Description of Males of Ptychoderes brevis and Ptychoderes jekeli, with a Cladistical Reanalysis of Ptychoderes (Coleoptera: Anthribidae)

Author(s): Jose Ricardo M. Mermudes and Ingrid Mattos Source: Annals of the Entomological Society of America, 105(4):523-531. 2010. Published By: Entomological Society of America DOI: http://dx.doi.org/10.1603/AN10016 URL: http://www.bioone.org/doi/full/10.1603/AN10016

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SYSTEMATICS

Description of Males of Ptychoderes brevis and Ptychoderes jekeli, With a Cladistical Reanalysis of Ptychoderes (Coleoptera: Anthribidae)
JOSE RICARDO M. MERMUDES1
AND

INGRID MATTOS1,2

Ann. Entomol. Soc. Am. 105(4): 523531 (2010); DOI: 10.1603/AN10016

ABSTRACT A recent study of the material housed in the Muse um National dHistoire Naturelle (Paris, France) provided new information to describe and illustrate the males of Ptychoderes brevis Jordan and Ptychoderes jekeli Mermudes & Napp (Coleoptera: Anthribidae). These males were previously unknown, and their characters had never been studied by the cladistical method. A reanalysis of the genus was carried out, which included 50 characters and 24 taxa (seven as outgroup). The resulting single most-parsimonious cladogram corroborates previous hypotheses of the relationships among the genera of Ptychoderini and groups of Ptychoderes species. The male of P. brevis is characterized by: rostrum 1.5 longer than wide across base; sides not swollen between scrobe and eye; scrobe with a deep narrow extension and a deep distal pit; antennae elongated, reaching middle of elytra; ventrite I swollen, with moderately developed setiferous sex patch, closer to apical margin, clothed with dense yellow pilosity; and ventrite V with lateroapical projections not acuminate. The male of P. jekeli has rostrum slightly longer than wide across base; sides not swollen and the extension shallow; antennae exceeding apical elytron by segment XI; ventrite I with well developed setiferous sex patch, closer to apical margin and with olive green pilosity; and ventrite V strongly impressed, with lateroapical angles not projected and somewhat expanded. The relationship between P. brevis and P. jekeli was conrmed, and this group belongs to the P. crustatus, corroborating the most basal lineage by the vestiture of the sides of prosternum with a vitta of the same color as the dorsal vestiture, and elytra transversely impressed behind the basal gibbosities. RESUMO Um estudo recente do material do Muse um National dHistorie Naturelle (Paris, France) possibilitou descrever e ilustrar machos de Ptychoderes brevis Jordan e Ptychoderes jekeli Mermudes & Napp (Coleoptera: Anthribidae) que eram desconhecidos e os caracteres nunca utilizados na clad stica. A reana lise do ge nero incluiu 50 caracteres e 24 ta xons (sete como grupo externo) e resultou em um cladograma mais parcimonioso corroborando hipo teses anteriores para as relac o es entre os ge neros neotropicais da tribo e grupos de espe cies em Ptychoderes. O macho de P. brevis e caracterizado pelo rostro 1.5 vezes mais longo que a largura basal; lados do rostro na o intumescidos; escrobo com extensa o profunda e estreita e com fo vea distal profunda; antena alongada, alcanc ando o meio do e litro; ventrito I intumescido com a mancha set gera moderadamente desenvolvida, pro xima da margem apical e com densa pilosidade amarelada; e ventrito V com projec o es la tero-apicais na o acuminadas. O macho de P. jekeli apresenta o rostro pouco mais longo que a largura basal, lados na o intumescidos, com extensa o do escrobo pouco profundo; antenas ultrapassam o a pice elitral pelo anteno mero XI; ventrito I com macha set gera desenvolvida, pro xima da margem apical e com pilosidade verde-clara; e ventrito V fortemente deprimido com a ngulos la tero-apicais na o projetados e algo expandidos lateralmente. A relac a o de P. brevis e P. jekeli e conrmada novamente pela presenc a de um u nico ponto grosso na a rea glabra do profe mur, pronoto com tube rculo arredondado na depressa o central, menos proeminente que as margens da depressa o central e ventrito I com mancha set gera pequena em P. brevis e pouco mais desenvolvida em P. jekeli. Este grupo tambe m foi conrmado como grupo irma o de P. crustatus, conrmado como o clado mais basal pelas seguintes sinapomoras: lados do prosterno com faixa concolor com o revestimento dorsal e e litros com depressa o transversa apo s as gibosidades. KEY WORDS Anthribinae, fungus weevils, morphology, phylogeny, Ptychoderini

1 Corresponding author: Laborato rio de Entomologia, Departamento de Zoologia, Universidade Federal do Rio de Janeiro, Caixa Postal 68044, 21941-971, Rio de Janeiro-RJ, Brazil (e-mail: jrmermudes@gmail.com). 2 Programa de Po s Graduac a o em Biocie ncias, Universidade do Estado do Rio de Janeiro, Rio de Janeiro-RJ, Brazil. (Capes Fellow.)

0013-8746/10/05230531$04.00/0 2010 Entomological Society of America

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The tribe Ptychoderini was proposed by Jekel (1855) based on Neotropical genera, when he revised the genus Ptychoderes Schoenherr, 1823 and described six species: Ptychoderes columbianus, from Colombia and Brazil, and Ptychoderes depressus, Ptychoderes callosus, Ptychoderes mixtus, Ptychoderes antiquus, and Ptychoderes obsoletus, all from Brazil. In addition, he proposed ve new Neotropical genera (Hypselotropis, Tribotropis, Ectatotropis, Tropipygus, and Stenometopus) for some of the Phloeotragus Schoenherr species described by Fahraeus in Schoenherr (1839). Eleven years later, Lacordaire (1866) established the tribe Phloeotragini for the same group proposed by Jekel (1855), adding non-Neotropical genera: Phloeotragus, Phloeopemon, and Cerambyrhynchus. Over time, other authors added some species, especially Jordan (1894), who described Ptychoderes brevis and Ptychoderes afnis from Brazil, Ptychoderes bivittatus from Mexico and Nicaragua, and Ptychoderes longicollis from Suriname. Later, Jordan (1895) described P. rugicollis from Panama and Nicaragua. In the same study, he reinstated the genus Hypselotropis, which had been synonymized under Ptychoderes by Lacordaire (1866). Recently, Alonso-Zarazaga and Lyal (1999) considered Phloeotragini Lacordaire, 1866 a synonym of Ptychoderini Jekel, 1855, retaining the non-Neotropical genera in the tribe. Studies of Neotropical fauna of Anthribidae have continued since the contributions by Mermudes (2002, 2003). Mermudes and Napp (2004) added two new species of Ptychoderes. Mermudes (2005) published a revision, cladistic analysis, and biogeography of the genera Hypselotropis and Tribotropis, proposing the latter as a synonym of the former. Mermudes and Napp (2006) studied the phylogenetic relationships among species of Ptychoderes. The name Tribotropis used by Mermudes and Napp (2006) is a lapse due to overlap of two recent articles. Currently, the tribe consists of six genera, three of which are restricted to the Neotropical region: Unanthribus Mermudes, 2003 (two species), Hypselotropis Jekel, 1855 (17 species), and Ptychoderes Schoenherr, 1823 (17 species); one Afrotropical genus, Phloeotragus Schoenherr, 1823 (13 species); Phloeopemon Schoenherr, 1839 (four species), with an Oriental and Indomalayan distribution; and a single species (Phloeopemon acuticornis) extending to Japan in the Palearctic region; and Cerambyrhynchus, a monotypic genus of the Australian region. At the time of the study by Mermudes and Napp (2006), the males of the species P. brevis and P. jekel were unknown, and their characters had never been studied using the cladistic methodology, resulting in uncertainty about basal lineages of the genus. These two species form a sister group to the clade P. crustatus, considered the most basal lineage. A recent study of material deposited in the Muse um National dHistoire Naturelle (Paris, France) allowed the recognition of the previously undescribed males of these two species. In this study, the males of P. brevis and P. jekel are described and illustrated for the rst time, and a new cladistic analysis for the genus is presented,

including characters of the external morphology of the males. Materials and Methods Material Examined. We studied specimens obtained through exchanges, loans, and donations from the following institutions (names of curators in parenthesis): BMNH, The Natural History Museum, London, England (C. Lyal, M. Barclay); BPBM, Bernice P. Bishop Museum, Honolulu, HI (G. A. Samuelson); CASC, California Academy of Sciences, San Francisco, CA, U.S.A. (Norman Penny); DEI, Deutsches Entomologisches Institut, Mu ncheberg, Germany (Dr. Lothar Zerche); DZUP, Departamento de Zoologia, Universidade Federal do Parana , Curitiba, Brazil (L. Marinoni); ISNB, Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium (J. Cools); IAHC, Instituto de Investigacio n de Recursos Biolo gicos Alexander von Humboldt, Vila de Leyva, Colombia (Y. M. Saba, F. Ferna ndez); INBC, Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica (A. Solis); INPA, Instituto Nacional de Pesquisas da Amazo nia, Manaus, Brazil (A. Henriques); MNHN, Muse um National dHistoire Naturelle, Paris, France (H. Perrin); MNRJ, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (M. A. Monne ); and MZUSP, Museu de Zoologia, Universidade de Sa o Paulo, Sa o Paulo, Brazil (C. A. Casari). The methods, terminology, and measurements used in this study follow Mermudes (2002, 2003, 2005) and Mermudes and Napp (2006). Cladistical Methods. All taxa and characters in a previous analysis (Mermudes and Napp 2006) were reexamined, with the addition of new character states for the males of P. brevis and P. jekeli (Table 1; see a list of characters in Appendix). The analyses with a data matrix compiled in NDE (Page 2001) were conducted with NONA, version 2.0 (Goloboff 1993) interface WINCLADA (Nixon 1999). Phylogenetic trees were rooted and character states polarized a posteriori using the outgroup (Nixon and Carpenter 1993). The outgroup included species of other genera of Ptychoderini. The search for the most parsimonious cladograms were carried out using the heuristic search option with tree bisection reconnection (TBR), and branch swapping with hold 100,000, 5000 replicates, and 30 trees retained for replica. Acctran and Deltran optimization was used. Clades are referred to using the name of the rst taxon included, followed by the plus sign (Amorim 1982). Systematics Ptychoderes brevis Jordan, 1894 (Figs. 1 4, 8 9) Description. Male. Vestiture (Figs. 1 4) exactly as described for female holotype (BMNH) by Mermudes and Napp (2006). The pale dorsal vitta with dense whitish scales, intermingled with olive green scales,

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Table 1.

MERMUDES AND MATTOS: MALES OF Ptychoderes AND CLADISTICAL REANALYSIS

Character matrix Taxa 1 0 0 0 0 0 0 0 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 3 0 0 0 0 0 0 0 0 0 0 1 1 0 0 2 2 1 2 0 0 1 1 0 0 4 0 0 0 0 0 0 1 3 3 3 2 1 1 2 2 2 2 1 1 1 1 2 1 5 0 0 0 0 0 1 1 1 2 1 1 1 2 1 2 2 2 2 2 1 1 1 1 1 6 0 0 0 0 0 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 7 0 1 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 8 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 9 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

525

10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 0 0 0 0 0 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 0 0 0 0 1 1 2 2 2 2 2 2 1 2 2 2 2 1 2 2 2 1 2 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 1 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 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 0 0 1 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 2 2 3 3 3 0 3 3 3 3 3 3 3 3 3 3 3 3 3 0 0 0 0 0 1 1 1 1 1 1 1 1 1 2 2 2 2 1 1 1 1 1 1 0 0 0 0 0 2 2 5 5 3 1 3 3 4 6 6 3 6 5 3 3 3 3 3 0 0 0 0 0 1 1 1 3 1 3 3 2 2 3 3 3 3 2 2 3 3 2 2 0 0 0 0 0 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 0 0 0 1 1 2 2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 0 0 0 0 0 0 0 0 0 2 0 1 0 0 3 2 2 3 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1

Cerambyrhynchus schoenherri Phloeopemon acuticornis Phloeotragus heros Hypselotropis prasinatus Hypselotropis annulicornis Unanthribus maximus Unanthribus grandis Ptychoderes brevis Ptychoderes jekeli Ptychoderes crustatus Ptychoderes bivittatus Ptychoderes callosus Ptychoderes jordani Ptychoderes depressus Ptychoderes elongatus Ptychoderes antiquus Ptychoderes viridanus Ptychoderes nebulosus Ptychoderes mixtus Ptychoderes obsoletus Ptychoderes longicollis Ptychoderes rugicollis Ptychoderes virgatus Ptychoderes magnus

26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 Cerambyrhynchus schoenherri Phloeopemon acuticornis Phloeotragus heros Hypselotropis prasinatus Hypselotropis annulicornis Unanthribus maximus Unanthribus grandis Ptychoderes brevis Ptychoderes jekeli Ptychoderes crustatus Ptychoderes bivittatus Ptychoderes callosus Ptychoderes jordani Ptychoderes depressus Ptychoderes elongatus Ptychoderes antiquus Ptychoderes viridanus Ptychoderes nebulosus Ptychoderes mixtus Ptychoderes obsoletus Ptychoderes longicollis Ptychoderes rugicollis Ptychoderes virgatus Ptychoderes magnus 0 0 0 0 0 ? 1 ? ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 1 1 1 1 1 1 2 2 1 1 2 1 1 1 1 1 1 0 0 0 0 0 1 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 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 2 3 3 3 3 1 1 1 1 1 1 0 0 0 0 0 1 1 0 0 0 0 3 0 2 4 0 0 4 2 2 0 3 2 2 0 0 0 0 0 0 0 1 1 1 2 3 1 1 3 3 3 3 2 1 2 3 1 1 0 0 0 0 0 1 1 2 1 2 1 1 1 1 3 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 1 1 0 0 2 1 0 2 2 2 2 1 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 2 2 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 ? 0 ? ? ? 1 1 2 2 1 1 1 1 1 2 2 1 2 2 0 0 0 1 1 ? 0 ? ? ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 1 ? ? ? 1 1 0 0 1 0 0 1 1 0 1 1 0 0 0 0 0 0 ? 1 ? ? ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 1 ? 0 ? ? ? 0 1 0 1 0 0 0 0 0 1 0 1 1 1 0 0 0 0 0 ? 0 ? ? ? 1 1 0 0 1 1 1 1 1 0 0 1 0 0 0 0 0 0 0 ? 0 ? ? ? 0 0 1 1 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 ? 0 ? ? 0 0 0 1 1 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 1 1 1 1 1 2 2 1 1 1 1 1 2 2 1 2 2 0 0 0 0 0 ? 0 ? ? 0 0 0 1 1 0 0 0 0 0 1 1 0 1 1 0 0 0 0 0 0 0 0 ? 0 0 0 1 1 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 ? 0 ? ? 1 2 2 1 1 2 2 2 2 2 1 1 2 1 1

decumbent and denser at sides of pronotum, elytra, and interstriae 1, 2, and 4 and apical declivity. Pygidium, ventral vestiture, and legs as in the female. Rostrum at least 1.5 times longer than wide across base. Sides of rostrum not swollen between scrobes and eyes, and tricarinate as in female. Scrobes with the same margins and a deep narrow extension, with a deep distal pit, slightly wider than extension. Antennae elongate, reaching middle of elytra. Segment II one third shorter than III; segments IVVII subequal in length, slightly longer than III; VIII one third shorter than VII; IX subequal to VIII; X slightly longer than wide; IX elongate, not narrowed, and slightly shorter than IX. Prothorax (Figs. 12) with pronotum, prosternum, and intercoxal process of mesosternum similar to ho-

lotype female. Elytra with surface rugose from base to middle. Outer apical glabrous area of profemur with a single deep puncture. Abdomen with ventrites attened; ventrite I (Figs. 3 and 4) swollen, with a setiferous sex patch moderately developed and clothed with dense yellow pilosity, closer to apical margin; ventrite V as long as IV, strongly depressed; with projected, not acuminated, lateroapical projections. Material Examined. FRENCH GUIANA: Route de Re gina. PK79, one male, 14-II-1994 LdA. Se ` ne uaux leg. and one female, Piste Corrlie. PK2, 2-II-1989, L. Se `ne uaux leg. (MNHN). Remarks. These are the third and fourth known specimens of P. brevis, and one of them is the rst male. It was discovered in MNHN. P. brevis belongs

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Figs. 14. P. brevis male. 1. Habitus. 2. Lateral view. 3. Meso- and metasternun, and ventrite I. 4. Detail of ventrite I and setiferous sex patch (arrowed). (Online gure in color.)

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Figures 57. P. jekeli male. 5. Habitus. 6. Lateral view. 7. Ventral view, ventrite I and setiferous sex patch (arrowed). (Online gure in color.)

to the P. crustatus group, being the sister group of P. jekeli (Mermudes and Napp 2004) by two synapomorphies according to Mermudes and Napp (2006): pronotal depression with a median, some-

what rounded, weakly prominent tubercle (205), and glabrous area of profemora with a single coarse puncture (362). Here, the ventrite I of males bears a small setiferous sex patch (341) corroborates this

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Fig. 8. Single most-parsimonious tree resulting from the reanalysis, showing the relationships among the Neotropical genera of Ptychoderini, and Ptychoderes interrelationships; 117 steps; Deltran optimization. CI 0.76 and RI 0.90. Synapomorphies are indicated by solid black circles; homoplasies by white circles.

lineage and occurs independently in P. jordani and P. mixtus. The pygidium not impressed medially near apex, and ventrite V with projected, not acuminate, lateroapical angles, both autapomorphies of P. brevis, were corroborated here. These characters differentiate this species from all other species of Ptychoderes. Distribution. Mermudes and Napp (2006) recorded P. brevis for Colombia and Brazil (Para and Mato Grosso). Here, the distribution is extended to French Guiana, being associated with the Amazon forest (Fig. 9). Ptychoderes jekeli Mermudes & Napp, 2004 (Figs. 57, 8 9) Description. Male. Vestiture (Figs. 57) with green scales somewhat lighter in color than in the female holotype and paratype housed in the BMNH. The vestiture of the sides of prosternum (Fig. 6), with a vitta of the same color as the dorsal vestiture. Rostrum slightly longer than wide across base; sides not swollen, and the extension shallow. Antennae elongate, reaching the elytral apex by segment XI; segment II one third shorter than III; IV and V subequal in length, slightly shorter than III; VIVII subequal to III; VIII one fourth shorter than VII; club weakly elongated; IX one third shorter than VIII; X longer than wide; XI as long as IX.

Prothorax (Figs. 57) with pronotum and carina similar to holotype and paratype females. Prosternum (Fig. 7) with coarse, deep dense punctures, some coalescent; antecoxal furrow wide and deep. Intercoxal process of mesosternum rounded at apex, impressed in the middle. Elytral surface rugose, with granules exceeding the middle (differing from P. brevis). Elytral apex with three to eight prominent interstriae (not tuberculate). Profemora with a single coarse puncture in glabrous area. Mesofemora with two coarse punctures in a distinct dark vitta. Abdomen with short pygidium, rounded at apex. Ventrite I with a developed setiferous sex patch, closer to the posterior margin, clothed with olive green pilosity; ventrite V strongly impressed, with lateroapical angles not projected and somewhat expanded. Material Examined. FRENCH GUIANA: Cayenne, one male, no other data (MNHN). Distribution. Mermudes and Napp (2004, 2006) recorded P. jekeli for French Guiana and Brazil (state of Para ). Remarks. P. jekeli belongs to the P. crustatus group, being the sister group of P. brevis by the synapomorphies mentioned above. It differs from P. brevis by the dorsal vestiture with dark green scales plus the autapomorphies: 1) mentum transversely carinate; 2) segments of antennal club robust and elongate; 3) prosternum transversely raised; and 4) glabrous area of mesofemora with two coarse punctures.

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Fig. 9. Geographical distribution of P. brevis (black squares) and P. jekeli (black triangles).

Discussion The cladistical analysis based on 24 terminal taxa and 50 characters (22 binary and 28 multistate) performed with heuristic searches of NONA, with 5,000 random-taxon-addition replicates, Deltran, Acctran, and Unambiguous optimizations, resulted in a single cladogram (Fig. 8) with 117 steps, consistency index (CI) 0.76, and retention index (RI) 0.90. This result differed from the previous analysis by only one additional step and an insignicant change in CI (see Mermudes and Napp (2006); 116 steps with CI 0.77]. The apomorphies indicated in cladogram (Fig. 8) using Deltran (or slow transformation) procedure for resulting ambiguous optimizations in Acctran already mentioned by Mermudes and Napp (2006). The main difference occurred with respect to character 34 (ventrite I of males with a setiferous sex patch). In the males of P. brevis and P. jekeli, the setiferous sex patch is closer to the posterior margin of ventrite I, being small in P. brevis and more developed in P. jekeli. Here, for both species, the state (1) was plotted in the data matrix. The analysis conrmed this state for the ancestor of these two species. The occurrence of this character independently in P. jordani and P. mixtus corroborates the homoplastic nature of this state as discussed by Mermudes and Napp (2006). Therefore, the index now changes for this character (CI 50 and RI 75). The following characters were conrmed for these species: 112, extension of scrobe shallow to deep in both sexes; 160, antennal segments IX and X neither grooved nor depressed; 170, segments IVVIII in an-

tennae of males, not grooved; 332, ventrite I with an anterior groove represented by a narrow line (only in P. brevis); 341, setiferous sex patch as discussed above; and 362, glabrous area on the outer apical area of profemora with a single coarse puncture. Unfortunately, it was not possible to dissect the only known males of these species, and some characters of the male genitalia could be not conrmed. Acknowledgments
We thank Helen Perrien (MNHN) and the curators of other institutions for help with material. This study was supported by Fundac a o de Amparo a ` Pesquisa do Estado de Sa o Paulo (FAPESP), Fundac a o de Amparo a ` Pesquisa do Estado do Rio de Janeiro (FAPERJ), and Conselho de Aperfeic oamento de Pessoal de N vel Superior (CAPES).

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Jordan, K. 1894. On Anthribidae in the museum of Honourable Walter Rothschild. Nov. Zool. 1: 591 651. Lacordaire, T. 1866. Histoire Naturelle des Insectes. Genera des Cole opte ` res, vol. 7. Roret, Paris, France. Mermudes, J.R.M. 2002. Systaltocerus platyrhinus Labram & Imhoff, 1840: redescric o es e considerac o es sobre a sinon mia com Homalorhamphus vestitus Haedo Rossi & Viana, 1957 (Coleoptera, Anthribidae, Anthribinae). Revta Bras. Entomol. 46: 579 590. Mermudes, J.R.M. 2003. Unanthribus, um novo ge nero Neotropical de Ptychoderini Jekel, 1855 (Coleoptera, Anthribidae, Anthribinae). Revta Bras. Entomol. 47: 239 244. Mermudes, J.R.M. 2005. Revisa o Sistema tica, Ana lise Clad stica e Biogeograa dos ge neros Tribotropis e Hypselotropis (Coleoptera, Anthribidae, Anthribiane, Ptychoderini). Revta Bras. Entomol. 49: 465511. Mermudes, J.R.M., and D. S. Napp. 2004. Duas novas espe cies de Ptychoderes Schoenherr, 1823 (Coleoptera, Anthribidae, Anthribinae). Revta Bras. Entomol. 48: 2730.

Mermudes, J.R.M., and D. S. Napp. 2006. Revision and cladistic analysis of the genus Ptychoderes Schoenherr, 1823 (Coleoptera, Anthribidae, Anthribinae, Ptychoderini) Zootaxa 1182: 1130. Nixon, K. C. 1999. Winclada. (beta) version 0.9.99 m24. Ithaca, NY. Nixon, K. C., and J. M. Carpenter. 1993. On outgroups. Cladistics 9: 413 426. Page, R.D.M. 2001. NDE (NEXUS data [ed.], for windows). Version 0.5.0. NDE. (http://taxonomy.zoology.gla.ac.uk/ rod/NDE/nde.html). Schoenherr, C. J. 1839. Genera et species Curculionidum cum synonymia hujus familiae: species novae aut hactenus minus cognitae, descriptionibus a Dom. Leonardo Gyllenhal, C. H. Boheman, et entomologis aliis ilustratae. 5: 1 456. Received 28 January 2010; accepted 20 April 2010.

Appendix List of characters used in the analysis (see Mermudes and Napp (2006) to discussion of states of characters). 1. Dorsal Vestiture. Without dorsal vitta (0); with a pale vitta formed by whitish scales intermingled with colored scales, extending medially on rostrum, pronotum and elytra (1); with a yellowish vitta on rostrum, sides of vertex and pronotum, reaching humeri (2). 2. Fusiform Scales on Elytra. Absent (0); present (1). 3. Dark Brownish Scales Spots at the Anterior Angles of Ventrites. absent (0); developed, on ventrites IV (1); small, on ventrites IIIV (2). 4. Vestiture of the Sides of Prosternum, below Lateral Carina. without vitta (0); with a dark vitta narrower posteriorly (1); with a dark vitta not narrowed posteriorly (2); with a vitta concolorous with the dorsal vestiture (3). 5. Small, Centered Spots on Outer Side of Meso- and Metafemora. Absent (0); two spots, one near dorsal side and one near ventral side (1); two spots connected forming a posmedian, narrow stripe (2). 6. Narrow Antebasal Stripe Plus a Posmedian, Subtriangular Spot on Tibiae. Absent (0); moderately developed (1); developed (2). 7. Apical Notch of Rostrum. Absent (0); deep and angulated (1); deep and almost rounded (2). 8. Dorsum of Rostrum. Longitudinally grooved (0); weakly depressed (1). 9. Rostrum. Shaped otherwise (0); Ptychoderes like (1). 10. Rostrum, dorsolateral carinae thickened and rounded on top. Absent (0); straight, close and subparallel to median carina (1); convergent, far from median carina at base of rostrum (2). 11. Extension of Scrobe. absent or vestigial in both sexes (0); vestigial in males and shallow in females (1); shallow to deep in both sexes (2).

12. Mentum. Shaped otherwise (0); Ptychoderes like (1). 13. Frons. Not carinate at sides (0); carinate (1). 14. Ommatidial Size. moderately ne (0); ne (1). 15. Antennae of male. Segments IIV thickened, cylindrical and depressed. absent (0); present (1). 16. Antennal Segments IX and X in Both Sexes. Neither grooved nor depressed (0); either depressed or grooved (1). 17. Segments IVVIII, in Males. Not grooved (0); grooved (1). 18. Central Depression on Pronotum. Absent (0); shallow, large, extending throughout the disk (1); posmedian (2); centrolongitudinal, deeper behind median tubercle (3). 19. Rugae of Pronotum. Absent (0); present throughout (1); present only inside the central depression (2). 20. Pronotum. Without tubercle or prominence (0); with a central prominence (1); with an antemedian prominence inside a depression (2); with a transverse, median tubercle inside a depression (3); with a central, rounded tubercle in the depression, as prominent as the margins of depression (4); with a central, rounded tubercle in the depression, less prominent than the margins of depression (5); with a central, rounded tubercle in the depression, at on top (6). 21. Lateral Carina of Prothorax. Short (0); elongate, straight (1); elongate, slightly sinuous (2); elongate, strongly sinuous (3). 22. Laterobasal Carina of Prothorax. Absent (0); weakly prominent (1); strongly prominent (2). 23. Prosternum. Not depressed or furrowed (0); depressed all over the posterior half (1); furrowed in front of procoxae, anterior margin broadly curved (2); furrowed in front of procoxae, anterior margin straight to sinuous (3).

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24. Prosternal Furrow Shallow. Absent (0); with a moderately developed projection at middle (1); sinuous (2); vestigial (3). 25. Prosternal Furrow Deep. Absent (0); with a developed projection at middle (1). 26. Radial Sector of the Wings. Absent (0); present (1). 27. Basal Two Thirds of Elytra. Not depressed (0); depressed along interstria two (1); depressed between suture and interstria three (2). 28. Basal Region of Elytra. Not raised (0); raised (1). 29. Elytra Behind the Basal Gibbosities. Not impressed (0); transversely impressed (1). 30. Elytral Surface. Not rugose (0); transversely rugose throughout basal region (1); transversely rugose on humeri, inconspicuously so near suture (2); inconspicuously rugose (3). 31. Apical Declivity of Elytra Swollen or Tuberculate between Interstriae 3 and 9. Absent (0); with interstriae three and nine swollen and linked (1); with interstriae 3, 5, 7, and 9 swollen (2); with rounded tubercles (3); with subtransverse tubercles (4). 32. Pygidium. Varied in shape, longitudinally convex or strongly depressed (0); longer than wide, depressed, slightly convergent at sides (1); as long as wide, weakly depressed (2); wider than long and convex (3). 33. Groove of Anterior Margin of Ventrite I. Shallow and wide (0); deep and wide (1); line shaped (2); vestigial (3). 34. Ventrite I of Males. Without setiferous sex patches (0); with a small setiferous sex patch (1); with a developed setiferous sex patch (2). 35. Ventrite V of Females. Not expanded at sides (0); expanded (1). 36. Glabrous Area on Profemora. Absent (0); present, nely punctate (1); present, with a single coarse puncture (2). 37. Glabrous Area on Meso- and Metafemora. Absent (0); present (1).

38. Tarsomere I. Longer than II (0); subequal to II (1); longer than II. 39. Tergite VIII of Males. Subquadrangular, apical margin sinuous (0); almost subparallel at sides, apical margin subtruncate or rounded (1); convergent at sides from base, apical margin rounded (2). 40. Aedeagus, Bridge between Apodemes. Absent (0); present (1). 41. Aedeagus, Arc between Apodemes. Absent (0); present (1). 42. Pedon, Body. Strongly or weakly constricted, truncate at apex (0); gradually attenuate apically, apex rounded (1). 43. Internal Sac. Pigmented rods of lateral membrane: absent (0); present (1). 44. Median Sclerites of the Internal Sac. Absent (0); present (1). 45. Basal Sclerites of the Internal Sac. Absent (0); present (1) (CI 100; RI 100). 46. Tergite VIII of Females. About as long as wide, with short, sparse pilosity (0); evidently longer than wide, with dense, elongate pilosity at sides (1). 47. Toothed Plate of the Ovipositor. with three or four teeth, the distal and proximal ones variously shaped (0); with four teeth: two subequal, developed, conchoidal distal teeth, one proximal, dorsolateral, moderately developed, another proximal, ventrolateral, very small (1); with three teeth moderately developed, successively smaller from basal to apical tooth (2). 48. Ovipositor, Median Rods. Thickened, rounded at proximal margin (0); slender, acuminate at proximal end (1). 49. Ovipositor, Ventral Median Lobe. Bilobate (0); acuminate (1). 50. Rectal Loop. Without dorsal projection (0); with a weak dorsal projection (1); with a strong dorsal projection (2).