Cotithene

Franz, Nico M., 2008, Revision, phylogeny and natural history of Cotithene Vo s s (Coleoptera: Curculionidae), Zootaxa 1782, pp. 1-33 : 27-29

Phylogeny of Cotithene – cladogram and character analysis

Cladogram

The successively refined character state assignments and optimizations on the resulting cladogram are presented simultaneously in this section. Cladistic analysis of a matrix with 12 terminal taxa and 32 characters, including two non-additive multistate characters ( Table 1), and performing an exhaustive search of the tree space produced a single most-parsimonious cladogram ( Fig. 14 ), with a length of 38 steps, consistency index (CI) of 89 and retention index (RI) of 93 (see Farris 1989). Three characters (4, 13 and 15) had multiple parsimonious optimizations, and in each case the fast (ACCTRAN) optimization was preferred, thereby preserving the original homology assessment (de Pinna 1991).

1. Size: (0) Small, length 1.6–3.4 mm; (1) larger, length 4.4–6.1 mm). Synapomorphy for the leptorhamphismelanoptera clade.

2. Maxilla, galeo-lacinial complex, apical region: (0) lacinia projecting along entire length of palpomere I; (1) lacinia apically truncate, projecting slightly beyond base of palpomere I ( Fig. 4 B). Synapomorphy for the Cyclanthura-Cotithene clade, corresponding to char. 7 in Franz (2006).

3. Labium, palpomere II, apical region: (0) apicolateral projection absent; (1) apicolateral projection present ( Fig. 4 C). Synapomorphy for the Cyclanthura-Cotithene clade, corresponding to char. 43 in Franz (2006).

4. Rostrum, dorsal view, basal region, male: (0) indistinct, subplane; (1) slightly impressed along basal 1/5. Fast optimization preferred (see above); synapomorphy for the globulicollis-melanoptera clade, with an apparent reversal in C. anaphalanta (ci = 50, ri = 0).

5. Rostrum, dorsal view, apical region, male: (0) indistinct, subplane; (1) with 2 apicolateral tubercles. Synapomorphy for Terioltes , corresponding to char. 53 in Franz (2006).

6. Rostrum, lateral view, male: (0) width similar throughout; (1) variously expanded near apex to basal 2/5 (e.g., Figs. 5 A, 8A). Synapomorphy for the Terioltes-Cotithene clade (see also char. 7).

7. Rostral expansion, lateral view, male: (0) apical to subapical; (1) near basal 2/5 of rostrum (e.g., Fig. 5 A); (2) near apical 2/5 of rostrum (e.g., Fig. 8 A). Inapplicable in taxa that lack a rostral expansion (see char. 6). Coded as nonadditive; synapomorphy for Cotithene (state 1), with an apparent subsequent transformation in the leptorhamphis-melanoptera clade (state 2). Refinement of char. 54 in Franz (2006).

8. Rostrum, dorsolateral margins, male: (0) indistinct, convex; (1) carinate. Synapomorphy for Cotithene , corresponding to char. 50 in Franz (2006).

9. Rostrum, lateral view, female: (0) slightly shorter to slightly longer than pronotum, r/p = 0.8–1.3, fairly broad to fairly narrow, slightly arcuate ( Figs. 2 D, 8B); (1) distinctly longer than pronotum, r/p = 1.7–2.1, very narrow, arcuate ( Figs. 6 B, 7B). Synapomorphy for the leptorhamphis-trigaea clade.

10. Antenna, scape: (0) reaching anterior margin of eye; (1) reaching subbase of rostrum. Synapomorphy for the leptorhamphis-trigaea clade.

11. Rostrum-head transition, lateral view, female: (0) contiguous, evenly convex in profile ( Figs. 2 D, 8B); (1) slightly angulate, concave in profile ( Figs. 6 B, 7B). Synapomorphy for the leptorhamphis-trigaea clade.

12. Head, dorsal region, lateral view, male: (0) evenly slightly convex between eyes ( Fig. 5 A); (1) elevated, convex, variously shaped and sculptured between eyes (e.g., Figs. 3 A, 8A). Synapomorphy for the stratiotricha-melanoptera clade.

13. Head, dorsal region, male: (0) posteriorly slightly, evenly convex; (1) posteromedially with small (large in C. stratiotricha ; see Fig. 3 D), slightly elevated, triangular, anteriorly directed projection. Fast optimization preferred (see above); synapomorphy for the stratiotricha-melanoptera clade, with an apparent reversal in the leptorhamphis-trigaea clade (ci = 50, ri = 66).

14. Head, anterodorsal region (frons), sculpture, male: (0) slightly convex; (1) rugulose to rugose, with several dense, very small, convex elevations. Synapomorphy for the leptorhamphis-trigaea clade.

15. Head, dorsal region (frons), male: (0) with sparse, short, appressed, recurvate vestiture ( Fig. 5 A); (1) with sparse to dense, longer, suberect to recurvate setae (e.g., Figs. 3 C, 8A). Fast optimization preferred (see above); synapomorphy for the stratiotricha-melanoptera clade, with an apparent reversal in the leptorhamphis-trigaea clade (ci = 50, ri = 66).

16. Head, ventral region (gena), male: (0) indistinct, glabrate and evenly convex; (1) with two large, transverse sulci. Synapomorphy for Terioltes , corresponding to char. 68 in Franz (2006).

17. Head, ventral region (gula), male: (0) in profile evenly convex; (1) in profile distinctly angulate, with transverse line separating anterior and posterior half, each region subplane (e.g., Figs. 3 C, 5A). Synapomorphy for Cotithene .

18. Prothorax, shape, dorsal view, male: (0) similar in size or slightly larger than in female and of similar proportions; (1) variously expanded, convex to globular (e.g., Figs. 1 A, 1B). Synapomorphy for the Ganglionus-Cotithene clade (see also char. 19). Reinterpretation of char. 72 in Franz (2006) to now also include Ganglionus Franz & O'Brien.

19. Pronotum, dorsolateral view, male: (0) slightly expanded, convex, ( Fig. 1 A), dorsolaterally slightly angled; (1) strongly expanded, convex ( Fig. 1 B), dorsolaterally rounded. Inapplicable in taxa that lack a pronotal expansion (see char. 18). Convergently present in Terioltes and in the leptorhamphis-melanoptera clade (ci = 50, ri = 50).

20. Prosternum, anterior margin, male: (0) with short, indistinct setation; (1) with row of dense, long, aurate setae directed anteriad (e.g., Figs. 3 C, 5A). Synapomorphy for Cotithene .

21. Prosternum, anterior region, male: (0) indistinct, slightly convex to slightly concave; (1) with small, obtuse elevation in front of each procoxal cavity. Synapomorphy for the globulicollis-melanoptera clade.

22. Procoxal cavities, female: (0) contiguous; (1) subcontiguous, separated by narrow septum; (2) separated by about width of antennal club. Coded as nonadditive; synapomorphy for Cotithene (state 1), with an apparent subsequent transformation in the leptorhamphis-melanoptera clade (state 2). Refinement of char. 81 in Franz (2006).

23. Elytra, color: (0) variously light to dark-reddish brown; (1) uniformly dark-reddish brown to black, other parts nearly orange. Synapomorphy for the anaphalanta-melanoptera clade.

24. Pygidium, orientation, lateral view, male: (0) subparallel to anterior tergites; (1) perpendicular to venter. Synapomorphy for the Cyclanthura-Cotithene clade, corresponding to char. 116 in Franz (2006).

25. Aedeagus, dorsal view: (0) apically gradually, evenly narrowed (e.g., Figs. 9 A, 9B); (1) constricted near apical third, thereafter slightly expanded ( Figs. 10 A, 10B). Synapomorphy for the leptorhamphis-trigaea clade.

26. Aedeagus, lateral view: (0) variously deflexed; (1) sinuate, basal 3/5 slightly deflexed, thereafter reflexed, very narrow. Synapomorphy for Terioltes .

27. Aedeagus, shape, subapical region, lateral view: (0) dorsal margins simple; (1) dorsal margins with separate inner rami, expanded, convex (e.g., Figs. 9 A, 10A). Synapomorphy for Cotithene .

28. Aedeagus, shape, apical region, lateral view: (0) dorsal and ventral margins variously slightly to distinctly arcuate (e.g., Figs. 9 B, 10B); (1) dorsal and ventral margins straight (e.g., Figs. 12 A, 12B). Synapomorphy for the globulicollis-melanoptera clade.

29. Aedeagus, internal sac, subapical region, dorsal view: (0) without paired sclerites; (1) with pair of variously shaped, elongate, subparallel sclerites (e.g., Figs. 9 A, 10A). Synapomorphy for the Terioltes- Cotithene clade, corresponding to char. 144 in Franz (2006).

30. Shape of aedeagal sclerites, dorsal view: (0) elongate, slightly overlapping, margins irregular-sinuate, orientation horizontal; (1) triangular, apically variously modified, subparallel to apically slightly converging, orientation vertical (e.g., Figs. 9 A, 12A). Inapplicable in taxa that lack aedeagal sclerites (see char. 29). Synapomorphy for Cotithene ; C. stratiotricha ( Fig. 9 B) presents several unique modifications that are presumed, in light of the overall tree structure, to have derived from the apomorphic state.

31. Shape of aedeagal sclerites, lateral view: (0) apically narrowed, simple (e.g., Figs. 9 A, 11); (1) apically expanded, with undulate margins ( Figs. 10 A, 10B). Inapplicable in taxa that lack aedeagal sclerites (see char. 29). Synapomorphy for the leptorhamphis-trigaea clade.

32. Aedeagus, internal sac, ventral view: (0) ventral region of internal sac membranous, denticulate, otherwise indistinct (e.g., Figs. 9 A, 11); (1) ventral region of internal sac with complex, partly membranous and party sclerotized, irregularly subquadrate to annulate structure, dorsally with elongate, retorted membranous connection to apical region of paired sclerites ( Figs. 12 A, 12B). Synapomorphy for the anaphalantamelanoptera clade.

Taxon/character 5 10 15 20 25 30

Notolomus basalis 0 0 0 0 0 0–000 0 0 0 0 0 000–0 0 0 0 0 0 0000– –0 Cyclanthura laticola 0 1100 0–000 0 0 0 0 0 000–0 0 0 0 10 0000– –0 Ganglionus undulatus 0 1100 0–000 0 0 0 0 0 0 0 100 0 0 0 10 0000– –0 Terioltes circumdatus 0 1101 10000 0 0 0 0 0 10110 0 0 0 10 10010 0 0 Terioltes nigripennis 0 1101 10000 0 0 0 0 0 10110 0 0 0 10 10010 0 0

Cotithene anaphalanta 11100 12100 0 1101 0 1111 12110 0 1111 0 1 Cotithene dicranopygia 0 1100 11100 0 0 0 0 0 0 1101 0 1010 0 1011 0 0 Cotithene globulicollis 11110 12100 0 1101 0 1111 12010 0 1111 0 0 Cotithene leptorhamphis 11100 12111 11010 0 1111 0 2011 0 1011 10 Cotithene melanoptera 11110 12100 0 1101 0 1111 12110 0 1111 0 1 Cotithene stratiotricha 0 1100 11100 0 1101 0 1101 0 1010 0 1011 0 0 Cotithene trigaea 11100 12111 11010 0 1111 0 2011 0 1011 10

Figures

FIGURE 1. Dorsal habitus of Cotithene, males. A, C. dicranopygia; B, C. globulicollis.

FIGURE 2. Habitus of Cotithene stratiotricha. A, male, dorsal view; B, male, lateral view; C, female, dorsal view; D, female, lateral view.

FIGURE 3. Variation of rostrum and head in males of Cotithene stratiotricha. A, small male, lateral view; B, intermediate-sized male, lateral view; C, large male, lateral view; D, large male, dorsal view.

FIGURE 4. Mouthparts of Cotithene globulicollis. A, left mandible; B, right maxilla; C, labium. Scale bars 0.1 mm. For further detail see also Franz (2006).

FIGURE 5. Rostrum and head of Cotithene males, lateral view. A, C. dicranopygia; B, C. stratiotricha.

FIGURE 6. Rostrum and head of Cotithene leptorhamphis, lateral view. A, male; B, female.

FIGURE 8. Rostrum and head of Cotithene globulicollis, lateral view. A, male; B, female.

FIGURE 9. Genital structures of Cotithene species, from left to right: aedeagus, ventral view; aedeagus, lateral view; posterior region of female sternum VIII, ventral view; spermatheca, lateral view; shaded areas depicting structures of the internal sac. A, C. dicranopygia; B, C. stratiotricha.

FIGURE 10. Genital structures of Cotithene species, from left to right: aedeagus, ventral view; aedeagus, lateral view; posterior region of female sternum VIII, ventral view; spermatheca, lateral view; shaded areas depicting structures of the internal sac; synapomorphic character states 31 (1) labelled. A, C. leptorhamphis; B, C. trigaea.

FIGURE 12. Genital structures of Cotithene species, from left to right: aedeagus, ventral view; aedeagus, lateral view; posterior region of female sternum VIII, ventral view; spermatheca, lateral view; shaded areas depicting structures of the internal sac; synapomorphic character states 32 (1) labelled. A, C. anaphalanta; B, C. melanoptera.

FIGURE 14. Phylogeny of Cotithene and selected outgroup taxa, most parsimonious cladogram (L = 38, CI = 89, CI = 93). Characters 4, 13 and 15 displayed under fast (ACCTRAN) optimization, all other characters have unambiguous optimizations. Black rectangles – single, non-homoplasious character state transformations, white rectangles – multiple, homoplasious character state transformations. Numbers above and below each rectangle correspond to characters and states, respectively; numbers at left end of each branch are Bremer support values (see text for details).