The opiine parasitoids of tephritids comprise several distinct lineages (Tobias 1977, Wharton 1997). It is likely that tephritid parasitism in the Opiinae represents multiple, independent evolutionary events. However, the alternative hypothesis that parasitism of tephritids is simply a plesiomorphic trait of the subfamily as a whole cannot be completely discounted.
The classification of the Opiinae used here is derived from that of Fischer (1972). Fischer (1972, 1977, 1987) treated the world species of Opiinae, and placed nearly all of the known species of tephritid parasitoids in two subgenera within Opius and in six other genera outside Opius. Recent works on the Opiinae by Tobias and Jakimavicius (1986), Wharton (1987, 1988), and van Achterberg and Maetô (1990), building on the work of Tobias (1977), Wharton and Marsh (1978), Wharton (1983), and Wharton and Gilstrap (1983), have led to substantial changes in the taxa to which these tephritid parasitoids were formerly assigned. Most notable among these changes are the restriction in the definition of Biosteres Förster, and elevation of Psyttalia Walker and Utetes Förster to generic rank. Those genera characterized by a short second submarginal cell (Diachasmimorpha, Doryctobracon, Fopius, and Rhynchosteres) have often had their species placed in Biosteres or elsewhere within Fischer’s subtribe Biosterina at one time or another.
Based on the classification used here, the parasitoids of fruit-infesting tephritids are hypothesized to fall into four major groups: 1) the Diachasmimorpha group of genera (Diachasmimorpha, Fopius , and the three species currently assigned to Diachasma), 2) the genus Psyttalia, 3) the genus Utetes + the Opius froggatti species group, and 4) the Opius bellus species group (treated as the subgenus Bellopius). Other opiines reared from fruit-infesting tephritids (e.g. Opius downesi Gahan and some of the species of Sternaulopius) may represent relatively isolated evolutionary forays into tephritid parasitism, but more host records are needed to confirm this. The genus Doryctobracon shares many features with the Diachasmimorpha group (Wharton 1997), but its relationship to other groups of tephritid parasitoids still needs to be explored further. Examples of additional publications on phylogeny of the Opiinae that include at least some exemplars from among the tephritid parasitoids are: Gimeno et al. (1997), Quicke et al. (1997), Wharton (2006), and Wharton et al. (2006).
More detailed information is provided under separate treatments for: Diachasma, Diachasmimorpha, Doryctobracon, Fopius, Opius, Psyttalia, Sternaulopius, Utetes and Bellopius).
APPENDICES
The following appendices treat all the names for Opiinae reared from fruit-infesting Tephritidae. They have been slightly updated from Wharton (1997). Appendix 1 is a list of the currently accepted species names for all described species with confirmed host records. Appendix 2 includes species group names (species, subspecies, and varieties) that are no longer in use, but which have been used for various tephritid parasitoids in the past. Appendix 3 gives the status of various generic names that have been applied to the species attacking Tephritidae.
Appendix 1
Valid species names of Opiinae associated with fruit, flower, and seed-infesting Tephritidae and their current generic placement. Parasitoids of leaf-mining tephritids are not included. See Appendix 3 for status of generic names used in association with tephritid parasitoids. Citations for original descriptions of these species can be found in Fischer (1971, 1972, 1977, 1984, 1986, 1987), Wharton and Gilstrap (1983), Wharton (1987,1997, 1999, 2009), Tobias (1998) van Achterberg (1999), Kimani-Njogu and Wharton (2002), Carmichael et al. (2005), and Fischer and Madl (2008).
acidoxanthicida (Fullaway, 1949) Psyttalia
aciurae (Fischer, 1964) Utetes
africanus (Szépligeti, 1910) Utetes
agreutretae (Wilkinson, 1927) Psyttalia
aino (Watanabe, 1938) Diachasmimorpha
albimanus (Szépligeti, 1905) Utetes
albobalteata (Cameron, 1912) Diachasmimorpha
alloeum (Muesebeck, 1956) Diachasma
alternatae (Tobias, 1977) Fopius
anastrephae (Viereck, 1913) Utetes
anastrephilus (Marsh, 1970) Doryctobracon
areolatus (Szépligeti, 1911) Doryctobracon
arisanus (Sonan, 1932) Fopius
auripennis (Muesebeck, 1958) Doryctobracon
baldufi Muesebeck, 1949 Opius
bellus Gahan, 1930 Opius
bevisi (Brues, 1926) Fopius
bianchii (Fullaway, 1951) Utetes
brasiliensis (Szépligeti, 1911) Doryctobracon
brevistyli (Paoli, 1934) Diachasmimorpha
brevitemporalis (Tobias, 1998) Psyttalia
bucki Costa Lima, 1938 Opius
budrysi van Achterberg, 1999 Diachasmimorpha
canaliculatus (Gahan, 1915) Utetes
capsicola (Muesebeck, 1958) Doryctobracon
carinata (Szépligeti, 1910) Diachasmimorpha
carpomyiae (Silvestri, 1916) Fopius
caudatus (Szépligeti, 1913) Fopius
ceratitivorus Wharton, 1999 Fopius
concolor (Szépligeti, 1910) Psyttalia
cosyrae (Wilkinson, 1927) Psyttalia
crawfordi (Viereck, 1911) Doryctobracon
dacicida (Silvestri, 1912) Psyttalia
dacusii (Cameron, 1906) Diachasmimorpha
deeralensis (Fullaway, 1950) Fopius
desideratus (Bridwell, 1919) Fopius
dexter (Silvestri, 1913) Psyttalia
distinguenda (Granger, 1949) Psyttalia
divergens (Muesebeck, 1958) Utetes
downesi Gahan, 1919 Opius
feijeni van Achterberg, 1999 Diachasmimorpha
ferrari Carmichael and Wharton, 2005 Fopius
ferrugineum (Gahan, 1915) Diachasma
fijiensis (Fullaway, 1936) Psyttalia
fletcheri (Silvestri, 1916) Psyttalia
fluminensis (Costa Lima, 1938) Doryctobracon
frequens (Fischer, 1964) Utetes
froggatti Fullaway, 1950 Opius
fullawayi (Silvestri, 1913) Diachasmimorpha
hageni (Fullaway, 1952) Diachasmimorpha
halidayi Wharton, 2009 Psyttalia
hirtus Fischer, 1963 Opius
humilis (Silvestri, 1913) Psyttalia
illusorius (Fischer, 1971) Fopius
incisi (Silvestri, 1916) Psyttalia
inconsueta (Silvestri, 1913) Psyttalia
insignipennis (Granger, 1949) Psyttalia
itatiayensis (Costa Lima, 1937) Utetes
juglandis (Muesebeck, 1961) Diachasmimorpha
juniperi (Fischer, 1964) Utetes
kraussii (Fullaway, 1951) Diachasmimorpha
kurentzovi (Tobias, 1977) Utetes
lectoides (Gahan, 1930) Utetes
leveri (Fullaway, 1953) Psyttalia
longicaudata (Ashmead, 1905) Diachasmimorpha
lounsburyi (Silvestri, 1913) Psyttalia
magnus (Fischer, 1958) Utetes
makii (Sonan, 1932) Psyttalia
mandibularis Kimani-Njogu & Wharton, 2002 Rhynchosteres
manii (Fullaway, 1951) Utetes
masneri Wharton, 2009 Psyttalia
mariae Tobias, 1977 Opius
mellea (Gahan, 1915) Diachasmimorpha
muesebecki (Fischer, 1963) Psyttalia
muliebre (Muesebeck, 1956) Diachasma
myolejae (Tobias, 1977) Fopius
niger (Szépligeti, 1913) Fopius
novaguineensis (Szépligeti,1900) Psyttalia
okekai Kimani-Njogu & Wharton, 2002 Fopius
ophthalmica (Tobias, 1977) Psyttalia
ottotomoanus (Fullaway, 1957) Fopius
paeoniae (Tobias, 1980) Diachasmimorpha
perkinsi (Fullaway, 1950) Utetes
perproxima (Silvestri, 1913) Psyttalia
persulcatus (Silvestri, 1916) Fopius
phaeostigma (Wilkinson, 1927) Psyttalia
ponerophaga (Silvestri, 1916) Psyttalia
rhagoleticola (Sachtleben, 1934) Psyttalia
richmondi (Gahan, 1919) Utetes
rosae (Tobias, 1977) Utetes
rosicola (Muesebeck, 1949) Utetes
sanctamariana (Fischer, 1980) Psyttalia
sanguinea (Ashmead, 1889) Diachasmimorpha
schlingeri Wharton, 1999 Fopius
scrutator (Tobias, 1977) Utetes
silvestrii (Wharton, 1987) Fopius
skinneri (Fullaway, 1951) Fopius
sublaevis (Wharton, 1978) Diachasmimorpha
subsulcata (Granger, 1949) Psyttalia
tabellariae (Fischer, 1964) Utetes
tafivallensis Fischer, 1968 Opius
testaceus (Wesmael, 1838) Utetes
tomoplagiae (Costa Lima, 1937) Utetes
toxotrypanae (Muesebeck, 1958) Doryctobracon
trinidadensis (Gahan, 1919) Doryctobracon
tryoni (Cameron, 1911) Diachasmimorpha
ussuriensis (Tobias, 1977) Utetes
vandenboschi (Fullaway, 1952) Fopius
vierecki (Gahan, 1915) Utetes
vittator (Brues, 1926) Psyttalia
walkeri (Muesbeck, 1931) Psyttalia
zeteki (Muesebeck, 1958) Doryctobracon
Appendix 2
Status of other species group names (i.e. species, subspecies, and varieties not included in Appendix 1) recorded as parasitoids of fruit-infesting Tephritidae. Citations for original descriptions can be found in Fischer (1971, 1972, 1977, 1984, 1987), Wharton and Marsh (1978), Wharton and Gilstrap (1983), and Wharton (1987).
acidusae Fischer, 1967
Junior subjective synonym of tryoni: Wharton and Marsh (1978). This is the type species, by original designation, of Parasteres Fischer.
anastrephae Brèthes, 1924 (not anastrephae Viereck)
Junior subjective synonym of brasiliensis Szépligeti, 1911: Wharton and Marsh (1978). Described by Brèthes (1924) as Coeloides anastrephae.
angaleti Fullaway, 1952
Junior subjective synonym of albobalteata: Wharton (1987).
argentina Brèthes, 1924
Junior subjective synonym of anastrephae Viereck: Wharton and Marsh (1978). This is the type species, by original designation, of Bracanastrepha Brèthes.
berberidis Fischer, 1964
Junior subjective synonym of downesi: Marsh (1974).
brasilianus Fischer, 1963
Unnecessary replacement name for brasiliensis Szépligeti, 1911 (not brasiliensis Szépligeti, 1902, now in a different genus): Fischer (1977).
cereus Gahan, 1919
Junior subjective synonym of areolatus Szépligeti, 1911: Wharton and Marsh (1978).
chocki Fullaway, 1953; proposed as variety of longicaudata
Junior subjective synonym of longicaudata: Wharton and Gilstrap (1983).
coffeae Fischer, 1962
Junior subjective synonym of caudatus: Wharton (1987).
compensans Silvestri, 1916
Junior subjective synonym of longicaudata: Wharton and Gilstrap (1983).
comperei Viereck, 1913
Junior subjective synonym of longicaudata: Wharton and Gilstrap (1983). This is the type species, by original designation, of Diachasmimorpha Viereck.
conjungens Enderlein, 1920
Junior subjective synonym of crawfordi: Wharton and Marsh (1978). This is the type species, by original designation, of Doryctobracon Enderlein.
curtiarticulatus Fischer, 1963
Available name, proposed as subspecies of persulcatus.
formosanus Fullaway, 1926
Junior subjective synonym of longicaudata: Wharton and Gilstrap (1983).
fuscipennis Szépligeti, 1914
Misidentification of caudatus in some of the early biological control programs: Wharton and Gilstrap (1983). Originally described by Szépligeti (1914) as Rhinoplus fuscipennis.
giffardii Silvestri, 1913
Junior subjective synonym of carinata: Wharton (1987). Originally described by Silvestri (1913) as Hedylus giffardii.
glasgowi Fischer, 1964
Junior subjective synonym of frequens: Wharton and Marsh (1978).
gomesi Costa Lima, 1938
Junior subjective synonym of bellus: Wharton and Marsh (1978).
javanus Fullaway, 1920
Preoccupied by javanus Szépligeti, 1908; replaced by vandenboschi: Wharton and Gilstrap (1983).
lectus Gahan, 1919
Junior subjective synonym of canaliculatus: Wharton and Marsh (1978).
lumpurensis Fischer, 1963
Available name, proposed as subspecies of persulcatus.
malaiaensis Fullaway, 1953; proposed as variety of longicaudata.
Junior subjective synonym of longicaudata: Wharton and Gilstrap (1983).
modestior Silvestri, 1913; proposed as variety of perproxima
Junior subjective synonym of perproxima: Wharton and Gilstrap (1983).
mombinpraeoptantis Fischer, 1966
Junior subjective synonym of anastrephae Viereck: Wharton and Marsh (1978).
novocaledonicus Fullaway, 1953; proposed as variety of longicaudata
Junior subjective synonym of longicaudata: Wharton and Gilstrap (1983).
oophilus Fullaway, 1951
Junior subjective synonym of arisanus: Wharton and Gilstrap (1983).
orientalis Silvestri, 1913; proposed as variety of africanus Szépligeti, 1910.
Available name, but not specifically treated since it was proposed.
rhagoletis Richmond, 1915
Junior subjective synonym of mellea: Gahan (1919).
robusta Silvestri, 1913; proposed as variety of fullawayi.
Available name, but not specifically treated since it was proposed.
saopaulensis Fischer, 1961
Unnecessary replacement name for areolatus Szépligeti, 1911 (not areolatus Szépligeti, 1902, now in a different genus): Wharton and Marsh (1983).
siculus Monastero, 1931
Junior subjective synonym of concolor: Wharton and Gilstrap (1983).
sayanicus Tobias, 1977
Junior subjective synonym of magnus: Fischer (1984).
substriatus Fischer, 1963
Available name, proposed as subspecies of persulcatus.
taiensis Fullaway, 1953; proposed as variety of longicaudata.
Junior subjective synonym of longicaudata: Wharton and Gilstrap (1983).
tephritivorus Wharton, 1987
Unnecessary replacement name for africanus Szépligeti, 1910 (not africanus Szépligeti, 1908, now in a different genus).
trimaculatus Spinola, 1851
Misidentification of anastrephae, bellus, or one of the species of Doryctobracon: Wharton and Gilstrap (1983).
tucumanus Blanchard, 1966
Junior subjective synonym of areolatus Szépligeti, 1911: Ovruski and Wharton (1996).
tucumanus Turica and Mallo, 1961
Nomen nudum, = tucumanus Blanchard: Ovruski and Wharton (1996).
turicai Blanchard, 1966
Junior subjective synonym of bellus: Wharton and Marsh (1978).
watersi Fullaway, 1951
Junior subjective synonym of dacusii: Wharton and Gilstrap (1983)
Appendix 3
Generic names used in association with tephritid parasitoids and their current status. For citations to original descriptions, see Fischer (1971, 1972, 1977, 1987).
Austroopius Szépligeti, 1900
Synonym of Psyttalia (Wharton 1987). Tephritid parasitoids formerly included here have usually been referred to as Opius in the biological control literature.
Biosteres Förster, 1862
Valid genus. Tephritid parasitoids formerly included here have now been transferred mostly to Diachasmimorpha and Fopius. See Wharton (1988).
Bracanastrepha Brèthes, 1924
Synonym of Utetes (Wharton 1987, 1988). Tephritid parasitoids formerly included here were referred to as Opius in most of the biological control literature prior to 1979.
Desmiostoma Förster, 1862
Synonym of Opius (Wharton 1983, 1988).
Diachasma Förster, 1862
Valid genus (but see Tobias 1977).
Diachasmimorpha Viereck, 1913
Valid genus for some of the most widely used parasitoids in tephritid biocontrol. Prior to 1987, most of these species were placed in either Biosteres or Opius, and more rarely Parasteres and Diachasma (Wharton 1987).
Doryctobracon Enderlein, 1920
Valid genus. Prior to 1978, these species were placed either in Opius or Parachasma (Fischer 1977, Wharton and Marsh 1978).
Fopius Wharton 1987
Valid genus, originally described as a subgenus of Rhynchosteres (Wharton 1987, van Achterberg and Maetô 1990). Includes some of the most significant parasitoids used in tephritid biocontrol. Most of these species have been placed in Biosteres in recent literature, and in Opius prior to about 1970.
Hedylus Marshall, 1891
A junior homonym of Hedylus Förster, 1862, an ichneumonid. One species (giffardii) was originally described in Hedylus.
Opius Wesmael, 1835
Valid genus. Most of the tephritid parasitoids formerly included in Opius have been transferred elsewhere. A few of the tephritid parasitoids are still placed in Opius pending further study of this very large and undoubtedly paraphyletic genus.
Parachasma Fischer, 1967
Junior subjective synonym of Doryctobracon (Fischer 1972, Wharton and Marsh 1978).
Parasteres Fischer, 1967
Junior subjective synonym of Diachasmimorpha (Wharton and Marsh 1978, Wharton 1987).
Psyttalia Walker, 1860
Valid genus for some of the most widely used parasitoids in tephritid biocontrol (Wharton 1987, 1988). Most of these species have been referred to as Opius in the biological control literature.
Utetes Förster, 1862
Valid genus. Tephritid parasitoids have been referred to as Opius in most of the literature prior to about 1990 (Wharton 1988). Treated by Fischer as a subgenus of Opius.
