**2.3 The family Tephritidae**

Tephritidae (true flies) is a very large family, which includes more than 4000 described species. Worldwide members of this family are among the most economically crucial insect pests of edible fruits and vegetables [20, 21]. The family can be characterized by an elaborate wing patterns and the possession of a telescopic ovipositor by the female. Tephritidae is known as one of the most ecologically diverse families of Diptera, and due to its size, it has been difficult to synthesize phylogenetic relationships among higher groups of the family [22]. Phylogenetic relationships of important genera of the family have been provided by [23]. Despite the lack of a conclusive phylogeny, the study of Tephritidae can be approached by looking separately at five different subfamilies; Blepharoneurinae, Phytalmiinae, Trypetinae, Dacinae and Tephritinae, all of which are well represented in the tropics.

The subfamily Blepharoneurinae represents flies of the tropical group, and composed of five main genera; *Ceratodacus, Problepharoneura, Blepharoneura, Baryglossa,* and *Hexaptilona.* The first three genera consist of species of the neotropical and afrotropical regions, while the last two genera include species of the palearctic regions. Although this subfamily is composed of a reduced number of described species, recent scrutiny on flies in the genus *Blepharoneura* suggests that there may be more than 200 species. This subfamily is interesting as the group appears to be one of the oldest lineages in Tephritidae. All the genera for which biological data have been gathered feed on plants and parts of plants in the family Cucurbitaceae. There is suggestive evidence that these flies have undergone rapid processes of

speciation, as much as more than one species can cohabit the same plant, exploiting different parts of it and exhibiting complex courtship behaviors [24].

Phytalmiinae is a subfamily comprising six genera; *Diplochorda, Ortaloptera, Phytalmia, Sessilinia, Tetrastiomyia,* and *Sophiria.* These are the flies with antennalike head projections and sometimes referred to as antler flies or deer flies (not to be confused with Tabanidae). Decaying plant material is the larval food across this subfamily. All described species of antler flies occur between the island of Borneo and the Cape York Peninsula of Australia. The few behavioral studies on this group suggest that antler flies have been evolved in the context of male intrasexual competition. Resource defense mating systems for this group have been described by [25], while [26] provided a review on current knowledge on the Phytalmiinae.

Trypetinae is a large subfamily that includes 19 known genera; *Carpomya, Cryptodacus, Goniglossum, Haywardina, Myiopardalis, Rhagoletis, Rhagoletotrypeta, Zonosemata, Acidia, Euleia, Strauzia, Trypeta, Anastrepha, Toxotrypana, Epochra, Paraterellia, Chetostoma, Oedicarena,* and *Myoleja*. The genera *Rhagoletis, Anastrepha,* and *Toxotrypana* include several species of major economic importance. While members of *Rhagoletis* are both holarctic and neotropical in distribution, *Anastrepha,* and *Toxotrypana* are restricted to the new world, while the rest of Trypetinae is especially diverse in the old-world tropics, which may be the center of origin [27]. Within Trypetinae, the subtribe Trypetina contains all the known leaf-mining species of tephritids, along with others by different larval feeding habits. A comprehensive account of this group of flies is provided by [28].

Tephritinae is considered the most specialized subfamily of Tephritidae. It is composed of six tribes with over 210 genera. Most species of Trypetini breed in flower heads, or form flower, stem, or root galls in plants of the family Asteraceae. Due to this habit, many of these tephritids have been used in biological control of weeds [29, 30]. Sexual behavior and biology of some members of this subfamily have been reviewed by previous authors [31].

Dacinae is a subfamily that contains only three genera *Bactrocera, Dacus,* and *Ceratitis* all of which include many species of major economic importance. All members of this subfamily are native to the Old World, despite the fact that the Mediterranean fruit fly, *C. capitata* has been established in South and Central America since the beginning of the 20th century, and there have been recurrent introductions and eradication efforts of this pest along with the Olive fruit fly*, B. oleae,* the Oriental fruit fly, *B. dorsalis* and others in North America.

#### **2.4 True fruit flies**

The term "fruit fly" is sometimes used for two distantly related groups of flies, namely the families Drosophilidae and Tephritidae (**Figure 4**). The Drosophilidae includes "fruit flies" of the geneticists, which are in reality, micro-fungi feeders that have acquired this name because of their habit of feeding on decaying fruit [32]. The Tephritidae is generally described to include the "true fruit flies" because most species attack living plant material, and an estimated 40% of the over 5000 described species attack intact and growing fruits. Females of fruit flies have an ovipositor, similar to the "sting" of a wasp, with which they puncture the skin of healthy fruits and lay their eggs therein. Larval development is completed within the fruit (which may become rotten as a result) and the fully-grown larvae then drop into the soil and form a puparium.

There are about 150 genera and 950 species of Tephritid fruit flies known in tropical Africa, most of which form a natural component of Africa's rich and varied

**151**

affected.

**Figure 4.**

**3.1 The adult**

significance [36–38].

margins of the third abdominal tergite.

**3. Functional morphology of Diptera**

*True fruit flies (Diptera: Tephritidae).*

group's developmental biology are considered here.

*Phylogeny and Functional Morphology of Diptera (Flies) DOI: http://dx.doi.org/10.5772/intechopen.90421*

biodiversity. About 70 species of fruit flies are considered important agricultural pests, and many others are minor or potential pests. Fruits and vegetables are the most important crops attacked [33–35], even though some seed crops are also

Morphology and anatomy of the different life stages of fruit flies, particularly their characters useful for taxonomic purposes, have been described in detail by previous authors. But only those aspects relevant for an understanding of the

Adults of Diptera have segmented bodies that include a head, thorax, and abdomen. The head has large eyes that cover the sides of the head with a small space between them in the front of the head. This helps them to see a wider area as they are flying (**Figure 5**). The adult body coloration of different dipteran species however varies from black through various shades of brown to orange or yellow. Yellow marks, particularly on the thorax, give many species a somewhat wasp-like appearance. This resemblance is particularly pronounced in certain *Bactrocera* subgenera and *Callantra* spp., which have petiolate abdomens, heavily fuscated costal stripes on the wings, and a jerky, wasp-like walk. The paired antennae each consist of three segments. Scanning electron microscope studies on *B. oleae* and *B. tryoni* indicate that the outer segment is covered with long cuticular spines interspersed with large numbers of chemosensilla of several distinct morphological types and functional

The general structure of dacine flies is fairly typical of cyclorrhaphan Diptera. Male dacines, except those of some groups such as Gymnodacus, typically have a pair of combs (or pectins) comprised of stiff curved bristles on the lateral hind

These combs function as stridulatory organs during courtship. Both sexes have a pair of tergal glands (ceromae) that open onto the surface of the fifth tergite. These

*Phylogeny and Functional Morphology of Diptera (Flies) DOI: http://dx.doi.org/10.5772/intechopen.90421*

*Life Cycle and Development of Diptera*

Phytalmiinae.

of flies is provided by [28].

**2.4 True fruit flies**

have been reviewed by previous authors [31].

drop into the soil and form a puparium.

speciation, as much as more than one species can cohabit the same plant, exploiting

Phytalmiinae is a subfamily comprising six genera; *Diplochorda, Ortaloptera, Phytalmia, Sessilinia, Tetrastiomyia,* and *Sophiria.* These are the flies with antennalike head projections and sometimes referred to as antler flies or deer flies (not to be confused with Tabanidae). Decaying plant material is the larval food across this subfamily. All described species of antler flies occur between the island of Borneo and the Cape York Peninsula of Australia. The few behavioral studies on this group suggest that antler flies have been evolved in the context of male intrasexual competition. Resource defense mating systems for this group have been described by [25], while [26] provided a review on current knowledge on the

different parts of it and exhibiting complex courtship behaviors [24].

Trypetinae is a large subfamily that includes 19 known genera; *Carpomya, Cryptodacus, Goniglossum, Haywardina, Myiopardalis, Rhagoletis, Rhagoletotrypeta, Zonosemata, Acidia, Euleia, Strauzia, Trypeta, Anastrepha, Toxotrypana, Epochra, Paraterellia, Chetostoma, Oedicarena,* and *Myoleja*. The genera *Rhagoletis, Anastrepha,* and *Toxotrypana* include several species of major economic importance. While members of *Rhagoletis* are both holarctic and neotropical in distribution, *Anastrepha,* and *Toxotrypana* are restricted to the new world, while the rest of Trypetinae is especially diverse in the old-world tropics, which may be the center of origin [27]. Within Trypetinae, the subtribe Trypetina contains all the known leaf-mining species of tephritids, along with others by different larval feeding habits. A comprehensive account of this group

Tephritinae is considered the most specialized subfamily of Tephritidae. It is composed of six tribes with over 210 genera. Most species of Trypetini breed in flower heads, or form flower, stem, or root galls in plants of the family Asteraceae. Due to this habit, many of these tephritids have been used in biological control of weeds [29, 30]. Sexual behavior and biology of some members of this subfamily

Dacinae is a subfamily that contains only three genera *Bactrocera, Dacus,* and *Ceratitis* all of which include many species of major economic importance. All members of this subfamily are native to the Old World, despite the fact that the Mediterranean fruit fly, *C. capitata* has been established in South and Central America since the beginning of the 20th century, and there have been recurrent introductions and eradication efforts of this pest along with the Olive fruit fly*,* 

The term "fruit fly" is sometimes used for two distantly related groups of flies, namely the families Drosophilidae and Tephritidae (**Figure 4**). The Drosophilidae includes "fruit flies" of the geneticists, which are in reality, micro-fungi feeders that have acquired this name because of their habit of feeding on decaying fruit [32]. The Tephritidae is generally described to include the "true fruit flies" because most species attack living plant material, and an estimated 40% of the over 5000 described species attack intact and growing fruits. Females of fruit flies have an ovipositor, similar to the "sting" of a wasp, with which they puncture the skin of healthy fruits and lay their eggs therein. Larval development is completed within the fruit (which may become rotten as a result) and the fully-grown larvae then

There are about 150 genera and 950 species of Tephritid fruit flies known in tropical Africa, most of which form a natural component of Africa's rich and varied

*B. oleae,* the Oriental fruit fly, *B. dorsalis* and others in North America.

**150**

#### **Figure 4.** *True fruit flies (Diptera: Tephritidae).*

biodiversity. About 70 species of fruit flies are considered important agricultural pests, and many others are minor or potential pests. Fruits and vegetables are the most important crops attacked [33–35], even though some seed crops are also affected.
