Chrysomya Robineau­Desvoidy, 1830: 44

Kurahashi, Hiromu & Kirk-Spriggs, Ashley H., 2006, The Calliphoridae of Namibia (Diptera: Oestroidea), Zootaxa 1322 (1), pp. 1-131 : 74-79

Genus: Chrysomya Robineau­Desvoidy, 1830: 44 .

TYPE SPECIES: Chrysomya marginalis (Wiedemann, 1830: 395) (Musca) , by designation of Coquillett (1910: 523).

NOTES: Chrysomya is the only genus of the Chrysomyinae represented in the mainland Afrotropical Region. The bionomics of several species, important as myiasis­producing ‘blowflies’, have been studied (vide Ferrar 1987: 88; Zumpt 1956: 180, 1965: 89–102, for review). Zumpt (1965: 89–90) keys the adults and larvae of species important in myiasis in the Old World. Chrysomya albiceps , C. chloropyga and C. marginalis (as C. regalis ) (among others) are known to cause faculitive traumatic myiasis in Africa (Zumpt 1956: 180). Diagnostic features to separate the 3 rd instar larvae of C. albiceps , C. chloropyga and C. putoria are provided by Erzinçlioğlu (1987).

Chrysomya albiceps (Wiedemann, 1819: 38) (Musca) .

‘Western banded blowfly’.

Fig. 75 .

TYPE LOCALITY: South Africa .

DISTRIBUTION: Widespread in Afrotropical Region, North Africa eastwards to Pakistan in Oriental Region. Neotropical Region: Brazil and Guatemala (introduced and established). Aldabra Is., Botswana, Cape Verde Is., Ethiopia, Kenya, Madagascar, Mauritius, Namibia, Réunion Is., Rodriquez Is., Seychelles Is., Socotra Is., South Africa (Cape, Natal, Transvaal),? St. Helena Is., Sudan, Yemen and Zimbabwe.

PREVIOUS RECORDS: Swakopmund [22°68'S, 14°53'E], 18.i.1970; Ongeama [22°22'S, 17°18'E], 25.i.1970 (Lindner 1976: 77).

MATERIAL: 1♂, Barby 26, 17–18.i.1972, [SMStaff], H6171 ; 2♂, same except: 2–7.x.1972, H9202; 2♀, Blinkoog , 14–17.x.1971, [SMStaff] ( MT), H4510 ; 1♂, Takuasa , 14–19.viii.1971, [SMStaff], H3027 ; 8♀, Otjikoko­Süd 61, 10–13.ii.1972, [SMStaff], H6453 ; 1♂, 2♀, Arandis , 6 km N, 3–31.vii.1984, Irish (1) & Liessner, H58519 ; 3♂, Windhoek (2), 1–4.xi.1973, [SMStaff], H14940 ; 1♂, Gorrasis 99, 12–15.ii.1973, [SMStaff], H11592 ; 1♀, Arnhem 222(1), 23–27.x.1972, [SMStaff], H9671 ; 1♂, Noachabeb , 7–12.i.1972, [SMStaff], H5867 ; 1♀, Okahandja 18.xi.[19]71, Gaerdes , H25144 ; 1♂, same except: 2.ii.[19]57, H25143 ; 2♂, 2♀, Hippo Pool , 15.i.1993, Marais ( LT) ; 3♂, 11♀, Viljoenskroon 507, 24–25.xii.1995, Marais ; 1♂, Rundu (Kavango Lodge ), 28.iii.2003, Kirk­Spriggs (1), decomposing fungi ; 1♂, Rundu , 20 km E, 17–18.x.1999, Kirk­Spriggs (1), Pape & Hauwanga ( YP) dry woodland ; 1♂, Klinghardt Mountains at(1): 27.viii–3.ix.1998, Kirk­Spriggs (1) & Marais ( YP) ; 3♂, Leeukop 664, 26–30.x.2001, Kirk­Spriggs (1) & Marais ( MT) riverbed ; 5♂, 1♀, Lekkerwater 406, 13.xii.1998, Mann & Kirk­Spriggs (1), beneath decomposing Small­spotted Genet ( Genetta genetta L.) (Vivarridae) on road [all with associated puparia] ; 5♂, Renostervlei , 2 km E, 26–27.xii.1999, Marais, Mann & Newman, MMN27, faeces ; 1♀, Tsisab, N end, 21.i–27.ii.1995, Marais ( PT) ; 3♂, 4♀, Wasserfallfläche (1), 20–22.iii.2001, Kirk­Spriggs (1) & Marais, decomposing Procavia capensis [all with associated puparia] ; 7♂, 25♀, Salambala campsite, 28.xii.2002, Kirk­Spriggs (2) ( HT) faeces ; 1♂, Upper Hungorob ravine at: 18.iv.2000, Meakin / Raleigh Int. ( LT) Bberg UV 24 ; 1♂, 1♀, Falls rock ravine at: 2.v.2000, Meakin / Raleigh Int ( YP) Bberg pan 52 ; 5♂, 1♀, Windhoek, S. Industrial Area, 1.ii.2004, Robberts, beneath human cadaver, ex National Forensic Science Institute, Ministry of Home Affairs , Windhoek .

NOTES: A very common fly, widespread over the whole of Africa. Cuthbertson (1933: 94) notes that in Zimbabwe the fly is regarded as beneficial, as its larvae prey upon those of other ‘blowfly’ larvae, including C. chloropyga . This is disputed by Zumpt (1965: 92), however, who notes that living tissues may also be consumed by larvae. Cuthbertson (1933: 94) records larvae from ‘under the tail’ of a heifer suffering from ‘scour’ in Zimbabwe. He describes the life history and notes that puparia are parasitised by Nasonia Ashmead (as Mormoniella ) ( Hymenoptera : Pteromalidae ). Zumpt (1965: 92) notes that the species is a typical secondary myiasis fly of sheep; the larvae not being able to ‘strike’ sheep, without the prior assistance of primary maggots (mainly Lucilia cuprina in southern Africa). In Namibia collected in pitfall, Malaise and yellow pan traps. In hanging traps baited with faeces and at faeces, at UV­light, and at decaying mushrooms. It has also been reared from larvae and puparia collected beneath a dead Smallspotted Genet Genetta maculata (Gray) (Viverridae) on a road and from beneath a dead Rock hyrax Procavia capensis (Pallas) (Procaviidae) . Occurring at high elevations on the Brandberg (1170 m, 1920 m). Apparently widely distributed throughout Namibia; occurring in all Namibian biomes ( Fig. 75 ). Recorded throughout the year, except in June and September; most abundantly in December (vide Table 2). The life history has been studied by Smit (1931) and Cuthbertson (1933: 94), who provide a habitus illustration of the 3 rd instar larva in lateral aspect (fig. 1, A), the anterior end in lateral aspect (fig. 2, A), and the posterior spiracles from behind (fig. 2, B). The larva is characteristic in having concentric rows of fleshy projections on all body compartments. For a summary of life history, immature stages and pathogenesis vide Zumpt (1965: 90–92), and for details of the 3 rd instar larva Erzinçlioğlu (1987).

FORENSIC SIGNIFICANCE: This is one of the three most important primary biodegraders of decomposing corpses and is the most prevalent fly during the summer months. The characteristic larvae, known as ‘hairy maggots’, are extremely aggressive and readily attack and consume other larvae, ensuring that all competition is eventually eliminated, especially on a dwindling resource. It tends to out­compete all other species because of it cannibalistic life strategy. Eggs are normally laid shortly after other species have oviposited, and this ensures an additional option for the larvae should the primary resource become depleted (M. Mansell pers. comm.). The forensic significance of the species is discussed in detail by Grassberger et al. (2003). The species has been identified as breeding in the cadaver of a murder victim in central Windhoek (vide ‘Material’ section above). For general information vide Greenberg and Kunich (2002).

Chrysomya chloropyga (Wiedemann, 1818: 44) (Musca) .

‘Copper­tailed blowfly’, ‘Winter blowfly’.

Fig. 76 .

TYPE LOCALITY: South Africa .

DISTRIBUTION: Widespread in Afrotropical Region: Cameroon, Democratic Republic of Congo, Ethiopia, Kenya, Lesotho, Namibia, St. Helena Is., South Africa, Tanzania and Zimbabwe (Rognes & Paterson 2005: 52–53).

MATERIAL: 1♀, Upper Ostrich Gorge, 8.v–5.vi.1984, Irish(1) & Liessner, H59299.

NOTES: For the most recent treatment of the taxonomy of this species and its separation from its sister species C. putoria vide Rognes and Paterson (2005) . A common sun­loving fly in southern Africa, attracted from great distances by carrion. Larvae feed on meat juice and the liquid meat ‘broth’ produced by their own feeding action. Any sweet liquid, such as sugar water, is also taken, but a protein meal is required by females for egg maturation (Zumpt 1965: 93–95). Cuthbertson (1933: 99) notes that the species is a serious pest of wooled sheep in Zimbabwe, while Zumpt (1965: 95) records it as a facultative cause of traumatic myiasis in sheep in South Africa, second in importance after Lucilia cuprina . He provides notes on the life history, immature stages and pathogenesis. Cuthbertson (1933: 99–100) notes a record of Bembix massaica Cameron ( Hymenoptera : Sphecidae ), carrying­off this species to its nest, and further records Nasonia Ashmead (as Mormoniella ) ( Hymenoptera : Pteromalidae ), as occasional parasites of the puparium. He provides illustrations of the anterior end of the 3 rd instar larva (fig. 6, C); and the posterior spiracles from behind (fig. 6, D). In Namibia, the species has only been collected in one locality, probably in a pitfall trap, in May. The single record is from the desert biome ( Fig. 76 ). Notes on the 3 rd instar larva are provided by Erzinçlioğlu (1987).

FORENSIC SIGNIFICANCE: This species, known as the ‘Copper­tailed’ or ‘Winter blowfly’, is the most prevalent forensic indicator and has been encountered on about 60% of cases investigated in South Africa over the past decade. Although present throughout the year, its phenology tends towards the colder months, when competition from other species is reduced. They are also capable of producing large populations on a decomposing corpse and compete directly with C. albiceps (M. Mansell pers. comm.). For general information vide Greenberg and Kunich (2002).

Chrysomya ? laxifrons Villeneuve, 1814 a: 178.

Fig. 77 .

TYPE LOCALITIES:? Uganda and Zaïre [= Democratic Republic of Congo] .

DISTRIBUTION: Widespread mainland Afrotropical Region: Cameroon, Democratic Republic of Congo, Equatorial Guinea, Namibia *, Sierra Leone, South Africa (Transvaal), Sudan, Tanzania, Togo, Uganda and Zimbabwe.

MATERIAL: 1♀, Kameseb, N of, 12.x–11.xi.1986, Griffin(1) ( PT).

NOTES: It appears that the life history and immature stages of this species are not known (vide Zumpt 1956: 184) and Zumpt (1965) does not record it as a myiasis­producing fly. In Namibia collected in a pitfall trap in December. The single record of this unconfirmed species is from the Etosha pan in the nama­karoo biome ( Fig. 77 ).

Chrysomya marginalis (Wiedemann, 1830: 395) (Musca) [specific name retained, I.C.Z.N. 1988: 236 (Opinion 1507)].

= regalis Robineau­Desvoidy, 1830: 449 .

‘Large blue blowfly’.

Fig. 78 .

TYPE LOCALITY: South Africa .

DISTRIBUTION: Widespread Afrotropical Region; Oriental Region: Pakistan. Benin, Kenya, Madagascar, Namibia, Nigeria, Rwanda, Senegal, Socotra Is., Somalia, South Africa (Cape, Natal, Transvaal), Tanzania, Yemen and Zimbabwe.

PREVIOUS RECORDS: Ongeama [22°22'S, 17°18'E], 25.i.1970; Krumhuk [22°44'S, 17°04'E], 28.i.1970 (Lindner 1976: 77); Anabib (Orupembe), 100 miles W Ohopoho [19°08'S, 13°44'E], 12–13.vi.1951 (Zumpt 1959b: 430).

KNOWN RECORDS: (all as C. regalis ) Aus [26°67'S, 16°27'E], i.1972, Eastwood; Okahandja [21°98'S, 16°92'E], Eastwood; Otjimbingwe [22°35'S, 16°13'E], 8.ix.1970; Outjo [20°12'S, 16°15'E], ix.1952; Tsumeb [19°23'S, 17°72'E], 17.ii.1970 (NMSA).

MATERIAL: (all labelled as C. regalis ) 4♂, 2♀, Barby 26, 17–18.i.1972, [SMStaff], H6171 ; 1♂, Arnhem 222(1), 23–28.x.1972, [SMStaff], H9670 ; 1♀, Claratal 18, 27.i.1971, [SMStaff], H1566 ; 3♂, 1♀, Wasservallei 382, 21–23.xii.1973, [SMStaff], H16263 ; 3♂, 2♀, Takuasa , 14–19.viii.1971, [SMStaff], H3028 ; 1♀, Windhoek (1), 20–27.xi.1973, [SMStaff], H15782 ; 1♀, same except: 13–15.xi.1973, H15056 ; 1♀, same except: 20–22.xi.1973, H15182 ; 2♀, same except: 13–30.iv.1974, H18611 ; 1♂, same except: v.1972, H8579; 1♂, same except: 1–4.xi.1973, H14939 ; 1♂, Windhoek (1), iii.1985, [SMStaff], H63969 ; 1♂, 3♀, Ruacana Falls , 26–27.viii.1973, [SMStaff], H13988 ; 2♂, Churutabis 108, 4–12.x.1974, [SMStaff], H20954 ; 1♂, Arnhem 222(1), 23–27.x.1972, [SMStaff], H9670 ; 1♂, same except: 23–28.x.1972, H9670; 4♀, Otjikoko­Süd 61, 10–13.ii.1972, [SMStaff], H6452 ; 1♂, Otjitembi 25, 14–15.ii.1972, [SMStaff], H6633 ; 1♀, Orupembe , 21 km SW, 16.viii.1973, [SMStaff], H13465 ; 1♀, Rundu , 80 km W, 2.iii.1973, [SMStaff], H11957 ; 1♀, Otjinhungwa , 17–22.xi.1970, [SMStaff], H10508 ; 1♂, 1♀, Barby 26, 2–7.x.1972, [SMStaff], H9201 ; 1♀, Riverside 135, 23–16.x.1971, [SMStaff], H4857 ; 1♂, Blinkoog , 14–17.x.1971, [SMStaff], collected near open water, H4509 ; 1♀, Otjiseva 45, 12.iii.1971, [SMStaff], H1903 ; 1♀, Swartbaas West 276, 19–22.iv.1972, [SMStaff], H7812 ; 1♀, Welverdiend 328, 8–13.x.1972, [SMStaff], H9519 ; 1♂, Waterberg Plato Park , 29.iv.1985, Rust (1) & Meyer, H64317 ; 1♀, Dorsland , 8.x.1986, Marais & Griffin (1) ; 1♀, Tsumkwe , 8–17.xi.1985, Rust (2), H64739 ; 2♂, Ubussis 3, 11–13.i.1985, Irish (2), H63121 ; 5♀, Okahandja, 2.ii.[19]57, Gaerdes , H25137 ; 1♀, same except: 27.xii.[19]51, H25134 ; 1♂, same except: 16.ix.[19]71, H25140 ; 1♀, same except: 4.i.[19]69, H25139 ; 1♀, same except: 17.xii.[19]68, H25138 ; 1♀, same except: 16.iii.[19]51, H25136 ; 1♀, Karasburg , 5 km NW, 15.iii.1988, Marais & Irish (1) ; 1♂, Orohona , 5–11.xi.1989, Roberts ; 1♀, Atlanta 618, 21.ii–21.iv.1995, Marais ( PT) ; 1♂, Ohamwaala , 21.i.1993, Marais ; 3♂, 1♀, Hippo Pool , 15.i.1993, Marais ( LT) ; 2♀, Dakota 424(1), 6.iii.1993, Pusch ; 1♂, 1♀, Varianto 771/2, 30.iii–1.iv.2003, Kirk­Spriggs (1) ( HT) fungi ; 1♀, same except: Kirk­ Spriggs (1) & Mey ( LT) ; 1♀, same except: 20–22.iii.2003, Kirk­Spriggs (1) ( MT) ; 1♂, Onangombe , 19.i.1993, Marais ; 1♂, 3♀, Viljoenskroon 507, 24–25.xii.1995, Marais ; 2♀, De Hoek 878, 3–6.ii.2001, Kirk­Spriggs(1) & Marais, decomposing fish bait; 1♂, dolomitic hill at: 25.ix.1997, Kirk­Spriggs(1) & Marais, on the wing; 1♂, Naukluft spring, 28–30.xi.1997, Kirk­ Spriggs (1) & Marais ( MT) ; 1♀, Ae­ams / Hoanib confluence, 28.xii.1999, Marais, Mann & Newman, MMN33, elephant dung ; 3♂, 2♀, Epupa , 35 km E: Kunene River, 9–11.x.1999, Kirk­Spriggs (1), Pape & Hauwanga ( MT) shaded woodland ; 1♂, Xawasha pan, 3 km E, 25.xii.1998, Kirk­Spriggs (1), Loxidonta africana dung ; 6♂, 5♀, Nama , 21–22.xii.1998, Marais, Kirk­Spriggs (1) & Mann ( HT) fruit ; 1♂, 1♀, same except: ( MT) ; 1♀, Hungorob ravine at(2): 17.iii.2001, Kirk­Spriggs (1) ( LT) ; 1♀, Messum Valley , 3.iv.1999, van Noort & Compton ( LT), sparsely vegetated river valley, Bushy Karoo­Namib shrubland, NA99–L01 ; 1♂, Okavango River at: 18–19.x.1999, Kirk­Spriggs (1), Pape & Hauwanga ( MT) ; 1♀, Viljoenskroon 507, 7–9.ii.1998, Kirk­Spriggs (1) & Marais ( MT) ; 2♀, Leeukop 664, 26–30.x.2001, Kirk­ Spriggs (1) & Marais ( MT) riverbed ; 3♂, Salambala campsite, 28–29.xii.2002, Kirk­Spriggs (2) ( HT) fish ; 7♂, 15♀, same except: 28.xii.2002, Kirk­Spriggs (2) ( HT) faeces ; 1♀, same except: 1–3.iii.2001, Kirk­Spriggs (1) ( HT) faeces ; 1♂, 2♀, same except: 3–4.iii.2001, Kirk­Spriggs (1) ( HT) fish ; 1♀, Salambala forest , 23–29.xii.2002, Kirk­Spriggs (2) ( MT) ; 2♂, Hungorob ravine at(1): 2.xi.1999, Meakin / Raleigh Int. ( LT) Bberg UV 2 ; 1♀, same except: 25.iv.2000, Meakin / Raleigh Int. ( LT) Bberg UV 31 ; 1♂, same except: 1.xi.1999, Meakin / Raleigh Int. ( LT) Bberg UV 1 ; 1♂, 1♀, Upper Hungorob ravine at: 28.iv.2000, Meakin / Raleigh Int. ( LT) Bberg UV 33 ; 1♂, Hungorob River at: 23.xi.1999, Meakin / Raleigh Int. ( PT) ; 3♂, 3♀, Naukluft campsite, 1–3.v.2005, Kirk­Spriggs (2) ( HT) faeces ; 3♂, 3♀, Windhoek, S. Industrial Area, 1.ii.2004, Robberts, beneath human cadaver, ex National Forensic Science Institute, Ministry of Home Affairs , Windhoek .

NOTES: A very distinctive fly that is common throughout the Afrotropical Region. Cuthbertson (1933: 101) observed adults feeding on wounds in cattle, but the species chiefly breeds in dead animals and adults are occasionally observed on flowers and at the honeydew of aphids and scale insects. Adults swarm on fresh cattle dung and faeces, and persistently feed on septic liquid exuding from sores and screw­worm infected wounds. In towns and cities they are a nui­ sance in butcher’s shops and abattoirs. He also notes that flies are attracted to sick animals (e.g. cattle) at the point of death and to carcasses. Zumpt (1956: 187, as C. regalis ) notes the species as causal agents of traumatic myiasis, but stated later (Zumpt 1965: 96) that it is almost certain that the fly does not attack sheep or other animals in southern Africa. In Namibia the species has been frequently hand­collected and caught on the wing; has been collected in pitfall and Malaise traps, and in hanging traps baited with rotting fungi and fish, faeces and fermenting fruit bait. It frequently comes to UV­light, and has been collected from fresh African elephant dung. Occurring at low and high elevations on the Brandberg (700 m, 1170 m). Apparently widely distributed throughout Namibia; occurring in all Namibian biomes, although rarer in the desert biome ( Fig. 78 ). Recorded throughout the year, except in June and July; most abundantly in December (vide Table 2). Cuthbertson (1933: 101) briefly describes the life history, and notes that migrating larvae fall prey to hymenopteran parasites, and bred Brachymeria varipes (Walker) (as Chalcis (B.)), Haltichella Spinola sp. and Hockeria Walker sp. (all Hymenoptera : Chalcididae ) from larvae, and noted that Nasonia Ashmead (as Mormoniella ) ( Hymenoptera : Pteromalidae ) occasionally attack puparia. The immature stages are incompletely known, but the larvae can be easily distinguished from those of other species of the genus by use of the key in Zumpt (1965: 89–90, as C. regalis ). The mature larvae are described by Prins (1979).

FORENSIC SIGNIFICANCE: This species is another primary biodegrader that is occasionally encountered in forensic investigations. It is mainly attracted to large mammal carcases, or bloated human corpses where there is a dependable resource. Eggs are laid in large clusters, which result in massive maggot infestations. It is probably about the fourth most important indicator species at crime scenes (M. Mansell pers. comm.). The species has been identified as breeding in the cadaver of a murder victim in central Windhoek (vide ‘Material’ section above). For general information vide Greenberg and Kunich (2002).

Chrysomya putoria (Wiedemann, 1830: 403) (Musca) .

Fig. 79 .

TYPE LOCALITY: Aus Sierra Leone .

DISTRIBUTION: Neotropical Region: Brazil, Colombia, Panama and Peru. Widespread Afrotropical Region: Botswana, Cameroon, Democratic Republic of Congo, Gambia, Ghana, Madagascar, Mauritius, Mozambique, Namibia *, Réunion Is., Senegal, Seychelles Is., Sierra Leone, South Africa, Swaziland, Tanzania, Uganda and Zamibia (Rognes & Paterson 2005: 52–53).

MATERIAL: 1♀, Skorpion area , 9–12.viii.1997, Marais & Kirk­Spriggs (1), on fresh Gemsbok dung ; 1♀, Rundu (Kavango Lodge ), 27–29.iii.2003, Kirk­Spriggs (1) & Mey ( LT) ; 1♂, Salambala campsite, 3–4.iii.2001, Kirk­Spriggs (1) ( HT) fish; 1♀, same except: 28–29.xii.2002, Kirk­ Spriggs (2) ( HT) fish; 3♂, 2♀, same except: 28.xii.2002, ( HT) faeces; 1♀, same except: 23–28.xii.2002, ( HT) mango; 1♀, Hippo Lodge (Zambezi River), 6–7.ii.2004, Kirk­Spriggs (1) ( HT) fruit.

NOTES: This has only recently been recognised as a distinct species from C. chloropyga (vide Rognes & Paterson 2005), based on characters of thoracic patterning, head setation and genital structure. The species appears to be widespread in the Afrotropical Region, and appears to share the same habits as C. chloropyga . Cuthbertson (1934: 38) notes that both sexes are attracted to the honeydew of aphids and scale insects (Homoptera: Coccidae ), on citrus and cotton and that males were observed on mango blossoms and flowers of Cymnospora sp. ( Celastraceae ) in Zimbabwe. Females are commonly found on fresh cattle dung, bird droppings, etc. In Namibia collected in Malaise traps and hanging traps baited with rotting fish, faeces, fermenting fruit bait and mangos, also at UV­light and alighting on fresh Gemsbok Oryx gazelle (L.) ( Bovidae ) dung. The few Namibian records are from the ‘mesic’ savanna biome in the north­east and the succulent karoo biome in the extreme south­west of the country ( Fig. 79 ). Recorded in February, March, August and December; most abundantly in December (vide Table 2). Additional notes on the 3 rd instar larva are provided by Erzinçlioğlu (1987).

SUBFAMILY: RHINIINAE

TRIBE: RHINIINI

Figures

FIGURES 75–80. Distributions of Namibian Calliphoridae species. 75. Chrysomya albiceps (Wiedemann). 76. C. chloropyga (Wiedemann). 77. C.?laxifrons Villeneuve. 78. C. marginalis (Wiedemann). 79. C. putoria (Wiedemann). 80. Rhinia apicalis (Wiedemann).