Figure 1.
Previous hypotheses of phylogenetic relationships in Tortricidae.
A. Powell (1964; [20]), B. Kuznetsov and Stekolnikov (1973; [21]), C. Razowski (1976; [23]), D. Kuznetsov and Stekolnikov (1977; [17]), E. Kuznetsov and Stekolnikov (1984; [22]), F. Safonkin (2007; [25]). Tree figures re-drawn, but nomenclature in each case follows the original.
Table 1.
Groups previously considered families by one or more tortricid workers.
Table 2.
Three recent classifications of Tortricidae and their agreement with molecular evidence. Number of exemplars in current study is given.
Figure 2.
Adult habitus of representatives of Tortricidae used in the analysis.
A–L, Tortricinae; M–R, Olethreutinae. A: Polyorthini, Histura perseavora Brown, Guatemala; B: Hilarographini, Hilarographa sp., Costa Rica; C: Chlidanothini, Auratonota sp., Costa Rica; D: Phricanthini, Phricanthes asperana Meyrick, Australia; E: Atterini, Anacrusis stapiana Felder & Rogenhofer, Costa Rica; F: Sparganothini, Amorbia humerosana Clemens, USA; G: Tortricini, Acleris semipurpurana Kearfott, USA; H: Archipini, Choristoneura rosaceana Harris, USA; I: Cnephasiini, Cnephasia alfacarana Razowski, Spain; J: Ceracini, Cerace sp., Japan; K: Euliini, Bonagota sp., Uruguay; L: Cochylini, Carolella sartana Hübner, USA; M: Microcorsini, Cryptaspasma bipenicilla Brown & Brown, USA; N: Olethreutini, Afroploce karsholti Aarvik, Tanzania; O: Bactrini, Bactra furfurana Haworth, Denmark; P: Enarmoniini, Ancylis sparulana Staudinger, Spain; Q: Eucosmini, Gypsonoma paradelta Meyrick, Tanzania; R: Grapholitini, Cydia pomonella Linnaeus, Spain. (A,C, E–H, J–M, US National Museum of Natural History, Smithsonian Institution; B, School of Biology, University of Costa Rica; D, Natural History Museum London; I, O, P, R, Cavanilles Institute of Biodiversity and Evolutionary Biology, University of Valencia; N, Q, Natural History Museum, University of Oslo). Scale bars: 5 mm.
Table 3.
Species sampled and their distribution across the current classification. Diversity numbers based on Baixeras et al. [2], distributions largely based on Horak [27], [29].
Table 4.
Outgroup species sampled.
Figure 3.
Maximum likelihood estimate of phylogenetic relationships in Tortricidae.
Tree was obtained from 1000 GARLI searches under a GTR+gamma+I model for all nucleotides (unpartitioned). Bootstrap support values (742–901 pseudoreplicates) above branches for: nt123 (19 genes), nt123_partitioned (19 genes), degen1 (19 genes), nt123 (5 genes), degen1 (5 genes). “[-]” = node not present on best ML tree for that analysis. Nodes within Tortricidae are numbered (to the right of node) for purposes of discussion.
Figure 4.
Phylogram presentation of best maximum likelihood tree obtained from 1000 GARLI searches.
Model used was a GTR+gamma+I model for all nucleotides (unpartitioned). Thickened branches are supported by ≥80% bootstrap in at least one analysis (see Fig. 3).
Figure 5.
Synopsis of species diversity, distribution and larval host-plant use for the tribes of Tortricidae studied, mapped onto a phylogeny condensed from Fig. 3.
The multiple branches leading to the names Eucosmini and Olethreutini s.s. denote the paraphyly of these tribes discovered here. Col. 1 = number of described species, taken from Table 3. Col. 2 = summary of geographic distributions, following Horak [29]. Cols. 3, 4, 5 = summary statements of predominant mode of larval feeding, host range, host taxa used and egg-laying habits, following Horak [29], Powell et al. [33] and J. Brown et al. [56]. ‘[]’ denotes assertion based on very few observations; preceding question mark denotes assertion based on uncertain or conflicting information; ‘( )’ denotes habits significantly represented but markedly less common than alternative; ‘;’ separates habits of similar frequency. In column 4, “oligophagous” = feeding on a single plant family, “polyphagous” = feeding on two or more plant families, as described in the text. Numbers in parentheses, compiled from Brown et al. [56], are number of polyphagous species/total number of species with two or more foodplant observations = % polyphagous species.