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Fruit Fly (Diptera: Tephritidae) Phylogeny
Although much remains unknown about fruit fly phylogeny, many groups of Tephritidae have been analyzed to at least some degree, especially during the past decade, when rigorous cladistic studies and molecular methods were first utilized within the family. Much additional work is needed in this area, however, as the relationships among many higher taxa are poorly resolved, and many large taxa have had limited or no analysis.
At the family level, Hennig (1958, 1973), Griffiths 1972, J.F. McAlpine (1977, 1989) and Korneyev (1992, 1999a) discussed the relationships of the Tephritidae to other families of Tephritoidea, which also includes the families Lonchaeidae, Pallopteridae, Piophilidae, Platystomatidae, Pyrgotidae, Richardiidae, and Otitidae (= Ulidiidae). The Tachiniscinae, long ranked as a family, were included in the Tephritidae by Korneyev (1999a,b). Also see Classification & Diversity.
Although there is some disagreement among the above-named authorities on the relationships of the families within the Tephritoidea, there is consensus that the Ulidiidae, Platystomatidae, Pyrgotidae and Tephritidae (including Tachiniscinae) are a monophyletic group. McAlpine (1989) and Korneyev (1999a), who termed these four families the "Higher Tephritoidea", indicated that they are a well defined monophyletic group and discussed the synapomorphies supporting this hypothesis of relationship. Within this group, the Platystomatidae, Pyrgotidae and Tephritidae are more closely related (Griffiths 1972, Korneyev 1999a). In all three of these families, the phallus of the male is coiled at rest dorsal to the postabdomen in a space between it and tergite 5 (the plesiomorphic condition being coiled ventrally in front of the epandrium and surstyli). Korneyev (1999a) discussed a number of other synapomorphies supporting the relationship of these three families. Relationships among them are not well resolved. Aczél (in Hardy 1957) believed that the Pyrgotidae and Tephritidae arose from the same ancestors, and J.F. McAlpine (1989) suggested that the Pyrgotidae + Tachiniscidae are the sister group of the Tephritidae, whereas Korneyev (1992) proposed that the Platystomatidae and Tephritidae are sister taxa. In the most thorough analysis, Korneyev (1999a) considered the Pyrgotidae and Tephritidae the most likely of the three families to be sister taxa.
Within the family Tephritidae, phylogenetic studies have been conducted on a wide number of taxa. The methods and rigor of the analyses have varied from intuitive discussions of the monophyly of certain groups and likely autapomorphies (e.g., Korneyev 1994, 1996, 1999b, Foote et al. 1993) to comprehensive analyses of large taxa using computer algorithms and rigorous outgroup testing of character polarities (Han 1999, Merz 1999, Freidberg & Norrbom 1999). Molecular analyses have been conducted on several taxa (Han & McPheron et al. 1999, Smith & Bush 1999, McPheron et al. 1999, Han 2000, Smith et al. 2003), but have been limited to some degree by unavailability of certain genera or species. The following is a brief overview of relationships within the family. Additional information and references can be found in Norrbom et al. (1999) and Korneyev (1999b,c).
Relationships among the subfamilies and tribes of Tephritidae were analyzed by Korneyev (1999b,c) based on morphology, and by Han & McPheron (1994, 1997, 1999) based on DNA sequence analysis. Hancock (1986) also proposed a phylogeny that included most tribes of Trypetinae and Phytalmiinae, but some of the lower clades are not supported by apomorphic characters. Despite the extensive efforts of these and other authors, relationships among the six subfamilies recognized by Korneyev (1999b) and the monophyly of some of them, particularly the Trypetinae, remain uncertain. The Tachiniscinae, Blepharoneurinae and Phytalmiinae are believed to be the more basal clades within the family.
The small subfamily Tachiniscinae is well supported as monophyletic by female genitalic characters, including a dorsoapical opening on the oviscape and a unique ventral cluster of scales on the eversible membrane (Korneyev 1999b). Relationships within the subfamily were discussed by Korneyev (1999b), and Norrbom (1994) analyzed the relationships among four of the five known genera of the tribe Ortalotrypetini, including the fossil genus Protortalotrypeta. The biology of the Tachiniscinae is poorly known. The only reared species is a parasitoid of a moth.
The small subfamily Blepharoneurinae is also clearly monophyletic, except perhaps the genus Ceratodacus. All of the genera have a distinctive anepisternal seta just anterior to the phragma, and the three largest genera have modified pseudotracheal ring tips on their labella (Munro 1957, Driscoll & Condon 1994). The relationships among the five included genera were analyzed by Norrbom & Condon (1999). The three genera of known biology are phytophagous and all of the known host plants are in the family Cucurbitaceae.
According to Korneyev (1999b), support for the monophyly of the Phytalmiinae includes: shape of the male terminalia (epandrium elongate dorsoventrally, lateral surstylus without anaterior lobe, medial surstylus narrow and weak (secondarily modified in Clusiosoma group)); hypandrium with lateral sclerites rudimentary, vanes of phallapodeme articulated with anterior end of hypandrium; and abdominal sternites 4-6 without anterior apodeme. The ocellar seta small or rudimentary is probably another synapomorphy. Phytalmiinae also share the following symplesiomorphies: aculeus usually "tactile", tip not completely fused to shaft; scutellum with 3 pairs of setae (in groundplan, reduced to 2 in some taxa); eversible membrane with no enlarged dentiform scales (in groundplan); and 3 spermathecae without surface denticles. Relationships within the subfamily Phytalmiinae were discussed by Korneyev (1999b), but based on a limited number of genera. The smaller tribes Phytalmiini, Epacrocerini, and Phascini appear to be well defined, but relationships within the Acanthonevrini remain poorly understood beyond the preliminary hypotheses proposed by Korneyev. He defined the Acanthonevrini on the basis of one synapomorphy: the sclerotized medioapical lobe of the oviscape is desclerotized medially and has a somewhat W-shaped appearance. Most genera of Phytalmiinae are saprophagous, although some are restricted to recently damaged or dead tissues of limited numbers of plant taxa.
The subfamilies Trypetinae, Dacinae and Tephritinae probably form a monophyletic group, but Korneyev (1999b) indicated that none of the character states likely to be synapomorphies are shared by all of the included genera. Whether this homoplasy is due to character reversal or convergence needs to be resolved to better determine the true relationships of these taxa.
The monophyly of the subfamily Dacinae is supported by morphological and biochemical data, although the important characters have been studied in few, or in some cases, no representatives of the tribe Gastrozonini. Hancock (1986) and Foote et al. (1993) suggested the monophyly of the Dacinae based on the shape of the lobe of cell bcu (often narrower at base than medially), spermathecae number reduced to two (considered a synapomorphy of Dacini + Tephritinae by Hancock 1986), and surstyli shape. Korneyev (1999b) also considered the large proctiger of the male to be a synapomorphy of the subfamily. Species of Dacini and Ceratitidini whose larvae have been described (no larvae of Gastrozonini have been described) have a ridge across the large caudal tubercle below the hind spiracle, an apomorphic state not reported in any other Tephritidae (Carroll 1992). Kitto (1983) suggested the close relationship of the few Dacini and Ceratitidini included in his immunological study (no Gastrozonini were studied), and the four species (2 Dacini, 1 Ceratitidini, 1 Gastrozonini) included by Han & McPheron (1997) were grouped in their neighbor-joining tree. Smith et al. (2002) provided better molecular support for the monophyly of the subfamily; the 16 species in four genera (at least one from each tribe) included in their study formed a well supported clade.
Relationships among the genera of Dacinae have not been rigorously analyzed. The Dacini and Ceratitidini each appear to be monophyletic, but the Gastrozonini may be paraphyletic (Norrbom et al. 1999, Korneyev 1999b). The Dacini and Ceratitidini breed in fruits or flowers, whereas many genera of Gastrozonini breed in shoots of bamboo or grasses (Hancock 1999, Hancock & Drew 1999). Norrbom et al. (1999) discussed the possible relationships among the three genera of Dacini, and Drew & Hancock (1999) discussed the possible relationships among the many subgenera of this group. White (1999) conducted a preliminary cladistic analysis of this group. Within the Ceratitidini, De Meyer (1999) analyzed relationships within the genus Ceratitis. Relationships among some species of Bactrocera were analyzed by Muraji & Nakahara (2002) and Smith et al. (2003).
The subfamily Trypetinae may be paraphyletic. Korneyev (1999b) suggested that based mainly on larval similarities, the tribes Trypetini and Zaceratini may be more closely related to the Tephritinae than to other Trypetinae. The relationship of the latter tribe to the Tephritinae is supported by morphological (the oval shape of the epandrium and surstyli) and molecular data (Korneyev 1999b, Han 1999).
The tribe Adramini has been defined mainly by a single synapomorphy, the presence of fine hairs on the anatergite (Korneyev 1994). Korneyev (1999b) discussed the possible relationships among many of the included genera.
The tribe Carpomyini, especially the subtribe Carpomyina, is among the best studied large groups of Tephritidae in terms of phylogenetic analysis. The subtribes Notommatina and Paraterelliina, which each include two genera, were included in the Carpomyini by Korneyev (1996), but he later considered their relationships to be less certain (Korneyev 1999b). Jenkins (1996) also discussed characters that support the monophyly of the Paraterelliina. Smith & Bush (1999) reviewed the known phylogenetic information for the Carpomyina and also reanalyzed morphological and molecular data sets for some taxa. Possible synapomorphies for the subtribe include the weakly sclerotized apical area on the female oviscape, the shape of the subapical lobe of the glans and the shape of the surstyli (with some secondary modification) in the male, and perhaps the presence of preoral teeth in the larvae (Norrbom 1989, Carroll 1992, Jenkins 1996, Smith & Bush 1999). The latter may be a synapomorphy for the entire Carpomyina or only some taxa within it. Jenkins (1996), McPheron & Han (1997), and Smith & Bush (1999) analyzed relationships among some included genera, and they suggested that the large genus Rhagoletis is paraphyletic. Relationships have been analyzed among the species within the following genera: Oedicarena (Norrbom et al. 1988), Cryptodacus, Haywardina and Rhagoletotrypeta (Norrbom 1994), and for many species of Rhagoletis (Jenkins 1996 and Smith & Bush 1999 using morphological data; Berlocher & Bush 1982 and Berlocher et al. 1993 using allozyme data; McPheron & Han 1997, Smith & Bush 1997, and Han & McPheron 1999 using DNA sequence data).
The tribe Rivelliomimini is a small monophyletic groups that is well defined by the following apomorphic characters: abdominal tergite 5 with a pair of black lateral swellings, and cell bcu acute apically but with vein Cu2 straight (Hancock 1986). Within the tribe Toxotrypanini, Anastrepha and Toxotrypana possess a number of synapomorphies involving the genitalia that clearly indicate their monophyly (Norrbom 1985, Hancock 1986, Norrbom et al. 1999). Hexachaeta is included mainly on the basis of molecular data (Han & McPheron 1997). Relationships within Anastrepha and Toxotrypana were analyzed based on morphology by Norrbom et al. (1999) and based on DNA sequence data by McPheron et al. (1999).
Relationships among the subtribes of the tribe Trypetini were discussed by Korneyev, who placed the Acidoxanthina, Nitrariomyiina, and Chetostomatina in a separate tribe Nitrariomyiini. The relationships of the Chetostomatina and Trypetina were analyzed by Han (1992, 1999) based on morphological data, and by Han (2000) and Han & Ro (2002) based on molecular data. Within the Trypetina, intrageneric analyses have been published for Cornutrypeta (Han et al. 1993) and Vidalia (Han et al. 1994, as Pseudina). The Trypetini include fruit and flower feeders as well as leaf and stem miners.
Relationships among the tribes of the subfamily Tephritinae were discussed by Korneyev (1999c). This group is probably monophyletic, although there is some homoplasy in both of the morphological synapomorphies listed by Korneyev: the apical part of the spermathecal duct is distinctly broader for a length at least equal to that of the spermatheca (Norrbom et al. 1999), although this character state occurs within the Ceratitidini and Gastrozonini; and the presence of pale, lanceolate setulae, which are common although not consistent in all Tephritinae. Female heterogamety could be another synapomorphy for the Tephritinae, and occurs at least in the Cecidocharini, Dithrycini, Eutretini and Tephritini (Bush 1966, Kitto 1983, Frias 1992, Foote et al. 1993), but representatives of the other tribes need to be studied. An analysis of DNA sequence data by Han & McPheron (1994) also strongly supported the monophyly of the Tephritinae, although no representatives of the Acrotaeniini, Schistopterini or Tephrellini were included in their study. The monophyly of the Tephritinae is further supported by the biology of the group. All Tephritinae breed in plants of the family Asteraceae as flower feeders, gall formers or stem miners, except for the tribe Tephrellini and a few species of Oedaspis (Dithrycini) and Eutreta (Eutretini) whose biologies are secondary shifts (Korneyev 1999c).
Korneyev (1999c) divided the Tephritinae into two groups: the Higher Tephritinae, which he considered monophyletic, including the tribes Dithrycini, Eutretini, Schistopterini, Tephrellini and Tephritinae; and the Lower Tephritinae, which is probably paraphyletic, including the tribes Terelliini, part of the Acrotaeniini (Tomoplagia group), Noeetini, Xyphosiini, Myopitini, and Cecidocharini. He placed most of the other genera of the Acrotaeniini within the Eutretini. This division of the subfamily was based mainly on the shape and color of the lateral vertical seta and the postocular setulae. Pending further testing of this classification, it has not been fully adopted here, as both of the preceeding characters vary more within the Acrotaeniini and Cecidocharini than was known to Korneyev.
Korneyev (1999c) considered the tribe Terelliini to be the sister group of the rest of the Tephritinae. He provided a detailed analysis of relationships among the genera, including various species groups within the probably paraphyletic genus Terellia. The hypothesis that the tribe is monophyletic is supported by two synapomorphies: the inclinate posterior orbital seta; and the lyre-shaped pattern on the scutum (Freidberg 1985), although Korneyev (1999c) questioned whether they might be convergent in the genus Neaspilota which lacks certain plesiomorphic genitalic character states. Relationships among the species of Craspedoxantha were analyzed by Freidberg (1985) and Freidberg & Mathis (1990).
The Acrotaeniini, as noted above, may include two unrelated groups of genera, and its monophyly needs to be further tested. Relationships within the genus Polionota were analyzed by Norrbom (1988).
The small tribe Xyphosiini was redefined by Korneyev (1999c). Some of the New World taxa currently included may belong elsewhere. Relationships among the species of Gymnocarena were analyzed by Norrbom (1992).
Korneyev (1999c) analyzed relationships among the genera of the tribe Noeetini. The barbed shape of the aculeus tip is presumed to be a synapomorphy, although it is secondarily lost in one species of Ensina. The long, slender shape of the acrophallus of the glans is a synapomorphy of the genera exclusive of Ensina. All species of Noeetini whose biology is known breed in flowers of Lactuceae (Asteraceae).
Relationships among the genera of Myopitini were analyzed by Freidberg & Norrbom (1999). This group is defined by the loss of the posterior orbital seta and the lobe on cell bcu (vein Cu2 is straight or convex). Each of these apomorphic character states occurs sporadically and independently in other Tephritinae, but this is hypothesized to be homoplasy.
The tribe Cecidocharini is defined by the wing pattern banded (plesiomorphic), scutellum swollen, and usually the mesonotum with white, lanceolate setulae in distinctive patterns, often in clusters (Foote et al. 1993). Korneyev (1999c) suggested that several of the smaller genera may belong in the Higher Tephritinae, but their relationships need further study.
The tribe Dithrycini was redefined by Korneyev (1999c) to include species with an unusual pattern of scales of the eversible membrane. Most included species also have a swollen scutellum and most are gall-formers. Korneyev (1999c) also conducted a preliminary analysis of relationships among the subtribes and many of the genera.
The tribe Eutretini may be a poly- or paraphyletic group and needs comprehensive analysis. Various characters used to delimit it, such as the usual presence of a parafacial spot, large palpus, eye banded or spotted, and erect setae on the scutellum occur sporadically in other Tephritinae and are absent in some included taxa. Freidberg & Kaplan (1993), Foote et al. (1993) and Korneyev (1999c) discussed possible relationships among some of the included genera. The tribe Schistopterini, which shares many characters with the Eutretini, may be related to part or all of that tribe. Schistopterini have one or more of the following apomorphic traits: preocellar setae; eye banded or spotted (Munro 1926); erect white setulae on disc of scutellum; a distinctive pattern of wing bullae; and/or a strong incision in the costal margin of the wing at the apex of the subcostal vein (Hancock 1986), and the distiphallus small, with little sclerotization (Norrbom et al. 1999).
The tribe Tephrellini has not been comprehensively analyzed. It has been defined partially by host relationships (the larvae breed in seed capsules of nonasteraceous plants - Acanthaceae, Lamiaceae, Verbenaceae), although most species have a typically shaped aculeus tip (very slender, with a slight, broad constriction well before apex), and a shining or sparsely mictrotrichose abdomen, which may be synapomorphies (Freidberg & Kaplan 1993, Norrbom et al. 1999, Korneyev 1999c). On the other hand, Hancock (1990) argued that the Platensinina (his Platensina group) belong in the Tephritini based on thoracic microtrichia density and differences in wing patterns from the other Tephrellini. He also suggested that the presence of a costal band in cell c (versus a medial spot) in many genera of the Tephrellina indicates their close relationship.
The Tephritini, the largest tribe of Tephritidae, was considered monophyletic by Korneyev (1999c), although none of the four synapomophies he listed are exclusive to the group or present in all included taxa. Several genus groups have been recognized within the tribe (Norrbom et al. 1999, Korneyev 1999c). Merz (1999) analyzed relationships among the Palearctic and Afrotropical genera of the Tephritis group. Intrageneric analyses have been published for Cryptophorellia (Freidberg & Hancock 1989) and Euaresta (Norrbom 1993).
See the Fruit Fly Bibliography Database for full information for cited references.
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Content by Allen L. Norrbom. Last Updated: April 30, 2004.