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Phylogeny of the Anastrepha daciformis species group

[add info from McPheron et al.; add images and trees and modify tables; replace citations with links]

Relationships among the species groups of Anastrepha were discussed by Norrbom et al. (1999) and McPheron et al. (1999)  (see Phylogeny of Anastrepha and Toxotrypana).  The relationships among the species of the daciformis group were analyzed by Norrbom (1998).  The following text is taken from these three references.  

The daciformis species group includes some of the most distinctive species of Anastrepha. All 13 species have dark brown markings of some type, and all but three have an uninterrupted marginal wing band, presumably for mimicry of vespid or other wasps. Four species in this group were placed in the subgenus or genus Pseudodacus by some authors (e.g., Hendel 1914a,b, Stone 1939, Blanchard 1961, Foote 1967), but Steyskal (1977a) synonymized this name with Anastrepha based on discovery of a fifth species intermediate in some of the characters used to differentiate Pseudodacus. Additional intermediate species were described by Norrbom (1998), lending further support to Steyskal’s decision.

The daciformis group is widespread. The overall range of the group extends from the West Indies and southern Texas to Argentina. Anastrepha antilliensis, murrayi, maculata and stonei are endemic to the Antilles. Anastrepha pallens is Mesoamerican, the daciformis complex is restricted to South America, and the macrura complex includes three Mesoamerican and two South American species. The known host plants (for 3 of the 13 spp.) are Sapotaceae. The larvae feed on the seeds.

The A. daciformis species group appears to be the sister group of the A. dentata species group, as indicated by the following synapomorphies: 1) phallus short, less than 1.25 mm long, and glans absent (Fig. 9E); 2) aculeus extremely slender (Fig. 16C-D), less than 0.05 mm wide except at base (reduction in aculeus width has occurred in certain other Anastrepha species, but whether this is convergence or a synapomorphy for some of these species and the daciformis + dentata groups remains uncertain); and 3) spermathecae weakly sclerotized. Norrbom (1998) suggested another possible synapomorphy: Third instar larva with hairs of hind spiracle relatively short, but this character is known for only one species in each group, A. pallens and A. sagittata (Baker et al. 1944, Phillips 1946), and these hairs are also short in Toxotrypana, and relatively short in A. interrupta (spatulata group) and A. limae (pseudoparallela group) (Baker et al. 1944, Steck & Wharton 1988), so this character may be homoplasious or have evolved at a lower level. The daciformis and dentata groups share one additional apomorphy: Lateral surstylus very short and rounded, barely extended beyond the prensisetae (Fig. 9F, H)), and subepandrial sclerite relatively posterior in position, in lateral view its apex usually at posterior margin of epandrium (Fig. 9E, G). But here again, the lateral surstylus is also rather short in Toxotrypana, the A. robusta group (especially in A. binodosa), and to a lesser extent in the A. punctata, leptozona and schausi groups, so it could possibly be a synapomorphy for some or all of these groups in addition to the A. daciformis and dentata groups.

The daciformis group is probably the most clearly monophyletic species group of Anastrepha, as indicated by the following unique apomorphies: 1) scutellum bicolored, with at least basal third on sides and dorsum distinctly darker than apex, and with basal scutellar seta within darker area (Fig. 1B), except usually in A. avispa (scutellar markings are present on only a few other Anastrepha species, and in different patterns, suggesting they are not homologous); 2) eversible membrane with unique pattern of dorsobasal teeth (Fig. 13B), small and weakly sclerotized except for medially interrupted apical row of large, strongly sclerotized, hook-like teeth; 3) spermathecae membranous; and, 4) phallus extremely short, less than 0.30 mm long (Fig. 9E).

Relationships within the daciformis group were analyzed by Norrbom (1998) using PAUP version 3.1.1. The characters used are listed in Table 3, and the character state distributions are shown in Table 4. The outgroup used for determining character polarities included Anastrepha sagittata (Stone), which belongs to the dentata species group (the hypothesized sister group of the daciformis group), and A. obliqua (Macquart), a more distantly related member of the genus. The heuristic search option was used, with stepwise and random addition yielding the same set of six trees of 76 steps (consistency index excluding uniformative characters = 0.651, retention index = 0.783). Two of these trees are shown in Figs. 3-4. They show the accelerated transformation character optimization, but the delayed transformation optimization did not produce trees differing in topology (i.e., the relationships of the species). The following clades were always consistent: maculata + stonei; the daciformis complex (with katiyari as the sister taxon of castanea + daciformis); and the macrura complex (aquila, avispa, bicolor, macrura, and zucchii, with the latter two always clustered and bicolor the basal taxon). The species with wasp mimicry wing patterns (the daciformis and macrura complexes) together are the sister group of A. pallens. The major differences among the six trees involved the placement of antilliensis, murrayi, and maculata + stonei, which either arise as the basal clades in that order (Fig. 3), or together form a monophyletic group that is the sister group of the rest of the daciformis group (Fig. 4). The other variation occurred within the macrura complex; avispa is either the sister taxon of aquila, in a trichotomy with aquila and macrura + zucchii, or is the sister taxon of all three of those species.

Table 3. Characters used in phylogenetic analysis of the daciformis group.

1. Predominant body color, except for pale areas and brown spots - 0) yellow to orange-brown; 1) red-brown to dark brown.

2. Frons color - 0) with lateral brown spot along eye margin on dorsal part; 1) unicolorous, except ocellar tubercle brown; 2) with transverse brown band or spot including ocellar spot; 3) with transverse brown band, at least sometimes with lateral anterior extension, broadest along eye margin. Brown markings may be reduced or absent in teneral specimens. Coded as ordered.

3. Gena color - 0) unicolorous; 1) with dark spot below eye (may be faint or absent in teneral specimens).

4. Occiput color - 0) entirely yellow; 1) with brown stripes or triangular marks on or near sutures of median occipital sclerite; 2) mostly brown except anterior margin. Coded as ordered.

5. Number of orbital setae - 0) 2; 1) 1, posterior orbital seta absent. This occurs as homoplasy in various other species of Anastrepha, e.g., see Norrbom (1991).

6. Antenna length - 0) not elongate nor extended to level of ventral margin of face, first flagellomere < 3 times as long as wide (measured on mesal side); 1) elongate, extended to or beyond level of ventral margin of face, first flagellomere usually > 3 times as long as wide.

7. Mesonotum with dark brown markings - 0) scutum and scutellum yellow to dark brown except for pale stripes and apex of scutellum (nonpale areas unicolorous); 1) scutum with triangular brown to dark brown markings, sometimes connected, near posterior margin and scutellum with lateral brown spots or transverse brown band at border of orange and white areas; 2) scutellum with black spots at border of orange and white areas, and scutum usually with dark brown spot mesal to postalar seta, sometimes with additional spots; 3) entire base of scutellum and large part of scutum black. Coded as unordered.

8. Mesonotal pale lateral presutural stripe, middle part (on scutum) - 0) complete; 1) narrowly interrupted; 2) absent. May be poorly differentiated in predominantly yellow to orange species. Coded as ordered.

9. Mesonotal pale lateral presutural stripe, posterior part (on posterior part of notopleuron) - 0) absent; 1) present. May be poorly differentiated in predominantly yellow to orange species.

10. Mesonotal pale presutural dorsocentral stripe - 0) absent; 1) present and connected anteriorly with pale area on postpronotal lobe; 2) separated anteriorly from pale area on postpronotal lobe. This character was difficult to observe for sagittata and antilliensis, as the surrounding area is pale yellow in these species, but the stripe appears to be absent. Coded as unordered.

11. Scutal microtrichia pattern - 0) entirely microtrichose, or at most with medial presutural bare area; 1) nonmicrotrichose medially, lateral and posterior margins microtrichose; 2) entirely or almost entirely nonmicrotrichose. Coded as unordered.

12. Scutellar microtrichia pattern - 0) disc entirely microtrichose; 1) disc mostly or entirely without microtrichia.

13. Scutellum color - 0) unicolorous or with dark markings only on extreme base; 1) bicolored, with at least basal third on sides and dorsum distinctly darker than apex.

14. Propleuron color - 0) yellow; 1) at least partly dark orange or brown.

15. Mesopleuron with dark brown spots - 0) without dark brown spots; 1) with small dark brown spot on anepimeron; 2) with small dark brown spots on anepisternum, katepisternum and anepimeron. Coded as ordered.

16. Cell r₁ with hyaline spot - 0) with hyaline spot or band at apex of R₁ (usually small in pallens); 1) without hyaline spot at apex of R₁, costal and S-bands completely fused in this area.

17. Costal band in cell br - 0) broadly extended to vein M along cell bm; 1) not extended to vein M.

18. Costal band in cell r₂₊₃ - 0) covering base of cell; 1) not covering base of cell.

19. S-band, basal section - 0) without posterior extension toward wing margin in cell a₁; 1) with posterior extension to or towards wing margin in cell a₁. The extension is not as broad or long in some specimens of stonei and is not fully extended to the margin in maculata, but I tentatively coded these species state 1.

20. S-band, basal section - 0) extended along vein Cu₁ beyond bm-cu, but no more than halfway to dm-cu, and cell bcu evenly infuscated; 1) not extended along vein Cu₁ beyond bm-cu, and posterior third or more of cell bcu hyaline or very faintly infuscated; 2) extended along vein Cu₁ to dm-cu, and cell bcu evenly infuscated. In teneral specimens of pallens and stonei, the band sometimes does not appear to extend along Cu₁ beyond bm-cu, but specimens with the wing pattern fully developed are clearly state 0, and I therefore coded these species 0. Coded as unordered.

21. S-band, middle section - 0) continuous; 1) separated from basal and apical sections; 2) absent. In bicolor the basal section is sometimes faintly extended into cell dm, but never connects with the apical section; I coded it state 0. Coded as ordered.

22. S-band, apical section width - 0) narrow or slightly broadened, apical half of cell r₂₊₃ with large hyaline area; 1) very broad, all of cell r₂₊₃ infuscated.

23. V-band, proximal arm - 0) complete, extended to vein R₄₊₅; 1) reduced, extended to or slightly anterior to vein M; 2) reduced and paler than costal band or absent. Coded as ordered.

24. V-band, distal arm - 0) present; 1) absent.

25. Medial vein ratio (distance along M from bm-cu to r-m/ distance from r-m to dm-cu) - 0) average greater than 1.5; 1) average less than 1.5. See Table for comparison of values.

26. Abdominal pattern - 0) banded (often poorly differentiated in species with predominantly yellow to orange-brown abdomens); 1) intermediate between banded and with T-shaped mark; 2) brown with T-shaped mark. Coded as ordered.

27. Outer surstylus shape - 0) long, somewhat truncate; 1) very short and rounded, extended only slightly beyond prensisetae.

28. Aedeagus - 0) longer than female oviscape, distiphallus present; 1) short, less than 1.3 mm long, distiphallus absent; 2) extremely short, less than 0.30 mm long, distiphallus absent. Coded as ordered.

29. Eversible membrane, pattern of dorsobasal scales - 0) at least 2 rows of similarly well sclerotized scales; 1) all scales weakly sclerotized; 2) weakly sclerotized except medially interrupted row of large, strongly sclerotized, scales. Coded as unordered.

30. Spermathecae - 0) moderately sclerotized; 1) weakly sclerotized; 2) membranous. Coded as ordered.

Table 4. Character state distributions in species of the daciformis group.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30

obliqua 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

sagittata 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1

antilliensis 0 1 0 0 0 0 3 0 0 0 2 1 1 0 1 0 0 0 1 0 0 0 0 1 0 0 1 2 2 2

murrayi 0 1 0 0 1 0 3 0 0 1 2 1 1 0 0 0 0 0 1 0 0 0 0 1 0 0 ? ? 2 ?

stonei 0 0 1 0 1 0 2 0 1 1 2 1 1 0 2 1 0 1 1 0 0-1 0 0 0 1 0 1 2 2 2

maculata 0 1 0 0 1 0 2 0 1 1 0 1 1 0 2 1 0 0 1 0 1 0 0 0 1 0 1 2 2 2

pallens 0 2 1 0 1 0 1 0 1 1 2 0 1 0 0 0 0 1 0 0 1 0 0 1 0 0 1 2 2 2

aquila 1 2 1 1 1 1 0 0-1 1 1 0 0 1 0 0 1 0 0 0 2 2 1 2 1 0 1 1 2 2 2

avispa 1 2 1 0-1 1 1 0 0 1 1 1 0 1 0 0 1 0 0 0 2 2 1 2 1 0 1 1 2 2 2

bicolor 1 2 1 0 1 1 0 0 1 1 1 0 1 0 0 1 1 0 0 0 2 1 2 1 0 0 1 2 2 2

macrura 1 2 1 1 1 1 0 2 1 1-2 0 0 1 1 0 1 1 0 0 0 2 1 2 1 0 2 1 2 2 2

zucchii 1 2 1 2 1 1 0 1 0 0 1 0 1 1 0 1 0 0 0 2 2 1 2 1 0 2 1 2 2 2

castanea 1 3 1 2 1 1 0 2 1 2 0 0 1 1 0 1 1 1 0 1 2 0 1 1 1 0 1 2 2 ?

daciformis 1 3 1 2 1 1 0 0 1 2 0 0 1 1 0 1 1 1 0 1 2 0 1 1 0 0 1 2 2 2

katiyari 1 3 1 1 0-1 1 0 0 1 1 1 0 1 0 0 1 1 1 0 1 2 0 1 1 1 0 1 2 2 2

References
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Baker, A. C., W. E. Stone, C. C. Plummer & M. McPhail. 1944. A review of studies on the Mexican fruitfly and related Mexican species. U.S. Dep. Agric. Misc. Publ. 531: 155 p.
Foote, R.H. 1967. Family Tephritidae (Trypetidae, Trupaneidae), Fasc. 57, 91 p. In: N. Papavero, ed., A catalogue of the Diptera of the Americas south of the United States. Departamento de Zoologia, Secretaria da Agricultura, São Paulo.
Hendel, F. 1914a. Analytische Übersicht über die Anastrepha-Arten (Dipt.). Wien. Entomol. Ztg. 33: 66-70.
Hendel, F. 1914b. Die Bohrfliegen Südamerikas. Übersicht und Katalog der bisher aus der neotropischen Region beschrieben Tephritinen. Abh. Ber. K. Zool. Anthrop. Ethnogr. Mus. (1912) 14: 1-84.
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. 1998. A revision of the Anastrepha daciformis species group (Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 100: 160-192.
Norrbom, A.L., R.A. Zucchi & V. Hernández-Ortiz. 1999. Phylogeny of the genera Anastrepha and Toxtrypana (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.
Phillips, V. T. 1946. The biology and identification of trypetid larvae (Diptera: Trypetidae). Mem. Am. Entomol. Soc. 12: 161 p.
Steck, G. J. & R. A. Wharton. 1988. Description of immature stages of Anastrepha interrupta, A. limae, and A. grandis (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 81: 994-1003.
Steyskal, G.C. 1977a. Two new neotropical fruitflies of the genus Anastrepha, with notes on generic synonymy (Diptera, Tephritidae). Proc. Entomol. Soc. Wash. 79: 75-81.
Stone, A. 1939. A revision of the genus Pseudodacus Hendel (Dipt. Trypetidae). Rev. Entomol. (Rio J.) 10: 282-289.