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Zonosemata Benjamin (Diptera: Tephritidae)

Zonosemata electa, dorsal habitus, female and male.

Recognition
Species of Zonosemata are mostly yellow, with a few dark brown markings on the abdomen and often on the thorax. The scutum has three white stripes, including a medial one, and often has dark brown markings at least posteriorly (often somewhat U-shaped). The wing is mostly hyaline with 4-5 narrow to moderately broad transverse bands in a "spider-mimic" pattern.
Additional useful diagnostic characters include: ocellar seta and posterior orbital seta well developed; usually 4 or more frontal setae; antennal first flagellomere with small acute dorsoapical lobe; arista pubescent; scutum and most of anepimeron without microtrichia; dorsocentral seta closer to level of intra-alar seta than to level of postsutural supra-alar seta; presutural supra-alar seta well developed; scutellum with 2 pairs of large marginal setae; crossvein R-M near midlength of cell dm and covered by discal band; crossvein DM-Cu covered by subapical band; vein Cu nonsetulose; lateral surstylus short and without medial mesal lobe; glans relatively simple, evenly sclerotized, with little distinct internal sclerotization; aculeus tip with small subapical notches; and spermathecae large, elongate, and weakly sclerotized.
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Classification and Evolutionary Relationships
Order: Diptera. Family: Tephritidae. Subfamily: Trypetinae. Tribe: Carpomyini. Genus: Zonosemata. Author: Benjamin.
Zonosemata belongs to the subtribe Carpomyina of the Trypetinae. Jenkins (1996) questioned whether the apical desclerotized area of the female oviscape (syntergosternite 7) considered a synapomorphy for the Carpomyina by Norrbom (1989), is present in the genus, but it is present (Norrbom 2002). The genitalia of the specimens from the USNM that were dissected by Jenkins are overcleared and this character can no longer be seen in them, but it is easily seen in less cleared dissections.
The phylogenetic relationships of the Carpomyina were reviewed by Smith & Bush (1999). The relationships among the genera of this tribe are not well resolved, but Zonosemata may be most closely related to Cryptodacus Hendel, Haywardina Aczél and Rhagoletotrypeta Aczél. All four genera are endemic to the Americas and possess a whitish medial scutal stripe or spot that may be a synapomorphy (Norrbom 1994). Few representatives of these genera have been included in molecular studies of the tribe; when included they tend to cluster together or sometimes also with certain species of Rhagoletis Loew, but further work and the inclusion of additional species are needed to test this hypothesis.  
Norrbom (1994), Jenkins (1996) and Smith & Bush (1999) listed the following probable synapomorphies for Zonosemata: scutum without microtrichia, which occurs as homoplasy in the Rhagoletis psalida species group, Haywardina cuculiformis and Rhagoleotrypeta xanthogastra (to this may be added, anepimeron mostly bare of microtrichia - this and additional areas of the pleuron are bare in the R. psalida group, but the anepimeron is microtrichose in H. cuculiformis and R. xanthogastra); lateral surstylus short and without medial mesal lobe; glans relatively simple, evenly sclerotized, with little distinct internal sclerotization; spermathecae large, elongate, and weakly sclerotized; and spermathecal duct annulated. Other possible synapomorphies (i.e., states of characters whose polarity depends upon the outgroup) are the posterior location of the dorsocentral seta and the notched aculeus tip. Bush (1966) also suggested that there are apomorphic traits in chromosome shape for Zonosemata, but karyotypes have not been studied in other carpomyine genera except Rhagoletis. Relationships among the species of Zonosemata have not been analyzed.

Diversity & Distribution
Zonosemata includes eight species that are endemic to the New World. Their combined range extends from southeastern Canada to Colombia and northeastern Brazil. One species is known from the Greater Antilles (Jamaica).
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Click here for list of included species and links to species pages.
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Names Used for this Genus
Zonosemata Benjamin 1934: 17 (Type species: Trypeta electa Say by original designation).
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Click here to link to fly names database for full nomenclatural data.

Biology
Zonosemata species breed in fruits of species of Solanaceae, mainly in the genus Solanum (Smith & Bush 1999, Norrbom 2002). All but one of the species have been reared from or at least associated as adults with solanaceous plants. Only Z. electa (Say), has been reared from genera other than Solanum. Smith and Bush (1999: 197) listed the known host plants of most of the species.
Various aspects of the biology of Zonosemata species were reviewed by Prokopy & Papaj (1999).  The behavior of Z.vittigera was studied by Greene et al. (1987) and Whitman et al. (1988) who indicated that the adults hold their banded wings such that they resembled jumping spiders, affording them some protection against these predators. The other species of Zonosemata, which are similarly marked, may also exhibit this behavior.

Economic Significance
Zonosemata electa (Say), commonly known as the pepper maggot, is considered a pest of pepper and eggplant in the eastern United States and Canada.

References
Key references are listed below. See fruit fly literature database for additional references.
Benjamin, F. H. 1934. Descriptions of some native trypetid flies with notes on their habits. U.S. Dep. Agric. Tech. Bull. 401: 95 p. [p. 17, description, taxonomy, immature stages]
Bush, G. L. 1966. The genus Zonosemata, with notes on the cytology of two species. Psyche (Cambridge) (1965) 72: 307-323. [revision, species descriptions]
Foote, R. H., Blanc, F. L. & Norrbom, A. L. 1993. Handbook of the fruit flies (Diptera: Tephritidae) of America north of Mexico. Comstock Publishing Associates, Ithaca. xii + 571 p. [2 spp. in U.S.A. and Canada]
Greene, E., L. J. Orsak & D. W. Whitman. 1987. A tephritid fly mimics the territorial display of its jumping spider predators. Science 236: 310-312. [biology]
Hernández-Ortiz, V. 1989. Una especie nueva de Zonosemata (Díptera: Tephritidae) y clave de identificación de las especies del género. Anales del Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Serie Zoologia 60: 205-210. [key, species description]
Jenkins, J. 1996. Systematic studies of Rhagoletis and related genera (Diptera: Tephritidae). Dissertation, Michigan State University, East Lansing, 184 p.
Norrbom, A. L. 1989. The status of Urophora acuticornis and U. sabroskyi (Diptera: Tephritidae). Entomological News 100: 59-66.
Norrbom, A. L. 1990. Notes on Zonosemata Benjamin (Diptera: Tephritidae) and the status of Cryptodacus scutellatus Hendel (= Z. ica Steyskal syn. n.). Annalen des Naturhistorisches Museums in Wien 91: 53-55. [synonymy, taxonomy, hosts, distribution]
Norrbom, A.L. 1994. New species and phylogenetic analysis of Cryptodacus, Haywardina and Rhagoletotrypeta (Diptera: Tephritidae). Insecta Mundi 8: 37-65. [generic relationships]
Norrbom, A. L. 2002. A new species and key for the genus Zonosemata Benjamin (Diptera: Tephritidae). Proceedings of the Entomological Society of Washington 104: 614-623. [species description, key, hosts, distribution]
Norrbom, A. L., L. E. Carroll, F. C. Thompson, I. M. White & A. Freidberg. 1999. Systematic database of names, pp. 65-251. In F. C. Thompson (ed.), Fruit Fly Expert Identification System and Systematic Information Database. Myia 9, vii + 524 pp. & Diptera Data Dissemination Disk (CD-ROM) (1998) 1. [p. 249, in catalog]
Prokopy, R. J. & D. R. Papaj. 1999. Behavior of flies of the genera Rhagoletis, Zonosemata, and Carpomya (Trypetinae: Carpomyina), p. 219-252. In M. Aluja & A. L. Norrbom, eds., Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC Press, Boca Raton. [16] + 944 p.
Smith, J. J. & G. L. Bush. 1999. Phylogeny of the subtribe Carpomyina (Trypetinae), emphasizing relationships of the genus Rhagoletis, p. 187-217. In M. Aluja and A. L. Norrbom, eds., Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC Press, Boca Raton. [16] + 944 p. [review, host plants]
Steyskal, G. C. 1975. Recognition characters for larvae of the genus Zonosemata (Diptera, Tephritidae). U. S. Department of Agriculture Cooperative Economic Insect Report 25 (12): 231-232. [larvae of 2 spp.]
Whitman, D. W., L. J. Orsak & E. Greene. 1988. Spider mimicry in fruit flies: further experiments on the deterence of jumping spiders (Araneae: Salticidae) by Zonosemata vittigera (Coquillett). Annals of the Entomological Society of America 81: 532-536. [biology]