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Fruit Fly Expert Identification System and Systematic Information

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Norrbom Status of Knowledge 19<br />

Incertae Sedis Tephritidae<br />

Genera: PA, 3 (3); OR, 1 (1); total, 4. Species: NT, 5 (5); PA,<br />

23 (23); AF, 1 (1); OR, 1 (1); AU, 4 (4); total, 34.<br />

Chejuparia: PA, 1 (1); total, 1.<br />

Oxyphora: PA, 1 (1); total, 1.<br />

Pseudorellia: OR, 1 (1); total, 1.<br />

Stylia: PA, 1 (1); total, 1.<br />

Unplaced species: NT, 5 (5); PA, 20 (20); AF, 1 (1); AU, 4 (4); total,<br />

30.<br />

Recent changes from previous classifications<br />

Tephritid higher classification is currently in a state of<br />

rapid flux. Numerous changes have been proposed since publication<br />

of the most recent regional catalogs (Cogan & Munro<br />

1980, Foote 1965[1502], 1967[1508], 1984, Hardy 1977,<br />

Hardy & Foote 1989), which themselves differ in classification.<br />

More emphasis on comprehensive, worldwide studies rather<br />

than regional faunas, more thorough study of genitalic characters,<br />

<strong>and</strong> introduction of phylogenetic analysis have contributed<br />

to these advances <strong>and</strong> doubtless will continue to improve our<br />

knowledge of fruit fly relationships. Here we attempt to list the<br />

most significant changes in higher classification since publication<br />

of the regional catalogs.<br />

We have generally avoided the recognition of monogeneric<br />

higher taxa in the above classification. Unless their sister<br />

group relationships are understood, such taxa add little predictive<br />

value to a classification. Genera whose relationships are<br />

uncertain are instead listed under “Incertae sedis” (unplaced).<br />

For the Phytalmyiinae, we generally follow the classification<br />

of Korneyev (1994[2744]), but we prefer to include within<br />

it several other taxa (Acanthonevrini, Blepharoneurini, Epacrocerini,<br />

<strong>and</strong> Phascini) that he ranked as separate subfamilies.<br />

These taxa have often been included within the Trypetinae.<br />

Hardy (1980[1949], 1986[1962]) <strong>and</strong> Hancock (1986[1890])<br />

also included important discussions of the classification of the<br />

Acanthonevrini, <strong>and</strong> Hardy (1983[1957]) <strong>and</strong> McAlpine &<br />

Schneider (1978) discussed the Phytalmiini. Following Korneyev<br />

(1994[2744]), Terastiomyiini is considered a synonym<br />

of Phytalmiini. According to Korneyev (pers. comm.), Homoiothemara<br />

also belongs in the Phascini.<br />

Within the subfamily Trypetinae, Euphrantini are no<br />

longer recognized as distinct from Adramini (White & Elson-<br />

Harris 1992, Korneyev 1994[2744]). Korneyev (1994[2744])<br />

<strong>and</strong> Hardy (1983[1958], 1986[1961]) discussed the limits <strong>and</strong><br />

included genera of this group. Genera that were previously<br />

included only on the basis of having reduced chaetotaxy <strong>and</strong> a<br />

strongly sclerotized bridge behind the metathoracic coxae are<br />

here mostly included in the Phytalmiini. The tribe Carpomyini<br />

was recently proposed as a subtribe of Trypetini (Norrbom<br />

1989[3653], 1994[3663]), but we follow Han & McPheron<br />

(1994) in ranking it as a tribe because its relationship with the<br />

Trypetini is uncertain. Korneyev (1996 [2747]) recently added<br />

two small subtribes, the Notommatina <strong>and</strong> Paraterelliina.<br />

The Dacina, usually previously ranked as a subfamily, or<br />

even as a separate family (Munro 1984), because of their<br />

distinctive appearance <strong>and</strong> the large size of the group, is now<br />

included as a subtribe of Trypetinae. It is included in the tribe<br />

Dacini with the Ceratitidina <strong>and</strong> Gastrozonina (Hancock<br />

1986[1890], Foote et al. 1993). Drew (1989[232], 1989[1231])<br />

<strong>and</strong> Drew & Hancock (1994[1239]) discussed the generic <strong>and</strong><br />

subgeneric classification (see also Hardy 1955[1927] <strong>and</strong><br />

Drew 1972[1216], 1979). The large genus Dacus has been<br />

divided, <strong>and</strong> many of its subgenera <strong>and</strong> species are now placed<br />

in Bactrocera. Munro (1984) proposed numerous subfamilies<br />

<strong>and</strong> tribes within the Dacina (as Dacidae), but his classification<br />

has not been accepted (Hancock 1986[1890], Drew<br />

1989[232]). We follow the synonymy suggested by White &<br />

Elson-Harris (1992) for the genus group names Munro proposed.<br />

The classification of the Ceratitidina was briefly discussed<br />

by Hancock (1984, 1987). The limits of this group <strong>and</strong> the<br />

Gastrozonina are poorly resolved. Hancock (1985[1889]) <strong>and</strong><br />

Hancock & Drew (1994[1901]) included several genera without<br />

a strongly apically pointed antennal first flagellomere that<br />

traditionally have been classified under Gastrozonina (Hardy<br />

1973, 1974[1943], 1988[1964]). The Gastrozonina have been<br />

considered as a subordinate taxon to Acanthonevrini (Hardy<br />

1988[1964]), but Hancock (1985 [1889]) considered them<br />

related to the Ceratitidina. Hancock (1985[1889], 1991[1895]),<br />

Hancock & Drew (1994[1901]), <strong>and</strong> Hardy (1988[1964]) discussed<br />

the definition of the group <strong>and</strong> the included genera. The<br />

correct stem for the family group name based on Ceratitis is<br />

Ceratitid-, rather than Ceratit-, <strong>and</strong> a name based on the latter<br />

stem is preoccupied by the ammonite name Ceratitidae Mojsisovics<br />

(D.L. Hancock, C.W. Sabrosky, pers. comm.).<br />

Hancock (1986[1890], 1991[1895]), Norrbom (1985,<br />

1994[3662]), <strong>and</strong> Norrbom & Foote (1989) discussed the classification<br />

of the Ortalotrypetini, Rivelliomimini, <strong>and</strong> Toxotrypanini,<br />

recognized here as tribes. Ischyropteron also belongs in<br />

the Ortalotrypetini (Norrbom, pers. obs.), <strong>and</strong> Hexachaeta is<br />

here included in the Toxotrypanini based on molecular data<br />

(Han & McPheron, 1997). A number of Neotropical genera<br />

previously included in the Trypetinae (R.H. Foote 1967[1508])<br />

have been transferred to the Tephritinae (Norrbom 1988, Foote<br />

et al. 1993).<br />

The Trypetini were recently redefined by Han (1992). His<br />

classification is used here, except that the subtribes Acidoxanthina<br />

<strong>and</strong> Nitrariomyiina recently proposed by Korneyev<br />

(1996 [2747]) <strong>and</strong> several additional genera are also included.<br />

Hancock (1986[1890]) suggested that the Acidoxanthina belong<br />

in the Ceratitidina.<br />

The Zaceratini (Hancock 1986[1890]) <strong>and</strong> Plioreoceptini<br />

(Korneyev 1987[2726]) were each proposed as monogeneric<br />

tribes. Their synonymy was noted by White & Elson-Harris<br />

(1992).<br />

The subfamily Tephritinae includes some taxa previously<br />

given subfamily rank: the Myopitini, Tephrellini (as Aciurinae),<br />

Terelliini, Oedaspidina, <strong>and</strong> Schistopterini (e.g., Hering<br />

1947, Cogan & Munro 1980). The Myopitini <strong>and</strong> Tephrellini<br />

have sometimes been included in the Trypetinae (e.g., Hardy<br />

1977). The limits of many of the tribes of the Tephritinae are<br />

vague (Hancock 1986[1891], Foote et al. 1993), <strong>and</strong> Hancock<br />

(1990) included all of them except the Myopitini, Terelliini,<br />

<strong>and</strong> some Tephrellini within the Tephritini.<br />

Hancock (1990) indicated that Tephrellini is the valid<br />

name for the group previously called Aciurini. In the classification<br />

followed here (Cogan & Munro 1980, Freidberg, pers.<br />

obs.), it also includes the Platensinina, sometimes recognized

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