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Phylogenetic Relationships Among the Species of Panthiades Hubner (Lycaenidae: Theclinae: Eumaein1)

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Phylogenetic Relationships Among the Species of Panthiades Hubner (Lycaenidae: Theclinae: Eumaein1) 30 June 2005 PROC. ENTOMOL. SOC. WASH. 107(3), 2005, pp. 501-509 PHYLOGENETIC RELATIONSHIPS AMONG THE SPECIES OF PANTHIADES HUBNER (LYCAENIDAE: THECLINAE: EUMAEIN1) ROBERT K. ROBBINS Department of Entomology, RO. Box 37012, NHB Stop 127, Smithsonian Institution, Washington, DC 20013-7012, U.S.A. (e-mail: [email protected]) Abstract.•A species level phylogenetic analysis of Panthiades Hubner was performed using twelve characters of wing pattern, androconia, and male and female genitalia. The purposes were to determine whether Panthiades is monophyletic without Cycnus Hubner and to provide a cladogram for a project on the evolution of "false head" wing patterns. Parsimony analysis with all characters unordered yielded two trees. One was the strict consensus of the two trees, and the other was the only most parsimonious tree when one of the multi-state characters was ordered. Panthiades is characterized by five hypothesized synapomorphies. If Cycnus is recognized, Panthiades is not monophyletic on either of the most parsimonious cladograms. The "classic" false head wing patterns in Panthiades appear to have evolved once. Key Words: hairstreaks, false head hypothesis, Cycnus Nicolay's (1976) taxonomic treatment of unclear whether the classic "false head" Panthiades Hubner and Cycnus Hubner has wing pattern evolved once or twice in the been stable with minor exceptions. Robbins Panthiades/Cycnus lineage, especially since (2004a, b) changed two specific epithets for Nicolay (1976) placed one in Cycnus and nomenclatural reasons and synonymized one in Panthiades. the monotypic Cycnus with Panthiades, In this paper, I infer phylogenetic rela- stating that Panthiades was probably not tions among the eight species of Panthiades monophyletic without Cycnus. Consistent for the purposes of assessing the monophy- with this synonymy, Nicolay (1976:3) had ly of Panthiades without Cycnus and of noted that "the entry of the ductus semin- providing a cladogram for ongoing studies alis on the ventral-lateral side of the corpus of the evolution of wing patterns associated bursae" is shared by Panthiades and Cyc- with the "false head" hypothesis. nus, but not by other close relatives. Panthiades is of biological interest be- MATERIALS AND METHODS cause it contains two species, P. hathildis Coded characters were derived from a (Reakirt) and P. phaleros (L.), that have comparison of adult morphology using wing patterns (Fig. 10) traditionally asso- 1,009 pinned specimens of Panthiades in ciated with the "false head" hypothesis of the National Museum of Natural History, predator avoidance (Robbins 1980). These Smithsonian Institution, Washington, DC, wing patterns show a significantly greater USA, plus numerous specimens borrowed incidence of unsuccessful predator attacks from other museums. In addition, 41 geni- directed to the "false head" than other ly- talic dissections of both sexes of the eight caenid wing patterns (Robbins 1981). It is Panthiades species were examined. Pan- 502 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OE WASHING TON Table 1. Character matrix for Panthiades, The outgroups arc the lypc species of Parrhasius, Thepytus, and Porthecla. Characters and their states are detailed in the text. Cha raelers 2 Sp ?CLOS i ! 5 (J • 4 7 8 V 10 11 12 Panthiades bilias 0 0 1 0 I 1 3 9 1 0 0 0 Panthiades hebraeus 0 0 1 0 I 1 3 2 1 0 0 0 Panthiades aeolus 0 0 0 ') 0 1 1 0 1 0 0 0 Panthiades horeas 9 0 1 1 0 0 3 2 1 0 0 0 Panthiades ochus 0 0 1 1 0 0 3 2 1 0 0 0 Panthiades paphlagon 0 0 1 1 0 0 3 2 1 0 0 0 Panthiades bathildis 9 1 1 0 0 1 2 1 1 0 0 0 Panthiades phaleros ? 1 1 0 0 1 2 1 1 1 0 0 Parrhasius polibetes 1 0 0 1 ? 1 0 ? 0 2 1 1 Thepytus epytus 1 0 0 0 0 1 n 0 1 2 1 1 Porthecla porthura 1 0 0 0 ? 1 0 0 0 2 1 1 thiades genitalia have been illustrated (Nic- cially the valvae, are phenetically most sim- olay 1976) except for the female of P. ho- ilar to those of Panthiades. reas (Felder and Fclder), which is figured The character state for each Panthiades in this paper. Genitalic terms follow those and outgroup species is listed in the char- in Klots (1970), as illustrated in Robbins acter matrix (Table I). I used the implicit and Nicolay (2002). Androconial terminol- enumeration option of Hennig86 software ogy follows Robbins (1991), and wing vein to derive a most parsimonious cladogram. names follow Nicolay (1971, 1977). A strict consensus tree was determined. To The terminal taxa are the eight species test the assumption of equally weighted that have been placed in Panthiades or Cyc- characters, successive weighting was per- nus (Nicolay 1976, Robbins 2004b). They formed (Farris 1969) and a consensus of the are: P. bitias (Cramer, 1777), P. hebraeus resulting trees was determined. All charac- (Hewitson, 1867), P. aeolus (Fabricius, ters were unordered, except for multi-state 1775), P. boreas (C. Felder and R. Felder, character 7. Because it forms a morphocline 1865), P. ochus (Godman and Salvin, (cf., Pogue and Mickevich 1990), it was an- 1887), P. paphlagon (C. Felder and R. alyzed ordered and unordered. Mapping of Felder, 1865), P. bathildis (C. Felder and characters on trees was done with Winclada R. Felder, 1865), and P. phaleros (Linnae- software (Nixon 2002) with the fast opti- us, 1767). These taxa can be identified us- mization option. Jackknife support was de- ing the key in Nicolay (1976). The adults termined in Winclada using Nona (1000 were illustrated in D'Abrera (1995). replications with mult* 10, memory 1000 One outgroup for the analysis is Par- trees). rhasius polibetes (Cramer), the type species of Parrhasius Hiibner, a genus that has been MORPHOLOGY AND CODED CHARACTERS considered to be congeneric with Panthia- Ventral wing pattern.•The ventral wing des (Clench 1961) or its closest relative pattern in Panthiades is highly variable. (Nicolay 1976). To test the robustness of However, determining homology among the the root, I also used as outgroups the type different wing pattern elements is difficult species of Thepytus Robbins, T. epytus because these elements are not recognizable (Godman and Salvin), and of Porthecla in some species. Further, I am interested in Robbins, P. porthura (H.H. Druce). Among using the resulting cladogram to examine the genera of the Panthiades Section (Rob- wing pattern evolution within Panthiades. bins 2004a,b), their male genitalia, espe- For these reasons, the only character coded VOLUME 107, NUMBER 3 503 has one state in Panthiades and a second 4). It is unclear whether these scales form state in all other Panthiades Section genera, a scent patch, but I tentatively treat them as so it does not affect the inferred phyloge- androconia because they are restricted to netic relations within the genus. males. They occur in three species of Pan- Character 1: Ventral hindwing postme- thiades, one Thepytus, and two Parrhasius dian line segment in cell Sc + Rl-Rs (0) co- (Nicolay 1979). The size and shape of the linear with remainder of postmedian line, black scales often varies geographically, at (1) basally displaced. In three Panthiades least in Panthiades and Parrhasius (Nico- species, the postmedian line is not recog- lay 1976, 1979). There is a patch of dark nizable, so these species were coded with a brown scales in P. aeolus between the scent question mark. pads on the disco-cellular veins and at the Dorsal wing pattern• Character 2: Dor- base of vein M3 (Fig. 3, arrow B) that could sal wings of female with (0) shining blue- possibly be homologous with the black green iridescence, (1) a varying amount of scales, for which reason I code the second dull, "chalky" blue scales. The second character below with a question mark for P. character state is restricted in Panthiades to aeolus. P. bathildis and P. phaleros, where it varies Character 3: Scent pad in the discal cell within each species from no blue to a dull surrounded by a ring of scales that are tight- blue sheen that is distinguishable from the ly attached to the wing membrane (0) ab- shining iridescent blue-green of the other sent (Fig. 3), (1) present (Fig. 4). species. Character 4: Black scent patch distal of Androconia.•Androconia in Panthiades the discal cell (0) absent, (1) present (Fig. are restricted to the dorsal forewing and are 4, letter B). composed of three parts. The first is a black Male genitalia.•As noted by Nicolay (rarely gray or tan) scent pad in the discal (1976), the genitalia of Panthiades are more cell surrounded by a conspicuous ring of interspecifically variable than those of scales (Figs. 1•2, also fig. 122 in Eliot many other eumaeine genera. Five charac- 1973), which are usually gray in color. ters are coded. These androconia and the surrounding ring Character 5: Number of cornuti in penis of scales are tightly attached to the wing (0) 1, (1) 2. Cornuti are usually easily membrane. They occur in all Panthiades except P. aeolus (Fig. 3), but nowhere else scored in eumaeines, including Panthiades. in the Eumaeini. However, outgroup genera Parrhasius and The second part is a gray to dark char- Porthecla are problematic in that the folded coal colored scent pad that is universal in vesica within the penis has patches of vary- the Panthiades Section. It is distal of the ing sclerotization, making it difficult to de- ring of scales (Figs. 1•2, 4) and is, in turn, termine what is and is not a cornutus. De- composed of two parts. The first is very spite this problem in Parrhasius and Por- roughly oval and usually covers the upper thecla, for which reason they are coded and middle disco-cellular veins and parts of with question marks, the two states within the wings basal and distal of these veins Panthiades are clear.
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