008; Rosa et al., 2009). Synanthropic studies in the Neotropical Region have examined the relationship between the species of this genus and the different environments (Carvalho et al., 1984; Costa et al., 2000; Ribeiro et al., 2000; Figueroa-Roa & Linhares, 2004).
The phylogenetic placement of the genus Ophyra has been the subject of debate for a long time. Older hypotheses place this genus in the Phaoniinae (Malloch, 1923; Séguy, 1937). Currently, this idea was rejected in several works (Hennig, 1965; Pont, 1973; Skidmore, 1985; Pamplona & Couri, 1989; Carvalho, 2002). On one hand, some authors consider the genus Ophyra as a junior synonym of Hydrotaea within the Azeliinae (Vockeroth, 1996, Savage & Wheeler, 2004). On the other hand, some authors considered Ophyra and Hydrotaea as separate Azeliinae genera (Skidmore, 1985; Pamplona & Couri, 1989; Carvalho, 2002). More recently, phylogenetic analyses based on morphological (Savage & Wheeler, 2004) and molecular characters (Schuehli et al., 2004; Schuehli et al., 2007) suggest the placement of Ophyra within the Muscinae.
In the Neotropical region, the genus is represented by seven species (Pamplona & Couri, 1989; Carvalho et al., 2005). In Argentina, the genus Ophyra has been scarcely studied in previous taxonomic works with a single species, Ophyra aenescens (Wiedeman), and listed for the local fauna (Nihei & Dominguez, 2008; Battan Horenstein et al., 2010). This species was recorded by Bigot (1885) as Ophyra argentina and by Shannon & Del Ponte (1926, 1928) as Ophyra carbonaria, and subsequent studies have used these synonymic names (Oliva, 1997, 2001, 2007; Centeno et al., 2002). On the other hand, unidentified species of Ophyra were reported by Oliva (2001), Laos et al. (2004), Battan Horenstein et al. (2005) and Aballay et al. (2008) for several locations in Argentina.
In the present study, the knowledge of the necrophagous genus Ophyra in Argentina is revised and first country records for Ophyra albuquerquei Lopes,
Ophyra capensis (Wiedemann), Ophyra chalcogaster (Wiedemann) and Ophyra solitaria Albuquerque are presented. A key for the Argentinean species is provided, and the range of distribution of the species is extended.

MATERIALS AND METHODS

The present study was based principally on specimens collected during ecological studies with a sweep net on attracting baits (meat, rotten cow liver and chicken viscera); and from autopsy samples analyzed between the years 1995 and 2008.
Identification of specimens was ensured by the use of original descriptions and redescriptions of the species (see remarks). All the specimens studied belong to the following institutions: ANLIS "Dr. Carlos G Malbrán", Departamento Vectores (ANLIS); Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Laboratorio de Entomología forense (MACN-EF); Facultad de Agronomía-UBA, Cátedra de Biología Agrícola (CBA-Agro); Fundación Miguel Lillo (FML); Instituto Argentino de Investigaciones de las Zonas Áridas (IADIZA). The specimens collected by the authors were deposited at ANLIS and MACN-EF. Acronyms used for collections are as follows: BMNH the Natural History Museum, London, United Kingdom; MNHN Muséum National d'Histoire Naturelle, Paris, France; MNRJ Universidade do Rio Janeiro, Museu Nacional, São Cristovão, Rio de Janeiro, Brazil; NMW Naturhistorisches Museum Wien, Wien, Austria; UMO  Oxford University Museum of Natural History, Oxford, United Kingdom; USNM National Museum of Natural History, Washington D.C., USA; ZMUC University of Copenhagen, Zoological  Museum, Copenhagen, Denmark.
The terminology used in key for the external morphology follows McAlpine (1981) and Huckett & Vockeroth (1987).

RESULTS

Ophyra Robineau-Desvoidy, 1830

Ophyra Robineau-Desvoidy, 1830: 516. Type-species, Ophyra nitida Robineau-Desvoidy (Rondani, 1866: 70, 84) [= Musca ignava Harris, 1780].

Ophyra aenescens (Wiedemann, 1830)

Anthomyia aenescens Wiedemann, 1830: 435. Type-locality: "New-Orleans", USA. Lectotype male (by designation of Pont, 1997: 88) (NMW), paralectotype (NMW).
Ophyra virescens Macquart, 1843: 321. Type-locality: "Guaratuba, Brésil". Syntype female (MNHN). Synonymy with aenescens by Pont, 1972: 13.
Anthomyia setia Walker, 1849: 956. Type-locality: "Galapagos". Holotype female (BMNH). Synonymy with aenescens by Aldrich, 1928: 4, 5 and confirmed by Curran, 1932: 360.
Ophyra argentina Bigot, 1885: 302. Type-locality:"Buenos-Aires". Holotype female (UMO). Synonymy with aenescens by Stein, 1907: 212.
Ophyra carbonaria Shannon & Del Ponte, 1926: 576. Type-locality: Argentina: Tucumán & Buenos Aires Syntypes male/female, (USNM and ANLIS). Synonymy with aenescens by Aldrich, 1928: 4 and by Shannon & Del Ponte, 1928: 142.

Type material. Syntypes of Ophyra carbonaria. Buenos Aires: San Isidro, 25-VIII-1926, Shannon leg., 2 males, 5 females (ANLIS); Tucuman: Tucumán, 21-VI-1926, Shannon leg., 1 male (ANLIS).
Other material examined. Buenos Aires: Balcarce, II-1947, Wappers leg., 1 female (ANLIS); Buenos Aires, 20-IV-1915, Bruch leg., 1 male (ANLIS); Buenos Aires, # 271, 7-X-2003, Oliva leg., 1 female (MACN-EF); Buenos Aires Emerg. 111, 5-I-2000, Oliva leg., 6 females (MACN-EF); Buenos Aires, 3-VI-1907, Oliva leg., 1 females (MACN-EF); Burzaco, 13-II-2007, Mulieri leg., 3 males, 13 females (ANLIS); Carlos Casares, 14-II-2006, Medan, Devoto & Fernandez leg., 1 male (CBA-Agro); Ciudad Universitaria, Capital Federal, 7-XII-2007, Patitucci leg., 8 females (ANLIS); Instituto Bacteriológico "Dr. Carlos G. Malbrán", Capital Federal, 9-VII-1938, 1 female (ANLIS); Laguna La Tablilla, Chascomús, 01-XII-2007, Torretta leg., 2 males, 1 female (CBA-Agro); Magdalena, 30-XI-2000, Torretta leg., 1 female (CBA-Agro); Mar del Plata, XI-2007, Patitucci leg., 1 male, 1 female (ANLIS); Ministro Rivadavia, 17-I-2007, Mulieri leg., 11 males, 53 females (ANLIS); Quequen, I-2010, Patitucci leg., 2 females (ANLIS); Tandil, 21-XI-2003, cerdo 52 d., Oliva leg., 1 male, 1 female (MACN-EF); Villa Elisa, La Plata, II-1982 Mariluis leg., 2 males, 2 females (ANLIS). Córdoba: Capilla del Monte, i-2001, Mariluis leg., 1 female (ANLIS). Corrientes: Ituzaingo, xii-1976, Mariluis leg., 2 males, 8 females (ANLIS). Mendoza: capital, IADIZA, 31-vii-2007, Aballay leg., 2 males (IADIZA). Misiones: Iguazú, 23-III-1987, Mariluis leg., 5 males, 3 females (ANLIS); Puerto Esperanza, X-1978, Mariluis leg., 1 male, 3 females (ANLIS). Santa Fe: Christophersen, General López, 25-IV-2009, Patitucci leg., 3 males, 14 females (ANLIS). Salta: La Caldera, XII-1986, Mariluis leg., 5 males, 9 females (ANLIS); El Maray, Depto Chicoana, XII-1986, Mariluis leg., 1 female (ANLIS). San Juan: Jachal, La Legua, 1164 m.a.s.l., 06-IV-2005, Aballay leg., 2 males (IADIZA); same data except., 27-III-2005, 1 female (IADIZA); Rivadavia, 674 m.a.s.l., 11-I-2006, Aballay leg., 1 male, 1 female (IADIZA); same data except., 10-I-2006, 2 females (IADIZA); same data except., 12-I-2006, 1 male (IADIZA); Rivadavia, Barrio Aramburu, 08-X-2005, 3 females (IADIZA); same data except., 666 m.a.s.l., 07-VI-2005, 1 female (IADIZA); Valle Fértil, 560 m.a.s.l., 30-VI-2006, Aballay leg., 2 males, 5 females (IADIZA)

Distribution. Buenos Aires, Córdoba, Corrientes (new record), Mendoza (new record), Misiones (new record), Santa Fe (new record), Salta (new record), San Juan (new record), Tucumán (Fig 1).

Figs. 1-4. Geographic distribution of Ophyra species in Argentina. 1, Ophyra aenescens (black spots = new record; black star = bibliographic record); 2, O. albuquerquei (black spots = new record); 3, O. capensis (black spots = new record); 4, O. chalcogaster (black spots = new record) and O. solitaria. (white triangle = new record).

Remarks. The specimens examined agree with the description given by Pamplona & Couri (1989). In addition, we compare the specimens captured with the type material of O. carbonaria Shannon & Del Ponte. Ophyra aenescens is a widespread synanthropic species, originally from the Neotropical Region (Hogsette & Washington, 1995). This species is associated with urban environments and is more abundant during the summer months (Linhares, 1981; Ribeiro et al., 2000). This species has been recorded breeding in numerous media as crab, liver cow, fish, shrimp and mouse carcass (d'Almeida, 1988; d'Almeida, 1994; Moura, 2004). Larvae are saprophagous during the first instar, but second- and third-instar larvae are predators on Musca domestica L. (d'Almeida et al., 1999). In forensic studies, the larvae are often found in body exudates that have soaked into soil beneath remains, and commonly associated with exposed gut contents (Byrd & Castner, 2001).

Ophyra albuquerquei Lopes, 1985

Ophyra albuquerquei Lopes, 1985: 117 (figs. 1-8). Type-locality: Brazil, Rio de Janeiro, Le Vallon, Alto da Mosela Holotype male (MNRJ), paratypes (MNRJ).

Material examined. Buenos Aires: Burzaco, 15-XII-2006, Mulieri leg., 1 female (ANLIS); Capital Federal, XII-2007, Patitucci leg., 1 female (ANLIS); Ministro Rivadavia, 15-I-2007, Mulieri leg., 15 females (ANLIS). Córdoba: Capilla del Monte, I-2001, Mariluis leg., 2 females (ANLIS). Misiones: Iguazú, 23-III-1987, Mariluis leg., 2 males, 4 females (ANLIS); Puerto Esperanza, X-1978, Mariluis leg., 2 males, 7 females (ANLIS). Salta: La Caldera, XII-1986, Mariluis leg., 5 males, 9 females (ANLIS); El Maray, Depto Chicoana, XII-1986, Mariluis leg., 1 male, 5 females (ANLIS). Tucumán: Padre Monti, Burruyacu, 7-II-1948, Golbach leg., 2 males (FML), Quebrada La Toma, 21-XII-1950, Golbach leg., 1 male (FML); San Pedro de Colalao, 1190 m., III-1979, Mariluis leg., 1 female (ANLIS).

Distribution. Buenos Aires (new record), Córdoba (new record), Misiones (new record), Salta (new record). Tucumán (new record) (Fig 2).

Remarks. Diagnosis and illustrations of the male and female genitalia can be found in Lopes (1985) and Pamplona & Couri (1989). Ophyra albuquerquei is a Neotropical species, previously known from Brazil (Carvalho et al., 2005). This species was associated to rural or wild environments (Costa et al., 2000). The seasonal fluctuation of this species was similar to O. aenescens in Brazil (Riveiro et al., 2000). The biological information of this species is poorly known, with studies referred to breed, longevity and oviposition (Krüger et al., 2003, 2004).

Ophyra capensis (Wiedemann, 1818)

Anthomyia capensis Wiedemann, 1818: 46. Type-locality: "Vorgebirge der guten Hoffnung", South Africa. Lectotype male (NMW) (by designation of Pont, 1997: 91).

Material examined. Buenos Aires: Capital Federal, Cementerio de la Chacarita, 8-VIII-1998, Oliva leg., 2 males (ANLIS); Capital Federal, 5-I-2000, Oliva leg., Emerged from beef bait rearing nº 111, 2 males, 5 females (ANLIS).

Distribution. Buenos Aires (new record) (Fig 3).

Remarks. The specimens examined agree with the description given by Pamplona & Couri (1989). Ophyra capensis is an exotic species from the Old World (Skidmore, 1985). This species has been reared from human faeces, carrion, nets of various mammals and birds (Couri et al., 2009). Ophyra capensis was found on human bodies kept indoors for several months, where blowflies (Diptera, Calliphoridae) do not have access to them (Bourel et al., 2004). Recently, paleopathologic studies found puparia and adult fragments of this species into a mummified body (Couri et al., 2008).

Ophyra chalcogaster (Wiedemann, 1824)

Anthomyia chalcogaster Wiedemann, 1824: 52. Type-locality: "Iava", Indonesia. Lectotype male (ZMUC) (by designation of Pont, 1973: 242).

Material examined. Buenos Aires: Burzaco, 15-XII-2006, Mulieri leg., 3 females (ANLIS); Buenos Aires, 20-X-1906, A Zotta leg., 1 male (MACN-EF); Buenos Aires 17-IV-2002, Oliva leg., # 219, 1 female (MACN-EF); Capital Federal, X-2007, Patitucci leg., 4 males, 3 females (ANLIS). Misiones: Iguazú, X-1988, Mariluis leg., 4 females (ANLIS).

Distribution. Buenos Aires (new record), Misiones (new record) (Fig 4).

Remarks.The specimens examined agree with the description given by Pamplona & Couri (1989). Ophyra chalcogaster is a cosmopolitan species associated to urban or suburban environments (Linhares, 1981; Costa et al., 2000). Different kinds of bait were used to collect and breed this species (human and dog faeces, fish, chicken viscera, mouse carcass) (d'Almeida et al., 1989; Mendes & Linhares, 1993; Moura, 2004). This species is less abundant than O. aenescens and O. albuquerquei in Brazil (Riveiro et al., 2000).

Ophyra solitaria Albuquerque, 1958

Ophyra solitaria Albuquerque, 1958: 7 (figs. 12-16). Type-locality: Brazil, Goiás, Anápolis. Holotype female (MNRJ).

Material examined. Misiones: Iguazú, 23-III-1987, Mariluis leg., 6 females (ANLIS); Puerto Esperenza, X-1978, Mariluis leg., 1 female (ANLIS).

Distribution. Misiones (new record) (Fig 4).

Remarks. Identification of Ophyra solitaria was based on Albuquerque (1958) for the females, and Lopes (1978) for males. In addition, Pamplona & Couri (1989) provided diagnosis of both sexes. This is a Neotropical species, previously known from Brazil (Carvalho et al., 2005). This species was associated to rural or wild environments (Costa et al., 2000). Ophyra solitaria was collected and reared from human faeces, crab, fish, shrimp and mouse carcass (d'Almeida, 1988; Leandro & d'Almeida, 2005). This species is less abundant than O. aenescens and O. albuquerquei in Brazil (Riveiro et al., 2000).

Key to the species of Ophyra in Argentina (modified from Carvalho, 2002)

1. Palpus orange-yellow. Female: ocellar triangle long and wide with a rounded apex reaching the lunule (Fig. 5). Male: hind trochanter on ventral surface with a tuft of fine setae ................................................................O. aenescens (Wiedemann)
- Palpus brown or black. Female: ocellar triangle shape not as above (Figs. 6-9). Male: hind trochanter on ventral surface without a tuft of fine setae ................................2

Figs. 5-9. Ocellar triangle, frontal view, female. 5, Ophyra aenescens; 6, O. albuquerquei; 7, O. capensis; 8, O. chalcogaster; 9, O. solitaria. Scale bars = 1 mm.

2. Lower calypter dark brown with dark brown margin. Female: ocellar triangle long and sharpened with a rounded apex reaching the lunule (Fig. 6). .............................................................................O. albuquerquei Lopes
-  Lower calypter yellowish or white with yellow or white margins. Female: ocellar triangle shape not as above (Fis. 7-9)..............................................................3

3. Arista yellow at the basal middle and black at the distal middle. Female: ocellar triangle short, with a triangular shape not reaching lunule (Fig. 7). Male: fore tarsi with tarsomeres yellow on ventral surface...........................O. chalcogaster (Wiedemann)
- Arista brown. Female: ocellar triangle shape not as above (Fis. 8-9). Male: fore tarsi with tarsomeres brown on ventral surface.......................................................4

4. Wing hialine. Female: ocellar triangle broad not reaching lunule (Fig. 8). Male: hind tibia on anteroventral, ventral and posteroventral with a series of long seta at apical two-thirds.....................................................................O. capensis (Wiedemann)
- Wing brown at the anterior middle. Female: ocellar triangle long with a rounded apex not reaching lunule (Fig. 9). Male: hind tibia with 2 setae at the middle on anteroventral, 1 seta on anterodorsal and 1 long seta posteroventral.................O. solitaria Albuquerque

DISCUSSION

The species of Ophyra, based on the literature and the examined material, were associated with different kinds of baits (Carvalho et al., 1984; d'Almeida et al., 1989; Costa et al., 2000). Specimens collected in Buenos Aires, as part of a comprehensive work on Muscidae assemblages along an urban-rural gradient, were mainly collected on meat, cow liver and chicken viscera. Phenology of the species seems to be restricted to the warmer period. The species were captured during November and December (late spring) in agreement with previous works (Costa et al.,2000; Carvalho et al., 1984). In addition, material of Ophyra spp. was found in autopsy samples along the whole year, but it was more abundant in November and December (late spring). Only 14 samples, out of 407 taken between 1995 and 2005, contained Ophyra spp. (Oliva, 2007; sub Hydrotaea argentina). In the period 1995-2008 there were 32 cases with Ophyra spp., of which 3 were exhumations. Adipocere was present in four cases (two exhumations, one corpse wrapped in plastic bags, one dismembered corpse inside a bag).
The distribution of these species was significantly extended. We registered the southernmost point of the South American distribution for O. aenescens in Quequen, Buenos Aires (38º 34' 12.68'' S, 58º 40' 24.52'' W). Also, our current findings expand the geographical distribution of the widespread species, O. capensis and O. chalcogater, reaching the area of Buenos Aires city. On the other hand, the Neotropical species O. albuquerquei, which had only been recorded from Brazil, was registered in Buenos Aires, Córdoba, Salta and Tucumán provinces expanding significantly its geographical distribution. Finally, a few specimens of O. solitaria were captured in the province of Misiones.
Muscid flies are of great forensic importance due to their wide distribution, ubiquitous nature, and close association with man (Byrd & Castner, 2001). Ophyra species have a specific forensic significance, appearing usually during the period of ammoniacal fermentation (Couri et al., 2008). The fact that species of Ophyra attack the corpses in a later succession wave suggests that probably the corpse has not been exposed to open air for some time (Couri et al., 2009). This theory was supported by Smith (1986), who makes clear that in the cold-temperate climate of the British Isles, Ophyra spp. will be found in warm, dry ambient, indoors or in coffins. However, in subtropical and tropical climates one may expect to find Ophyra spp. in large numbers outdoors.
The species of Ophyra occur in different environments along an urban-rural gradient (Linhares, 1981; Costa et al., 2000). The Neotropical species of this genus (O. albuquerquei and O. solitaria) were collected in rural and wild environments. On the contrary, the cosmopolitan species (O. aenescens, O. capensis, and O. chalcogaster) were associated to urban areas (Linhares, 1981; Costa et al., 2000). In Buenos Aires, Ophyra spp. appear in large numbers on disinterred corpses and indoors in cases of undetected natural deaths (Oliva, 2007). In contrast, Oliva (2001) found Ophyra sp. in samples obtained on an outdoors corpse in December (early summer), in Chajarí, Entre Ríos province. These contrasting results may be explained by the presence of different species in different environmental conditions.
The knowledge concerning with the relative abundance and the seasonal fluctuation of species of Ophyra in different environments could provide new tools for forensic entomology.

ACKNOWLEDGEMENTS

Thanks to the curators Guillermo L. Claps (FML) and Juan P. Torretta (FAUBA) for the loan material. We also thank Fernando H. Aballay for providing San Juan and Mendoza specimens. We appreciate the comments made by Gustavo E. Flores.

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