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Anastrepha bezzii Lima

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Recognition
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Anastrepha bezzii differs from all other Anastrepha species in its elongate pattern of toothlike scales on the eversible membrane, and in the shape of the aculeus tip. Species of the doryphoros group have a somewhat similar pattern of eversible membrane scales, but not as elongate and with the scales themselves much shorter. Anastrepha bezzii differs from most Anastrepha species in having a complete S-band that is broadly fused to the C-band along the costa, so that there is no hyaline area in cell r1 distal to the apex of vein R1. It differs from the other species with this type of wing pattern in having a hyaline area in cell r2+3 between crossveins BM-Cu and R-M; the section of vein M between BM-Cu and R-M less than 1.85 times as long as section between R-M and DM-Cu; and vein R2+3 usually undulant.

Classification and Evolutionary Relationships
Order: Diptera. Family: Tephritidae. Genus: Anastrepha. Species: bezzii. Author: Lima.
Relationships among the species of Anastrepha were analyzed by Norrbom et al. (1999) and McPheron et al. (1999). Click here for more detailed discussion of Anastrepha phylogeny. Anastrepha bezzii has been placed in the mucronota species group.

Names Used for this Species
Anastrepha bezzii Lima 1934: 498.
Anastrepha balloui Stone 1942: 12. Synonymy (Norrbom 1991).
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Type Data
bezzii: Holotype - Male (Instituto Oswaldo Cruz, no. 1796), "Brazil, Rio de Janeiro, Manguinhos, 24.II.1902". Zucchi (1978) reexamined it and reported its date of collection as 24.III.1917. Lima dissected and mounted the abdomen and a wing on slides 1876 and 1877.
balloui: Holotype - Female (USNM), Venezuela: Guarico: San Juan de los Morros, on Terminalia catappa, 12.IV.1938, C. H. Ballou. It bears a labels with "S. J. los Morros, Venez, 249 C.H. Ballou, 4-12-1938" and "on Terminalia catappa", a red "Type No. 51652 U.S.N.M" label, and a Stone determination label with "Anastrepha balloui Stone". Stone dissected and mounted the terminalia on slide 39.IV.17^a.

Distribution
Anastrepha bezzii is known from Mexico (Chiapas) south to Venezuela, following the Andes to southern Brazil (Mato Grosso do Sul, Goiás, Minas Gerais, Rio de Janeiro).
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Biology
The only well confirmed host plants of Anastrepha bezzii are Sterculia apetala (Jacq.) Karst. (Caraballo 1981) and S. chicha St. Hil. (Santos et al. 1993), although Guagliumi (1966) also listed Terminalia catappa L. as a host. The latter record may be erroneously based on Stone's (1942) report that some of the types of balloui were collected "on" T. catappa, but Guagliumi's listing (p. 232) of two Opius species as "Parassita di Anastrepha balloui Stone s/ Terminalia" suggests that he might have seen reared specimens. Fernandez-Yepez (in Teran 1980) considered this record doubtful, and Caraballo (1981) was unable to confirm it. The larvae reared from S. chica in Brazil fed inside the seed rather than in the pulp of the fruit.

Economic Significance
Anastrepha bezzii is not considered a pest species.

Comments
There is considerable variation in terminalia length among specimens that Norrbom (1991) recognized as bezzii (Table 1, Fig. 5). Stone (1942) and subsequent authors recognized populations from Mexico to Venezuela and Peru as a distinct species, A. balloui, because they have the oviscape, the aculeus, the part of the eversible membrane with dorsal scales, and the phallus shorter than in the few Brazilian specimens previously known. The lengths of all of these structures are correlated, however, and these differences are all the result of variation in terminalia length. The distinctive shape of the aculeus tip, the pattern of the scales of the eversible membrane, the shape of the surstyli, and the wing pattern and venation are consistent in the Brazilian and non-Brazilian specimens.

Table 1 and Figure 5 of Norrbom (1991) present an analysis of the variation in the length and relative length (ratio to mesonotum length) of the oviscape between samples from Mexico-Panama, Venezuela, and Brazil. All measurements were taken from examined specimens with the addition of those for single females from Viçosa, Brazil (Zucchi 1984) and Venezuela (Caraballo 1981). The measurements of the female reported by Lima (1937) from Campinas, Goiás, Brazil, which unfortunately has been lost (Zucchi 1984), are plotted for comparison, but are not included in the analysis. Caraballo (1981) found greater variation in oviscape length in a larger sample of females from Venezuela (5.56-7.30 mm; n=75). She did not report the relative length, but her range of mesonotal lengths suggest that it is consistent with that in the specimens Norrbom examined.

Oviscape length and relative length are significantly different (Ttest, p<0.01) between all samples, as is the arcsine transformation of the inverse of the latter ratio. Mesonotum length is significantly different between Brazil and Venezuela (p<0.01) and Brazil and Mexico-Panama (p<0.02), but not between Venezuela and Mexico-Panama. Thus, although the Brazilian females have longer terminalia, their relative terminalia length is closer to that of the Mexico-Panama sample than to that of the Venezuela sample or the Campinas, Brazil female of Lima (1937) (Fig. 5B). Similarly, the regression lines for these two samples are closer than that for the Venezuela sample (Fig. 5A). There does not appear to be an allometric relationship, at least within samples (Fig. 5B).

Norrbom (1991) did not examine the male holotype of bezzii which is assumed to be conspecific with the Viçosa sample. The type locality of Rio de Janeiro is much closer to Viçosa than to Campinas. At least the Mexico-Panama populations seem to be conspecific, and although the relative terminalia length of the Venezuelan sample is significantly different statistically, whether there is any biological significance is questionable. The status of the Campinas female is less certain since Lima's measurements cannot be verified, but if they are accurate, it is at least as different from the other Brazilian specimens as the Venezuela population.

Norrbom (1991) did not believe there was enough evidence to justify the recognition of the Venezuelan population as a distinct species, and therefore regarded balloui Stone a synonym of bezzii. Continued use of this name is not justifiable without also regarding the Campinas female as representative of another separate species. Unless other characters or behavioral or biological studies demonstrate otherwise, all of these specimens should be considered conspecific. Additional collecting, an analysis of larger sample sizes, and biochemical or behavioral studies to test the statistical and biological significance of the differences among them would certainly be useful.

The scutal color pattern in bezzii is faint and difficult to see. It is often obscured by underlying tissues in preserved specimens, especially those preserved in fluid prior to pinning.

References
Key references are listed below. See fruit fly literature database for additional references.
Aluja, M., M. Cabrera, E. Rios, J. Guillén, H. Celedonio, J. Hendrichs, and P. Liedo. 1987. A survey of the economically important fruit flies (Diptera: Tephritidae) present in Chiapas and a few other fruit growing regions in Mexico. Fla. Entomol. 70: 320-329. [Mexico]
Malavasi, A. & R. A. Zucchi, eds. 2000. Moscas-das-frutas de importância econômica no Brasil. Conhecimento básico e aplicado. Holos, Riberão Preto. 327 p. [Brazil]
McPheron, B. A., H.-Y. Han, J. G. Silva & A. L. Norrbom. 1999. Phylogeny of the genera Anastrepha and Toxotrypana (Trypetinae: Toxotrypanini) based upon 16S rRNA mitochondrial DNA sequences, p. 343-361. In M. Aluja & A. L. Norrbom, eds., Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC Press, Boca Raton. [16] + 944 p. [phylogeny]
Norrbom, A. L. 1991. The species of Anastrepha (Diptera: Tephritidae) with a grandis-type wing pattern. Proc. Entomol. Soc. Wash. 93: 101-124. [revision]
Norrbom, A. L., R. A. Zucchi & V. Hernández-Ortiz. 1999. Phylogeny of the genera Anastrepha and Toxotrypana (Trypetinae: Toxotrypanini) based on morphology, p. 299-342. In M. Aluja & A. L. Norrbom, eds., Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC Press, Boca Raton. [16] + 944 p.  [classification & phylogeny]
Santos, G. P., Anjos, N., Zanuncio, J. C. & Assis Júnior, S. L. 1993. Danos e aspectos biológicos de Anastrepha bezzii Lima, 1934 (Diptera, Tephritidae) em sementes de Sterculia chicha St. Hill. (Sterculiaceae). Rev. Bras. Entomol. 37: 15-18.
Teran B., J. 1980. Lista preliminar de Hymenoptera parasitos de otros insectos en Venezuela. Rev. Fac. Agron. (Maracay) 10 (14): 283-389.