Fruit Fly Expert Identification System and Systematic Information

Norrbom Status of Knowledge 31
Table 6. Number of species of Tephritidae for which some immature stages have been described.
Adults Immatures
Region Genera Species Genera (%) Species (%)
Afrotropical 151 920 20 (13.2) 35 (3.8)
Australasian 144 762 3 (2.1) 20 (2.6)
Nearctic 60 358 30 (50.0) 80 (22.3)
Neotropical 68 717 9 (13.2) 31 (4.3)
Oriental 155 943 4 (2.6) 12 (1.3)
Palearctic 126 827 39 (31.0) 109 (13.2)
World 471 4,257 91 (19.3) 291 (6.8)
Numbers for immatures based on 1992 data
some species of Tephritinae. The anterior spiracles are sessile
or nearly so, and usually fan-shaped or bimodal, with 2-53
papillae (Ferrar 1987). The cephalopharyngeal skeletons of all
species so far described are typically muscomorph, with right
and left mandibles equally developed, and sometimes with 1-2
subapical teeth persisting into the third instar (as far as known,
subapical teeth are always well-developed in earlier instars).
Fruit fly puparia are usually barrel-shaped, but rarely are beanshaped
or have a flattened caudal segment. They may be smooth
or wrinkled, with distinct or unclear segmentation. They range
in color from white to yellow, brown or black, and their color
may vary from the anterior to the posterior end.
Morphology & Terminology
McAlpine (1981) proposed a morphological terminology
for adult Diptera that is standardized with that of other insects
and is well illustrated. Proposed modifications to this terminology
or more specific applications of it to the Tephritidae include
Steyskal (1984), Norrbom & Kim (1988[3666]), Freidberg &
Mathis (1986), White (1988), White & Elson-Harris (1992) and
Foote et al. (1993). White & Elson-Harris (1992) provided an
extensive glossary. Extensive discussions and/or illustrations
of morphological terminology were also included by Hendel
(1927[2109]; Hering and Ito used this terminology), Shiraki
(1933), Munro (1947, 1984), Foote (1980), Freidberg &
Kugler (1989), Kapoor (1993; includes most terms used by
Hardy) and Merz (1994[3343]). Some additional useful references
on tephritid adult morphology include: general -Aczél
(1955[28], May (1963[3232]), Zaka-ur-Rab (1971), Valdez-
Carrasco & Prado-Beltran (1991); digestive system - Dean
(1933), Kobayashi (1934), Zaka-ur-Rab (1971); antenna -
Giannakakis & Fletcher (1985), Vasey & Ritter (1987), Dickens
et al. (1988), Bigiani et al. (1989), Crnjar et al. (1989, 1989),
Mayo et al. (1987); compound eye - Agee et al. (1977), Davis
et al. (1983); head -Nayar (1961[3567]); integument - Evans
(1967[1358]); mouthparts - Driscoll & Condon (1994); musculature
and internal anatomy - Zaka-ur-Rab (1971), Berube &
Zacharuk (1983[429]), Valdez-Carrasco & Prado-Beltran
(1991); nervous system -Kobayashi (1934), Zaka-ur-Rab
(1971); reproductive systems and genitalia -Kobayashi (1934),
Dean (1935), Hanna (1938), Drew (1969), Zaka-ur-Rab (1971),
Dodson (1978[1193]), Korneyev (1979, 1985[2715]),
Zacharuk et al. (1986), Stoffolano & Yin (1987), Norrbom &
Kim (1988[3666]), Williamson (1989), Eberhard & Pereira
(1993), De Carlo et al. (1994); and thorax - Nayar (1962),
Verma (1985).
Freidberg & Kugler (1989) defined various types of tephritid
wing patterns, and terminologies for the parts of various
banded patterns were proposed by Lima (1934[2954]) and
Stone (1942[4674]) for Anastrepha, Bush (1966[683]),
Steyskal (1979), Foote (1981), and Freidberg & Hancock
(1989) for Rhagoletis, Cryptophorellia and Urophora, which
are similar in wing pattern, Munro (1984) for the Dacina,
Freidberg (1991) for Ceratitis, and Freidberg & Kaplan (1992)
for the Oedaspidina. White & Elson-Harris (1992) used standardized
terms for similar bands in several economic genera.
Terminology for the larvae of Diptera was recently reviewed
by Teskey (1981, 1991), and White & Elson-Harris
(1992) provided an extensive glossary of morphological terms
for larval Tephritidae. Phillips (1946) summarized previous
terminologies for the cephalopharyngeal skeleton and presented
a terminology for the caudal segment. Carroll & Wharton
(1989) proposed a terminology for specific sensilla.
Terminology for sense organs of the head is confusing; four
different systems have been used in the Tephritidae. That of
Teskey (1981), following Weismann (1864), has been the most
common and was used by Efflatoun (1927), Kandybina (1977),
Steck & Wharton (1986, 1988), Steck & Malavasi (1988),
Carroll & Wharton (1989), Carroll (1992) and White & Elson-
Harris (1992). The other terminologies include: Snodgrass
(1924), used by Phillips (1946) and Exley (1955); Bolwig
(1946), used by Headrick, Goeden and coauthors (e.g., Goeden
& Headrick (1990)); and Snodgrass (1953), used by Novak &
Foote (1968, 1975, 1980) and Steck (1984).
Detailed, comprehensive anatomical studies of tephritid
larvae include Snodgrass (1924), Knell & Stoffolano (1973),
Zaka-ur-Rab (1978[5283], 1978[5284], cephalopharyngeal
musculature, tracheal system), Dean (1932, alimentary canal,
1942, reproductive system), Bates (1934[351], peristigmal
gland cells), and Butt (1937) and Jones & Kim (1988) (posterior
stigmatic apparatus).
Taxonomic Knowledge of Immature Stages
In spite of the economic importance of Tephritidae, the
immature stages (eggs, larvae, puparia) are poorly known. The
figures in the classification table are overestimated, as many of