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Phylogeny of Cryptodacus, Haywardina and Rhagoletotrypeta

Table 1. Characters studied in cladistic analysis. Unless otherwise indicated, state 0 is plesiomorphic and multistate characters are linear transformation series. See Table 2 for distribution of character states among species.

1. Frons color - 0) yellow except ocellar tubercle; 1) with brown mark extended laterally from ocellar tubercle.

2. Number of frontal setae - 0) 2-3; 1) 4 or more. R. striatella was coded 0 in the Hennig86 matrix because there are only 3 frontal setae in other species of the striatella group, so the occasional occurence of 4 in striatella is probably homoplasy.

3. Orbital ratio (distance between orbital setae/ distance from anterior seta to eye margin) - 0) < or = 1; 1) > 1.

4. Ocellar seta - 0) at least as large as posterior orbital seta; 1) smaller than posterior orbital seta.

5. Face color - 0) entirely yellow; 1) with 1-2 brown spots; 2) with 4 spots or 2 elongate spots; 3) with 3 spots arranged in inverted triangle or fused into U-shaped mark.

6. Facial ratio (medial height/ ventral width of face) - 0) at least 0.80; 1) less than 0.70.

7. First flagellomere length/width (lateral view) - 0) < 3.5; 1) > 4.0.

8. First flagellomere shape - 0) tapered, tip with distinct dorsoapical pointed lobe; 1) tapered, tip at most slightly flattened, rounded or with weak dorsoapical point; 2) not tapered, tip strongly flattened, rounded or slightly expanded, without dorsoapical point. With Paraterellia as outgroup, state 1 is plesiomorphic, with states 0 and 2 independently derived. With either Rhagoletis species as outgroup, state 0 is plesiomorphic.

9. First flagellomere color - 0) yellow; 1) at least tip brown.

10. Occiput and postgena color - 0) without brown areas; 1) with large brown area(s). The coding for R. annulata slightly effects tree topology (see discussion of annulata group).

11. Dorsocentral seta location - 0) closer to level of postalar seta than to level of postsutural supra-alar seta; 1) closer to level of postsutural supra-alar seta than level of postalar seta. With either Rhagoletis species as outgroup, state 1 is plesiomorphic.

12. Scutal microtrichia - 0) entirely bare; 1) evenly microtrichose; 2) with 4 stripes of more dense microtrichia; 3) with only 2 submedial denser stripes; 4) with submedial stripes and with sublateral bare areas. With Paraterellia as outgroup, state 1 is plesiomorphic; with either Rhagoletis species, state 2 is plesiomorphic.

13. Postpronotal lobe - 0) mostly or entirely white; 1) mostly or entirely brown. If R. xanthogastra is coded 0 vs. 1 in the Hennig86 matrix, the trees have the same topology but are 1 step shorter.

14. Notopleuron - 0) entirely brown or yellow; 1) with white posterior area.

15. Thorax color (excluding white markings) - 0) mostly yellow or orange; 1) mostly brown. With R. striatella as outgroup, state 1 is plesiomorphic.

16. Medial scutal white stripe or spot - 0) absent; 1) present. P. immaculata Blanc and varipennis (Coquillett) have a broad, faint, posterior white mark that probably is not homologous with state 1.

17. Lateral scutal white stripe - 0) absent; 1) present, at least as anterior or posterior spot; 2) slender, not extended along transverse suture. If Paraterellia is coded 0 vs. 1 in the Hennig86 matrix, the same trees result except they are 1 step longer, with 17.1 a basal synapomorphy.

18. Lateral scutal white stripe - 0) extended to transverse suture (or absent); 1) extended to postsutural supra-alar seta, but not to transverse suture; 2) reduced to white spot at or posterior to intra-alar seta. C. silvai was coded 1 in the Hennig86 matrix.

19. Postsutural scutal brown spots - 0) scutum brown or orange except for white areas, or with U-shaped brown spot extended from posterior margin; 1) scutum with isolated brown spot bordering lateral white stripe.

20. Anepisternum color (excluding white area) - 0) mostly brown or yellow (if yellow with brown spot, spot located medially or ventrally); 1) yellow with brown dorsal spot.

21. Anepisternum dorsal white area - 0) linear or narrowly triangular, entire dorsal margin, but no more than 1/2 of posterior margin white; 1) broadly triangular, entire dorsal margin and > 1/2 of posterior margin white; 2) anterior part of dorsal margin dark, most or all of posterior margin white. H. cuculi, which has brown spot extended to dorsal margin anteriorly, but white area narrow posteriorly, was coded 0.

22. Katepisternum dorsal margin - 0) brown or yellow; 1) with broad white spot or stripe.

23. Scutellum color - 0) white area small, basal seta in brown area on sides and disc; 1) white area large, basal seta not in brown areas on sides and disc; 2) sides with large brown basal area, but disc yellow basally and medially. State 1 is plesiomorphic, with states 0 and 2 derived independently.

24. Length of vein M from BM-Cu to R-M/ length from BM-Cu to DM-Cu - 0) < 0.60; 1) > 0.63.

25. Vein Cu1 with brown border between bm-cu and discal band - 0) no; 1) yes. If C. tau is coded 0 vs. 1 in the Hennig86 matrix, the trees have same topology but are 1 step longer. P. immaculata sometimes has a band along Cu1 that is doubtfully homologous with state 1.

26. Wing band width - 0) at least subbasal band and usually others moderately to very broad; 1) all bands narrow, including subbasal band in cells bm and bcu. P. superba Foote has narrow bands, but this is probably homoplasy.

27. Discal band - 0) not crossing DM-Cu; 1) crossing at least posterior end of DM-Cu.

28. Subapical band - 0) covering all of DM-Cu; 1) ending at anterior end of DM-Cu or in cell r4+5.

29. Subapical band - 0) as strong anteriorly as posteriorly; 1) faint or absent anterior to vein R4+5. Paraterellia was coded 0 in the Hennig86 matrix because this state is found in most Carpomyina.

30. Accessory costal band - 0) present; 1) absent. Coded state 0 in C. obliquus. State 1 is plesiomorphic with R. striatella as outgroup. Paraterellia was coded 0 because the anterior end of the discal band (the apical fork in P. ypsilon Foote) may be homologous with the accessory band. If it is coded state 1 (or the polarity is reversed), tree topology is not effected.

31. Abdomen color - 0) mostly orange, at least tergite 5 usually with paired basal submedial brown spots; 1) mostly orange, often with lateral brown spots; 2) mostly brown. Coded nonadditive. State 1 is plesiomorphic with R. ferruginea as outgroup; state 2 with R. striatella. The coding for R. pastranai slightly effects tree topology (see discussion of xanthogastra group).

32. Apical white bands on tergites - 0) present except sometimes on apical tergite; 1) present on syntergite 1+2 and tergite 3; 2) only syntergite 1+2 with apical white band; 3) absent. Species with orange abdomens were coded state 0, because the bands are difficult to see. They are faint or possibly absent in Paraterellia, Haywardina and Zonosemata.

33. Abdominal medial white stripe - 0) present only on apical tergite or absent; 1) present on tergites 3-5 (male) or tergites 3-6 (female); 2) faint or reduced to posterior spot on tergites 3-4.

34. Sublateral white areas on tergites - 0) absent; 1) present on at least tergites 5-6.

35. Male sternite 6 with medial, usually mostly membranous lobe - 0) no; 1) yes.

36. Lateral surstylus, shape apical to medial surstylus (posterior view) - 0) tapered or truncate; 1) slightly expanded apically.

37. Lateral surstylus, mesal lobe - 0) short and sharply delimited; 1) broad and gradually arising from apical part of surstylus. The lobe is absent in Zonosemata, which was coded state 0.

38. Lateral surstylus, length apical to mesal lobe - 0) at least 2x width of lateral surstylus; 1) < 2x width of lateral surstylus; 2) absent. State 1 is plesiomorphic with R. striatella as outgroup.

39. Medial surstylus - 0) relatively long (in lateral view, distance from margin of epandrium to apex of medial surstylus at least 2x width of lateral surstylus); 1) relatively short (above ratio < or = 1).

40. Glans - 0) with distinct, complex internal sclerotization; 1) relatively simple, with little distinct internal sclerotization.

41. Glans - 0) short to moderately long, without sinuous internal tube; 1) elongate, with sinuous internal tube.

42. Apical membranous part of glans - 0) broad in ventral and/or lateral views; 1) tapered to single lobe slender in both views.

43. Aculeus tip length/ aculeus length - 0) < 0.4; 1) > 0.4.

44. Aculeus with lateral barb basal to tip - 0) no; 1) yes.

45. Aculeus tip sagittate and finely serrate - 0) no; 1) yes.

46. Aculeus tip with small subapical notches - 0) yes; 1) no. State 1 is plesiomorphic with either Rhagoletis species as outgroup.

47. Aculeus tip deeply trilobed in ventral view - 0) no; 1) yes.

48. Aculeus tip flattened in sagittal plane and trilobed in lateral view - 0) no; 1) yes.

49. Number of spermathecae - 0) 3; 1) 2. State 1 is plesiomorphic with either Rhagoletis species as outgroup.

50. Spermathecal shape - 0) small, spherical, usually with short neck, normally sclerotized; 1) large, elongate, and weakly sclerotized.


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Content by Allen L. Norrbom. Last Updated: January 31, 2002.