Zootaxa, an Essay on the Tribe Xylographellini (Coleoptera

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An essay on the tribe Xylographellini (Coleoptera: Tenebrionoidea: Ciidae)

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An essay on the tribe Xylographellini (Coleoptera: Tenebrionoidea: Ciidae)

CRISTIANO LOPES-ANDRADE

Magnolia Press Auckland, New Zealand CRISTIANO LOPES-ANDRADE An essay on the tribe Xylographellini (Coleoptera: Tenebrionoidea: Ciidae) (Zootaxa 1832) 110 pp.; 30 cm. 30 July 2008 ISBN 978-1-86977-241-3 (paperback) ISBN 978-1-86977-242-0 (Online edition)

FIRST PUBLISHED IN 2008 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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2 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE Zootaxa 1832: 1–110 (2008) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2008 · Magnolia Press ISSN 1175-5334 (online edition)

An essay on the tribe Xylographellini (Coleoptera: Tenebrionoidea: Ciidae)

CRISTIANO LOPES-ANDRADE Programa de Pós-Graduação em Entomologia, Departamento de Biologia Animal, Universidade Federal de Viçosa, 36570-000, Viçosa, MG, Brasil; e-mail: [email protected]

Table of contents

Abstract ...... 4 Resumo ...... 4 Resúmen ...... 4 Introduction ...... 5 Material and Methods ...... 5 Tribe Xylographellini Kawanabe & Miyatake, 1996 (Figs 1–159) ...... 9 Identification key to the subtribes and genera of Xylographellini ...... 9 Subtribe Xylographellina Kawanabe & Miyatake, 1996 (Figs 1–6, 9–10, 13–135, 159) ...... 9 Xylographella Miyatake, 1985 (Figs 5, 9, 13–26) ...... 10 Identification key to the species of Xylographella Miyatake ...... 10 Xylographella punctata Miyatake, 1985 (Figs 5, 9, 13–15, 19, 21A, 22A, 23–24, 26) ...... 11 Xylographella speciosa Lopes-Andrade sp. nov. (Figs 16–18, 20, 21B, 22B, 25–26) ...... 11 Scolytocis Blair, 1928 (Figs 1–4, 6, 10, 27–135) ...... 12 Identification keys to the species of Scolytocis Blair ...... 14 Northern Neotropical species of Scolytocis Blair ...... 16 Scolytocis bouchardi Lopes-Andrade sp. nov. (Figs 27–29, 54, 63A, 66A, 69, 80) ...... 16 Scolytocis cariborum Lopes-Andrade sp. nov. (Figs 30–32, 55, 63B, 66B, 70, 81) ...... 17 Scolytocis danielssoni Lopes-Andrade sp. nov. (Figs 33–35, 56, 63C, 66C, 80) ...... 19 Scolytocis difficillimus Lopes-Andrade sp. nov. (Figs 36–38, 57, 64A, 67A, 71, 80) ...... 20 Scolytocis howdeni Lopes-Andrade sp. nov. (Figs 39–41, 58, 64B, 67B, 72, 80) ...... 21 Scolytocis indecisus Lopes-Andrade sp. nov. (Figs 42–44, 59, 64C, 67C, 73, 80) ...... 22 Scolytocis kiskeyensis Lopes-Andrade sp. nov. (Figs 45–47, 60, 65A, 68A, 74–76, 81) ...... 24 Scolytocis lawrencei Lopes-Andrade sp. nov. (Figs 48–50, 61, 65B, 68B, 77–78, 80) ...... 25 Scolytocis panamensis Lopes-Andrade sp. nov. (Figs 51–53, 62, 65C, 68C, 79, 80) ...... 27 Southern Neotropical species of Scolytocis Blair ...... 28 Scolytocis fritzplaumanni Lopes-Andrade sp. nov. (Figs 82-84, 91, 94A, 95A, 96-97, 101) ...... 28 Scolytocis furieriae Lopes-Andrade sp. nov. (Figs 10, 85–87, 92, 94B, 95B, 98, 101) ...... 29 Scolytocis paschoali Lopes-Andrade sp. nov. (Figs 1–4, 88–90, 93, 94C, 95C, 99–101) ...... 31 Indo-Malayan, Australasian and Oceanic species of Scolytocis Blair ...... 32 Scolytocis malayanus Lopes-Andrade sp. nov. (Figs 102-104, 120, 126A, 128A, 130, 134) ...... 32 Scolytocis novaezelandiae Lopes-Andrade sp. nov. (Figs 105–107, 121, 126B, 128B, 135) ...... 33 Scolytocis philippinensis Lopes-Andrade sp. nov. (Figs 108–110, 122, 126C, 128C, 131, 134) ...... 35 Scolytocis samoensis Blair, 1928 ...... 36 Scolytocis thayerae Lopes-Andrade sp. nov. (Figs 111–113, 123, 127A, 129A, 132, 134) ...... 36 Scolytocis werneri Lopes-Andrade sp. nov. (Figs 114–116, 124, 127B, 129B, 133, 134) ...... 37 Scolytocis zimmermani Lopes-Andrade sp. nov. (Figs 117–119, 125, 127C, 129C, 135) ...... 39 Syncosmetina Lopes-Andrade subtr. nov. (Figs 7–8, 11–12, 136–159) ...... 40 Syncosmetus Sharp, 1891 (Figs 11, 113–122) ...... 40 Identification key to the species of Syncosmetus Sharp ...... 40 Syncosmetus japonicus Sharp, 1891 (Figs 11, 136–138, 142A, 143A, 144–145) ...... 41 Syncosmetus reticulatus Miyatake, 1985 (Figs 7, 139–141, 142B, 143B, 145) ...... 41 Tropicis Scott, 1926 (8, 12, 146–158) ...... 42

Accepted by D. Pollock: 11 Apr. 2008; published: 30 Jul. 2008 3 Identification key to the species of Tropicis Scott ...... 42 Tropicis brevicarinatus Scott, 1926 ...... 42 Tropicis cuccodoroi Lopes-Andrade sp. nov. (Figs. 146–148, 152, 154A, 155A, 156, 158) ...... 42 Tropicis flexicarinatus Scott, 1926 ...... 44 Tropicis sexcarinatus (Waterhouse, 1876) (Figs 8, 12, 149–151, 153, 154B, 155B, 157–158) ...... 44 Discussion ...... 45 Acknowledgements ...... 49 References ...... 49

Abstract

The tribe Xylographellini is redefined and three genera are formally transferred to it: Scolytocis Blair, to the subtribe Xylographellina; Syncosmetus Sharp and Tropicis Scott, to the subtribe Syncosmetina subtr. nov. The genus Scolytocis is redescribed and diagnoses are provided for the other genera of the tribe. Twenty new species are described: Xylographella speciosa sp. nov.; Scol. bouchardi sp. nov.; Scol. cariborum sp. nov.; Scol. danielssoni sp. nov.; Scol. difficillimus sp. nov.; Scol. fritzplaumanni sp. nov.; Scol. furieriae sp. nov.; Scol. howdeni sp. nov.; Scol. indecisus sp. nov.; Scol. kiskeyensis sp. nov.; Scol. lawrencei sp. nov.; Scol. malayanus sp. nov.; Scol. novaezelandiae sp. nov.; Scol. panamensis sp. nov.; Scol. paschoali sp. nov.; Scol. philippinensis sp. nov.; Scol. thayerae sp. nov.; Scol. werneri sp. nov.; Scol. zimmermani sp. nov.; and Trop. cuccodoroi sp. nov. A key to the subtribes and genera of Xylographellini is provided, as well as keys to the species of Scolytocis, Syncosmetus, Tropicis and Xylographella Miyatake. The taxonomic limits and geographic distribution of the tribe, its subtribes and genera are discussed in light of the new data provided here. The morphological similarities between Xylographellini and Orophiini are briefly discussed.

Key words: Ciidae, Xylographellini, new subtribe, new species, Oriental, Afrotropical, Neotropical

Resumo

A tribo Xylographellini é redefinida e três gêneros são formalmente transferidos para ela: Scolytocis Blair, para a subtribo Xylographellina; Syncosmetus Sharp e Tropicis Scott, para a subtribo Syncosmetina subtr. nov. O gênero Scolytocis é redescrito e diagnoses são fornecidas para os outros gêneros da tribo. Vinte espécies novas são descritas: Xylographella speciosa sp. nov.; Scol. bouchardi sp. nov.; Scol. cariborum sp. nov.; Scol. danielssoni sp. nov.; Scol. difficillimus sp. nov.; Scol. fritzplaumanni sp. nov.; Scol. furieriae sp. nov.; Scol. howdeni sp. nov.; Scol. indecisus sp. nov.; Scol. kiskeyensis sp. nov.; Scol. lawrencei sp. nov.; Scol. malayanus sp. nov.; Scol. novaezelandiae sp. nov.; Scol. panamensis sp. nov.; Scol. paschoali sp. nov.; Scol. philippinensis sp. nov.; Scol. thayerae sp. nov.; Scol. werneri sp. nov.; Scol. zimmermani sp. nov.; e Trop. cuccodoroi sp. nov. Uma chave para as subtribos e gêneros de Xylographellini é fornecida, assim como chaves para as espécies de Scolytocis, Syncosmetus, Tropicis e de Xylographella Miyatake. Os limites tax- onômicos e distribuição geográfica da tribo, subtribos e gêneros são discutidos a partir dos novos dados apresentados aqui. As similaridades morfológicas entre Xylographellini e Orophiini são brevemente discuticas.

Palavras-chave: Ciidae, Xylographellini, nova subtribo, novas espécies, Oriental, Afrotropical, Neotropical

Resúmen

Se redefine a la tribu Xylographellini y se transfieren a tres géneros hacia ella: Scolytocis Blair, a la subtribu Xylographellina; Syncosmetus Sharp y Tropicis Scott, a la subtribu Syncosmetina subtr. nov. Se redescribe al género Scolytocis y se presentan diagnosis para los otros géneros de la tribu. Se describen veinte nuevas especies: Xylographella speciosa sp. nov.; Scol. bouchardi sp. nov.; Scol. cariborum sp. nov.; Scol. danielssoni sp. nov.; Scol. difficillimus sp. nov.; Scol. fritzplaumanni sp. nov.; Scol. furieriae sp. nov.; Scol. howdeni sp. nov.; Scol. indecisus sp. nov.; Scol. kiskeyensis sp. nov.; Scol. lawrencei sp. nov.; Scol. malayanus sp. nov.; Scol. novaezelandiae sp. nov.; Scol. panamensis sp. nov.; Scol. paschoali sp. nov.; Scol. philippinensis sp. nov.; Scol. thayerae sp. nov.; Scol. werneri sp. nov.; Scol. zimmermani sp. nov.; y Trop. cuccodoroi sp. nov. Se presenta una clave para las subtribus y géneros de Xylographellini, y claves para las especies de Scolytocis, Syncosmetus, Tropicis y Xylographella Miyatake. Se discuten los límites tax- onómicos y distribución geográfica de la tribu, sus subtribus y géneros, a partir de nuevos datos presentados en este tra- bajo. Las similitudes morfológicas entre Xylographellini y Orophiini son presentadas y discutidas.

Introduction

The tribe Xylographellini (Ciidae: Ciinae) was established by Kawanabe & Miyatake (1996) for the genus Xylographella Miyatake, comprising only the type species from Japan. Along with Sphindociinae, Xylographellini is regarded as being monospecific and both are the unique suprageneric Ciidae taxa defined by robust morphological characters (Lawrence 1974; Kawanabe & Miyatake 1996; Lopes-Andrade 2007a, b). However, while no other Sphindociinae species came to light in the last decades, many undescribed Xylographellini species are known from museum and private collections. Moreover, other described Ciidae genera that possess the diagnostic characters of Xylographellini should be included in the tribe. One of these genera, the monospecific Scolytocis Blair, was included in the tribe by Kawanabe and Miyatake (1996) with some doubt. Besides it, the genera Syncosmetus Sharp and Tropicis Scott also share some characters with Xylographella and Scolytocis. In the description of Scolytocis, Blair (1928) cited that it resembles Tropicis in the "short antennae with broad compact club, and also in the denticulations of the outer edge of the tibiae". The drawings and descriptions of Tropicis by Scott (1926) and Syncosmetus Sharp by Miyatake (1954, as Omogocis Miyatake) provided important cues on the morphological similarities of both genera, and also on their relationship with Scolytocis and Xylographella. Kawanabe (2003) recognized some similarities of Syn- cosmetus and Xylographella, mainly regarding the morphology of mouthparts and male pregenital segments. However, he has neither included Syncosmetus in any existing tribe of Ciidae nor described any new suprageneric taxon for it. In the last nine years, large series of new Scolytocis species were collected in Brazil, and other specimens were made available from private and institutional collections. A good amount of material was brought together (including specimens of Syncosmetus, Tropicis and Xylographella), allowing comparative studies, and clarification of the generic and suprageneric relationships of Xylographellini. I do not intend to provide an exhaustive taxonomic work here, but to make available some important information on the distribution and morphology of this barely known tribe. Xylographellini is the unique widespread and speciose major group of Ciidae not treated in the first revision of the family by Mellié (1849), and the importance of this tribe has been concealed by the fact that it is currently believed to be monospecific. There are certainly other new Xylographellini that I have not examined for this work. However, at this moment, I prefer to accomplish this work, transferring some taxa to Xylographellini, describing several new species and providing morphological descriptions to allow the adequate recognition of the tribe by other researchers. In this work, I formally include Scolytocis in Xylographellini. I also transfer Syncosmetus and Tropicis to the tribe (in the new subtribe Syncosmetina subtr. nov.), redefine Xylographellini and provide identification keys for its genera and subtribes, and for the species of each genera. Twenty new species (19 of Xylographell- ina and one belonging to Syncosmetina subtr. nov.) are described, and six morphological species-groups are proposed for the species of Scolytocis. The morphology, host-use and geographic distribution of Xylographellini are briefly discussed in light of the new information provided here.

Material and Methods

Morphological examinations More than 1,000 specimens were examined, belonging to 24 species: four already described and 20 described here as new. Among the 27 Xylographellini species dealt with in this work, I could not examine three: Scol. samoensis Blair; Trop. brevicarinatus Scott; and Trop. flexicarinatus Scott. Scolytocis samoensis is the unique previously described species of the genus, and therefore the type species by monotypy. Even without examining it, I have decided to include 18 new species in the genus as they fit the description of Blair

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 5 (1928). Moreover, the genus is easily recognizable among the Xylographellini as it is unique in possessing nine antennomeres (rather than ten). Around 80 specimens were dissected, including specimens of all available species, and permanent slide preparations followed the protocol described by Lopes-Andrade et al. (2003a). In some cases, female terminalia were stained with Chlorazol Black E (Direct Black 38, C.I. No. 30235) following the protocol of Carayon (1969). Examination and photography of slide preparations were made under an Olympus BX41 or a Leica DM 2500 compound microscope. Measurements of body length, pronotum and elytra were taken under an MBS-9 stereomicroscope with a scale ocular. Measurements of antennae, hindwings and genitalia were taken under a Leica DM 2500 with a scale ocular. Final comparisons and descriptions of general external morphology were made under a Leica MZ16. Digital photographs of slide preparations and holotypes were taken with a Canon S70. Final images of dorsal, lateral and ventral surfaces of body were the result of joining 20 to 40 photographs in different focus using the automontage freeware CombineZM (Hadley 2006). Scanning Elec- tron micrographs (SEMs) were obtained using a LEO 435VP under variable pressure (which does not require prior metal coating). Images were arranged and standardized using Corel Photo Paint and Corel Draw X3 software. Terms for external morphology, male genitalia and pregenital segments of ciids are explained and discussed by Lopes-Andrade & Lawrence (2005). The terms mesoventrite and metaventrite (introduced by Lawrence 1999) have been used in place of the misapplied terms mesosternum and metasternum (see Lopes- Andrade & Lawrence 2005; Lopes-Andrade 2007a). Terms for female terminalia are those used by Lopes- Andrade (2007a, b). The term “sex patch” is used to refer to the structure present in the first abdominal ventrite of most male ciids (Faustini & Halstead 1982). In some cases, the number of available specimens of each locality was low, and so I preferred not to dissect all specimens in an attempt to find a male: the specimens could be damaged during dissection, with the risk of not finding any male. I did not dissect any specimens chosen as holotype, so their gender could not be determined with accuracy. While separating the species of this group, I have given priority first to characters of the exposed surface of body, then to the morphology of male genitalia. The male genitalia was very small and barely sclerotized in most species examined and, in some cases, of little utility for taxonomic purposes. However, in cases when the male genitalia was well sclerotized or modified enough, characters of this structure were used for the diagnosis of the species or even for the identification keys. Among the new species of Scolytocis, I found males of 15 species and describe their genitalia here. Males of three Scolytocis species are unknown (or were not dissected), as follows: Scol. danielssoni sp. nov., Scol. novaezelandiae sp. nov. and Scol. zimmermani sp. nov. These three species are sufficiently different to allow recognition by other external morphological features. Males of Tropicis cuccodoroi sp. nov. were dissected and their genitalia examined, and comparisons were made with the male genitalia of the other available species of the genus, Trop. sexcarinatus (Waterhouse). The male genitalia and female terminalia of Sync. japonicus Sharp could also be examined. Unfortunately, I had only one female Sync. reticulatus Miyatake for examination, which was completely dissected, with no males for comparison. Two females and one male of Xyl. punctata Miyatake were dissected. The female terminalia of Xylographellini vary little between species of the same genus, but the comparison of this structure revealed some important characters, which are diagnostic at the generic level. Therefore, I provide images of only one species of each genus (Figs 9–12). A maximum of ten specimens from each locality were measured. For each species, the range, mean and standard deviation are given for measurements and ratios of a subsample or the whole type series (including the holotype). Measurements of antennomeres were taken only from the holotypes and are provided in each description. The following abbreviations are used for measurements and ratios: TL, total length (including head measured from above); PL, pronotal length along midline; PW, greatest pronotal width; EL, elytral length (median length from base of scutellum to elytral apex); EW, greatest elytral width; GD, greatest depth; FL, antennal funicle length; CL, antennal club length. The ratio GD/EW was taken as an indication of degree

6 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE of convexity; TL/EW indicates degree of body elongation. Species descriptions are based on the holotype that, in each case, is a fully pigmented adult. All characters mentioned in each description refer to the holotype, except characters of hindwing and genitalia that were all observed in paratypes. Measurements of tibiae and antennae are of holotypes, but the corresponding figures are of paratypes. Differences among paratypes, if any, are given in the sections on Variation, together with standard measurements and ratios of the holotype and paratypes. Images of antennae, hindwings, metatibiae and male genitalia are of paratypes from the type locality, unless otherwise specified.

Species boundaries Besides other external morphological characters, I carefully compared the male genitalia (whenever possible), antennae, hindwings and tibiae of all the subsamples of the new species described here. The morphology of these structures was consistent throughout the range of each species (in case of species known from more than one locality and/or field collection). The comparison of subsamples of widely distributed species (mainly of the genus Scolytocis) allowed me to recognize characters that vary more interspecifically than intraspecifically in the Xylographellini genera. Therefore, I could better characterize species known from only one locality. Species boundaries were judged only by morphological criteria, and the geographic distributions of the species were interpreted a posteriori (to avoid redundancy while discussing the distribution of the Xylographellini taxa). When there were doubts on the boundaries of two or more species, I preferred to treat and describe them as separate taxa. Further field collections, mainly in the known geographic gaps, would clarify these matters. Among all the specimens examined, only two specimens of Scolytocis could not be included in any of the species described here, as follows: one specimen (CNCI) from Ecuador labeled /ECU. Limoncocha Napo Prov. 250m 22.VI.76, Ber. 356, S&J Peck / Scolytocis sp. [handwritten] C. Lopes-Andrade det. 2007 [printed]/; one specimen (CMNC) from Colombia labeled /3000', Rio Jamundi, 10mi.S. Cali, Valle, Colom- bia, II.16.1970 Henry F. Howden / Scolytocis sp. [handwritten] C. Lopes-Andrade det. 2008 [printed]/. In both cases, the specimen was damaged and no other specimen was known from the locality or its vicinity. However, as the tribe and genus could be determined with accuracy, the locality data of these specimens were included in the distributional map of Xylographellini (Fig. 159).

Identification keys The identification keys provided here are based mostly on external and exposed morphological characters of adults, and work for both male and female. I attempted to include the most robust and easiest to observe features in the identification key to the subtribes and genera. I do not provide a key to the major Ciidae taxa because that is not possible at this moment. Such a key would work well for regional faunas, but not for the worldwide ciid fauna. Moreover, I intend to make changes in the suprageneric classification of Ciidae and describe some genera in forthcoming works, and then provide a key to the suprageneric and generic taxa of the family. The genera Syncosmetus, Tropicis and Xylographella are less speciose, though I provide one key for each genus. The key for Tropicis is based on the one provided by Scott (1926), with modifications and the addition of a new species. Scolytocis is the most speciose and widespread genus of the tribe, so I arrange the species in major geographic regions and provide two separate keys: one for the twelve Neotropical species; another for the Indo-Malayan, Australasian and Oceanic species (not including Scol. samoensis Blair). Although all the Neotropical species are included in one joint identification key, I treat these species as two separate biogeographic groups in the text: the northern Neotropical; and the southern Neotropical Scolytocis. The reason for this is that the southern Neotropical Scolytocis species constitute a very particular group, without close morphological affinities to the other faunas. The northern Neotropical species have several affinities to the Indo-Malayan and Australasian faunas. Moreover, there is a wide geographic gap between the northern

Pin label data: host-fungi and localities Data of pin labels, given in the section on Specimens examined, are cited ipsis litteris. Data of each label are delimited by a forward slash. All labels are printed on white paper, unless otherwise specified. If a label is mixed printed and handwritten, or has a distinct colour (yellow or red) the information is cited inside square brackets immediately following the label wording. Information on host fungi from pin label data were updated following the database Species Fungorum (Index Fungorum Partnership 2007) available online, and are given in the sections on Host-fungi. Geographic coordinates were estimated for all the locality data available from the literature and from pin label data of the specimens examined. Two main data sources were consulted (Global Gazetteer 2007; Google Earth 2007), and the estimated geographic coordinates were plotted in distributional maps using the software ArcMap 9.0. Most of the terms used here for major biogeographic areas are those adopted by Olson et al. (2001). However, in the case of Latin America and the Caribbean Islands, I followed the biogeographic areas and transition zones discussed by Morrone (2006) based on analyses of the entomofauna. In both cases, my choices were arbitrary and are used here only to illustrate and discuss the major distributional patterns of Xylographellini. The term “Region” here is used as a synonym of “Ecozone” and “Realm”.

Depositories Specimens have been deposited in (or belong to) the following institutional and private collections, (with acronyms used in this paper):

ANIC Australian National Insect Collection, CSIRO Entomology (Canberra, Australia) BPBM Bernice P. Bishop Museum (Honolulu, Hawaii, USA) CAMB Coleção Ayr de Moura Bello (Rio de Janeiro, RJ, Brazil) CMNC Canadian Museum of Nature (Ottawa, Ontario, Canada) CNCI Canadian National Collection of Insects (Ottawa, Ontario, Canada) FMNH Field Museum of Natural History (Chicago, Illinois, USA) GOPC Glenda Orledge Private Collection (Bath, UK) ISNB Institut Royal des Sciences Naturelles de Belgique (Bruxelles, Belgium) KMMA Koninklijk Museum voor Midden Afrika (Tervuren, Belgium) LAPC Cristiano Lopes-Andrade Private Collection (Viçosa, MG, Brazil) MCNZ Museu de Ciências Naturais da Fundaçao Zoobotânica (Porto Alegre, RS, Brazil) MCZ Museum of Comparative Zoology, Harvard University (Cambridge, Massachusetts, USA) MHNG Muséum d'Histoire Naturelle (Geneva, Switzerland) MNNC Museo Nacional de Historia Natural (Santiago, Chile) MZLU Museum of Zoology, Lund University (Lund, Sweden) MZSP Museu de Zoologia da Universidade de São Paulo (São Paulo, SP, Brazil) RRPC Rafal Ruta Private Collection (Poland) SMTD Staatliches Museum für Tierkunde (Dresden, Germany) UFES Coleção Entomológica da Universidade Federal do Espírito Santo (Vitória, ES, Brazil)

The holotypes of the new Brazilian species are deposited in MZSP. Holotypes of the species from the Philippines belong to the FMNH. The holotypes of Scol. bouchardi sp. nov. and Scol. howdeni sp. nov. belong to the CNCI, and the ones of the remaining new Scolytocis belong to the ANIC. The holotype of Trop- icis cuccodoroi sp. nov. belongs to the MHNG.

Included taxa Xylographella Miyatake, 1985 (type genus) Scolytocis Blair, 1928 Syncosmetus Sharp, 1891 Tropicis Scott, 1926

Diagnosis and comments Xylographellini may be distinguished from the other Ciidae by the following combination of characters: (i) each antenna with seven (antennomeres of club fused; Fig. 65C), or nine to ten antennomeres (club with three antennomeres; Figs 2, 21, 63A–65B, 94, 126–127, 142, 154); (ii) antennal club distinctly compact, at least 1.2X longer than the funicle; (iii) at least the apical antennomere bearing more than four sensillifers (Fig. 2), usually more than seven, sometimes coalescent and difficult to be counted; (iv) insertion of labial palpi near the middle of prementum (Fig. 5); (v) prementum with a longitudinal sulcus in frontal view (Fig. 3); (vi) apical labial palpomere widened (Figs 5–8); (vii) prosternum short, being concave, biconcave, flat or slightly tumid; (viii) each tibia usually expanded near the middle or the apex, bearing spines on its outer margin (Figs 22, 66–68, 95, 128–129, 143, 155); (ix) ninth segment of male abdomen (pregenital segment) always Y- shaped (Figs 24–25, 69, 73, 76, 78–79, 96–99, 130–132, 144,156–157); (x) male genitalia with tegmen and median lobe elongate, cylindrical, at least 4X longer than large; (xi) absence of conspicuous secondary sexual characters in male, except for a small sex patch in some species (in Syncosmetina subtr. nov.); (xii) female terminalia (Figs 9–12) with gonocoxites divided in two parts, always longer than paraprocts; (xiii) gonostyli usually absent (Figs 9–10, 12), and small gonostyli present only in Syncosmetus (Fig. 11). Characters ii to vi and ix are restricted to Xylographellini. Character viii in Xylographellini and Orophiini. Character xii occurs in the latter two tribes and also in some genera formally included in Ciini (Ceracis Mel- lié, Phellinocis Lopes-Andrade & Lawrence, Sulcacis Dury and related genera), which shows that Ciini may be polyphyletic. In the other hand, the female terminalia of the Ciini genera close related to Cis Latreille (Ennearthron Mellié, Strigocis Dury, Falsocis Pic and so on) all have the basic morphology found in many tenebrionoid families (Tanner 1927; Tschinkel & Doyen 1980; Banaszkiewicz 2006; Lopes-Andrade 2007a, b).

Identification key to the subtribes and genera of Xylographellini

1 Width of scutellum at base more than 0.10X the greatest elytral width (EW). Prosternal process laminate. Procoxae subconical, extending below the plane of the prosternal process. Tibiae (Figs 22, 66–68, 95, 128–129) with outer edge broadly rounded and/or tibiae distinctly expanded to apex; spines conspicuous...... 2 (Xylographellina Kawanabe & Miyatake) - Width of scutellum at base less than 0.10X the greatest elytral width (EW). Prosternal process broad. Pro- coxae subcylindrical and at the same level of the prosternal process. Tibiae (Figs 143, 155) not conspicuously expanded to apex; outer edge of tibiae straight or slightly rounded, bearing few small, inconspicuous spines ...... 3 (Syncosmetina Lopes-Andrade subtr. nov.) 2 Antennal funicle with five antennomeres (Fig. 21). Lateral margins of pronotum finely crenulated. Japan and the Philippines...... Xylographella Miyatake - Antennal funicle with four antennomeres (Figs 63–65, 94, 126–127). Lateral margins of pronotum smooth. Indo-Malayan, Australasian, Oceanic and Neotropical Region...... Scolytocis Blair 3 Body subparallel-sided. Humeri angulate. Hindwings well developed. Female terminalia devoid of gonostyli. Western Indian Ocean (Seychelles and the Mascarene Islands)...... Tropicis Scott - Body ovoid. Humeri rounded. Hindwings extremely reduced (almost absent). Female terminalia bearing gonostyli. Japan ...... Syncosmetus Sharp

Included genera Xylographella Miyatake, 1985 (type genus) Scolytocis Blair, 1928

Diagnosis This subtribe is characterized by the combination of the following characters: (i) scutellum conspicuous, its basal width more than 0.10X the greatest elytral width; (ii) prosternal process laminate; (iii) procoxae subconical, extending below the plane of the prosternal process; (iv) apex of prementum acute; (v) female terminalia devoid of gonostyli.

Distribution Xylographellina is the most widespread subtribe of Xylographellini (Figs 26, 80–81, 101, 134, 159), occurring throughout the Neotropical Region and in parts of the Palearctic (restricted to Japan), Oceanic, Aus- tralasian (restricted to New Zealand) and Indo-Malayan Regions.

Xylographella Miyatake, 1985 (Figs 5, 9, 13–26)

Included species Xylographella punctata Miyatake, 1985 (type species, by monotypy) Xylographella speciosa Lopes-Andrade sp. nov.

Diagnosis The genus Xylographella was redescribed by Kawanabe & Miyatake (1996) in the work delimiting the tribe Xylographellini, and it is not necessary to provide another redescription here. The genus is closely related to Scolytocis, but differs mainly by the combination of the following features: (i) antennal funicle with five antennomeres; (ii) lateral margins of pronotum finely crenulated; (iii) pronotum and elytra with very coarse and dense punctation, the punctures being separated by a distance of one puncture width or less.

Comments Since its description, the genus was formally known as monospecific and restricted to Japan. A new species is described here, based on specimens from the Philippines. Some species of Scolytocis may be confused with Xylographella due to their coarse pronotal and elytral punctation. However, they are easily distinguished by the diagnostic characters of the genera.

Identification key to the species of Xylographella Miyatake

1 Pronotum with interstice between punctures microreticulate. Elytral apices devoid of raised keels. Total length < 1.70mm. Japan...... Xylographella punctata Miyatake - Interstice between punctures smooth. Elytra with apical declivity bearing longitudinal raised keels. Total length> 2.05mm. Philippines...... Xylographella speciosa sp. nov.

Diagnosis Small species, TL usually less than 1.7mm. Pronotum with interstice between punctures microreticulate. Elytra devoid of raised keels.

Host fungi Fomes fomentarius (L.) J.J. Kickx (Polyporaceae) and Elfvingia applanata (Pers.) P. Karst. (Ganoder- mataceae) (Kawanabe & Miyatake 1996; Kawanabe 1998). Regarded as belonging to the Ganoderma host- use group (Orledge & Reynolds 2005).

Distribution Japan, at the islands of Honshu and Shikoku (Fig. 26).

Specimens examined Japan: 1% labeled (ANIC) /Japan: Chu-Zenji 21.vii.09 E. Gallus [handwritten] / 604 [handwritten] / Xylographella punctata Miyatake, 1985 C. Lopes-Andrade det. [printed]/, 2&& (1 ANIC, 1 LAPC) labeled / Japan: Chu-Zenji 21.vii.09 E. Gallus [printed] / Xylographella punctata Miyatake, 1985 C. Lopes-Andrade det. [printed] /; 2 specimens (LAPC) / Japan, Nara Pref. Mt. Obako-dake, 1300m near Nosegawa-mura vill. 25 iii 2006, leg. P. Jaloszynski [printed] / Xylographella punctata det. Rafal Ruta 2005 [sic] [printed] /.

Xylographella speciosa Lopes-Andrade sp. nov. (Figs 16–18, 20, 21B, 22B, 25–26)

Etymology The specific epithet is a latin adjective meaning “impressive”.

Diagnosis Large species, total length more than 2.00mm. Interstice between punctures smooth. Elytra with apical declivity bearing longitudinal raised keels.

Description Holotype. (Figs 16–18) Measurements in mm: TL 2.58; PL 0.84; PW 1.11; EL 1.68; EW 1.21; GD 1.21. Ratios: PL/PW 0.76; EL/EW 1.39; EL/PL 2.00; GD/EW 1.00; TL/EW 2.13. Body with dorsal and ventral surfaces dark reddish brown; mouthparts, antennae and legs reddish brown. Head coarsely and shallowly punctate on dorsum; punctures sparse, with interstice conspicuously microreticulate. Each antenna (Fig. 21B) with ten antennomeres (FL 0.17mm; CL 0.22mm; CL/FL 1.26); length of the antennomeres (in mm) as follows: 0.118; 0.055; 0.073; 0.036; 0.018; 0.023; 0.023; 0.055; 0.055; 0.109. Eyes with greatest width 1.0X the basal width of scutellum. Pronotum with deep punctures; punctures coarse, separated by a distance of one puncture width or less, but not coalescent; interstice between punctures smooth; anterolateral angles broadly rounded; lateral margins crenulated. Scutellum impunctate, smooth; basal width 1.5X the EW. Elytra with confused punctation; punctures as coarse as the ones on pronotum, separated by a distance of one puncture width or less; interstice smooth; elytral apices truncate; apical declivity (posterior one-third of elytra) bearing twelve raised keels (six in each elytron) converging to apex. Each hindwing (Fig. 20) with apical area bearing two distinct pigmented lines, one near the anterior margin and the other almost reaching the posterior margin. Prosternum biconcave, with a thin longitudinal carina at midline. Metaventrite glabrous, either side bearing large obsolescent punctures; disc smooth, dull; discrimen absent. Abdominal ventrites glabrous. Each metati-

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 11 bia (Fig. 22B) slender, near 4.5X as long as broad, slightly expanded to the basal one third, then almost parallel-sided; outer lateral edge somewhat straight, just slightly sinuous, bearing around ten spines; outer apex angulate; apical edge bearing around five distinct spines. Male genitalia (in a paratype). (Fig. 25) sclerotized and large (length more than 1.2mm); tegmen as long as median lobe, with apex expanded and globose; median lobe slender, elongate, narrower than the tegmen.

Type series Holotype. (FMNH) Philippines: /CAMP MERAN E [handwritten] slope Mt. Apo, Davo Prov. Mindanao, P.I. [printed] XI-46 [handwritten] / F.G. Werner Collector / Xylographella speciosa Lopes-Andrade HOLO- TYPUS [printed on red paper]/. Paratypes. Philippines: 8 specimens (4 FMNH, 4 LAPC), same data as holotype; 1 specimen (FMNH) labeled / E. slope Mt. McKinley, Davo Prov., Mindanao, P.I. 6600’ IX-2-1946 [handwritten] / From Polypore [handwritten] / Heyneman Collector [handwritten]/. All paratypes labeled / Xylographella speciosa Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 7, including the holotype): TL 2.05–2.95 (2.68 ± 0.32); PL 0.68–1.00 (0.89 ± 0.13); PW 0.84–1.21 (1.11 ± 0.13); EL 1.32–1.84 (1.72 ± 0.19); EW 0.89–1.32 (1.19 ± 0.14); GD 0.89–1.26 (1.17 ± 0.12). Ratios: PL/PW 0.71–0.86 (0.80 ± 0.05); EL/EW 1.36–1.52 (1.45 ± 0.06); EL/PL 1.79–2.20 (1.93 ± 0.14); GD/EW 0.92–1.00 (0.98 ± 0.03); TL/EW 2.13–2.35 (2.26 ± 0.07).

Distribution Known only from Mindanao and Luzon provinces, in the Philippines (Fig. 26).

Host fungi Unidentified bracket fungus.

Comments A species very distinct from Xyl. punctata. Besides the morphological differences, they are quite distinct in size: the biggest Xyl. punctata is considerably smaller than the smallest available specimen of Xyl. speciosa sp. nov. The description of this new species helps to reinforce the limits of the genus Xylographella, sustain- ing its current separation from Scolytocis. Some species of Tropicis (belonging to Syncosmetina subtr. nov.) also have similar longitudinal raised keels on elytra. However, these Tropicis and Xyl. speciosa sp. nov. are easily distinguished by the diagnostic characters of the subtribes of Xylographellini.

Scolytocis Blair, 1928 (Figs 1–4, 6, 10, 27–135)

Included species Scolytocis bouchardi sp. nov. Scolytocis cariborum sp. nov. Scolytocis danielssoni sp. nov. Scolytocis difficillimus sp. nov. Scolytocis fritzplaumanni sp. nov. Scolytocis furieriae sp. nov. Scolytocis howdeni sp. nov. Scolytocis indecisus sp. nov.

12 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE Scolytocis kiskeyensis sp. nov. Scolytocis lawrencei sp. nov. Scolytocis malayanus sp. nov. Scolytocis novaezelandiae sp. nov. Scolytocis panamensis sp. nov. Scolytocis paschoali sp. nov. Scolytocis philippinensis sp. nov. Scolytocis samoensis Blair, 1928 (type species, by monotypy) Scolytocis thayerae sp. nov. Scolytocis werneri sp. nov. Scolytocis zimmermani sp. nov.

Diagnosis Scolytocis may be distinguished from the other Ciidae by the combination of the diagnostic characters of Xylographellini, mainly the compact antennal club in which each antennomere bears more than four sensillifers, and the longitudinally fissured prementum with labial palpi inserted at its middle. Scolytocis differs from the other Xylographellini genera by the combination of the following characters: (i) antennal funicle with four antennomeres; (ii) lateral pronotal margins smooth; (iii) prosternal process laminate; (iv) procoxae subconical. Among the Xylographellini, character i is restricted to Scolytocis. Character ii separates Scolytocis from Xylographella. Characters i, iii and iv separate Scolytocis from the other Xylographellini.

Redescription Measurements in mm (n = 204; including all species, excepting Scol. samoensis Blair): TL 1.00–2.05; PL 0.32–0.79; PW 0.42–0.89; EL 0.58–1.26; EW 0.42–0.95; GD 0.42–0.89. Body light brown to black, shiny to dull on dorsum, subglabrous, vestiture consisting of minute, inconspicuous fine setae, sometimes with moder- ately long setae on the ventral surface. Head strongly declined, barely visible from above. Frontoclypeal region simple in both genders. Labium (Figs 3, 6) with prementum elongate, subpentagonal, apex subacute, external surface bearing a longitudinal sulcus at midline, with labial palpi inserted at its middle and apical palpomere widely expanded. Antennae inserted in front of eye (Fig. 1); fossa deep; funicle always bearing four antennomeres; club compact (with three antennomeres; Figs 2, 63A–65B, 94, 126–127) or with the antennomeres completely fused to each (so that it appears to be one large antennomeres; 65C); apical antennomere of the club bearing at least five sensillifers (Fig. 2, arrows) formed by a group of short and cylindrical sensilla. Pronotum with anterior edge simple in both sexes, broadly rounded, anterolateral angles obtuse, not produced; anterolateral margins straight or slightly arched inwards; posterolateral angles broadly rounded or angulate; lateral pronotal carinae smooth, not visible for their entire lengths from above; posterior margin sometimes bearing a rugose border along it. Scutellum conspicuous, triangular, glabrous, punctate or not; basal width more than 0.10X the EW. Elytra subquadrate to suboval; apex truncate or acute, convex or slightly concave, sometimes bearing small cuticular globules (Figs 82–84, small arrows; each globule bearing a small fine seta); posterolateral corners bending dorsally, lateral margins usually not visible from above; suture margined or not, but never deflexed at apex; punctation single, seriate to confuse. Hindwings always fully developed, sometimes extremely membranous; radial cell present and conspicuous; apical area always longer than basal area, without pigmented patches or with one to two pigmented lines/patches (complete or incomplete; distinct or vague). Metaventrite suture (discrimen) absent, or inconspicuously short to long, usually restricted to the basal region (Fig. 4). Prosternum very short; concave, biconcave or triconcave; with or without longitudinal carina at midline; prosternal process laminate and slightly curved. All tibiae expanded to middle or to apex, outer margin bearing a row of several spines (Figs 66–68, 95, 128–129). Outer face of protibiae setose along the inner margin; inner face of meso and metatibiae with a setose region near the inner margin, restricted to

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 13 apical third or extending to base. Femorae subglabrous on internal and external surfaces. Each tarsus formed by four tarsomeres. Male without abdominal setose patch or with a very small, inconspicuous setose patch in the midline of the first abdominal ventrite, barely visible even in high magnifications (around 100X) or in slide preparations. Male genitalia. Ninth segment Y-shaped (Figs 69, 73, 76, 78–79, 96–99, 130–132). Teg- men and median lobe elongate, cylindrical (69–75, 77–79, 96–100, 130–133). Tegmen with sclerotized apex and membranous basal portion; median lobe as long as tegmen or longer, membranous to barely sclerotized. Female terminalia. (Fig. 10) Eighth sternite broadly rounded or slightly arched inwards at apex. Spiculum ventrale slightly longer to twice as long as ovipositor. Gonostyli of ovipositor always absent.

Distribution Scolytocis is found on all the regions where the subtribe Xylographellina occurs, except on Japan. How- ever, it is most diversified in the neotropics, with 12 known species (all these species being described here). Comments Six morphological species groups are recognizable, and are arbitrarily named here as follows: (i) the bouchardi group, comprising the species with triconcave prosternum and larger hindwings (in comparison to the species of the lawrencei group), and including Scol. bouchardi sp. nov. and Scol. difficillimus sp. nov.; (ii) the lawrencei group, comprising the small species with tumid prosternum and small hindwings, and including Scol. cariborum sp. nov., Scol. kiskeyensis sp. nov., Scol. lawrencei sp. nov., Scol. malayanus sp. nov. and Scol. samoensis Blair; (iii) the danielssoni group, comprising the large Central-American species with a conspicuous rugose border along the posterior pronotal margin and a biconcave prosternum bearing a narrow longitudinal carina at midline, including Scol. danielssoni sp. nov., Scol. howdeni sp. nov., Scol. indecisus sp. nov. and Scol. panamensis sp. nov.; (iv) fritzplaumanni group, comprising the species with smooth posterior pronotal border and a biconcave prosternum, and including the Brazilian Scol. fritzplaumanni sp. nov., Scol. furieriae sp. nov., and Scol. paschoali sp. nov.; (v) the werneri group, comprising the species with a concave prosternum and a rugose border along the posterior pronotal margin, including Scol. novaezelandiae sp. nov. and Scol. werneri sp. nov.; (vi) the zimmermani group, comprising the species with each metatibia expanded to the middle, with its outer margin broadly rounded and bearing several spines very close to each other at the apical half, and including Scol. philippinensis sp. nov., Scol. thayerae sp. nov. Scol. zimmermani sp. nov. The species-groups are just taxonomic tools, and they cannot be considered a priori to be monophyletic taxa (Lopes-Andrade et al. 2003a).

Identification key to the Neotropical species of Scolytocis Blair

1 Posterior margin of pronotum bearing a rugose border...... 9 - Posterior margin of pronotum without a rugose border...... 2 2 Prosternum conspicuously biconcave...... 3 - Prosternum concave to slightly tumid at midline, or triconcave ...... 5 3 Pronotal punctation fine and sparse; punctures separated by one to three puncture widths...... 4 - Pronotal punctation coarse and dense; punctures separated by a distance of one puncture width of less. South Region of Brazil (Fig. 101) ...... Scol. fritzplaumanni sp. nov. 4 Antennal club less than 1.6X the funicle length. Total length usually less than 1.5mm. Southeast and Northeast Regions of Brazil (Fig. 101)...... Scol. furieriae sp. nov. - Antennal club at least 2X longer than the funicle. Total length usually more than 1.5mm. Southeast Region of Brazil (Fig. 101)...... Scol. paschoali sp. nov. 5 Prosternum concave to slightly tumid at midline, but not triconcave ...... 7 - Disc of prosternum in a different plane from the sides, so that the prosternum appears to be triconcave .. 6 6 Scutellum less than 0.18X the greatest elytral width. Known from Guatemala and Mexico (Fig. 80)......

14 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE ...... Scol. bouchardi sp. nov. - Scutellum more than 0.18X the greatest elytral width. Known from Ecuador (Fig. 80)...... Scol. difficillimus sp. nov. 7 Elytral apex almost glabrous ...... 8 - Elytral apex with concavity bearing several setae (best seen in lateral view). Known from Dominica, in the insular Northern Neotropical Region (Fig. 81)...... Scol. cariborum sp. nov. 8 Metaventrite and abdominal ventrites glabrous. Discrimen of metaventrite absent. Known from the Dominican Republic and Puerto Rico, in the insular Northern Neotropical Region (Fig. 81)...... Scol. kiskeyensis sp. nov. - Metaventrite and abdominal ventrites bearing several setae (best seen in lateral view). Discrimen of metaventrite present, being usually one-fourth the length of the metaventrite at midline. Known from Mexico, Costa Rica and Panamá (Fig. 80) ...... Scol. lawrencei sp. nov. 9 Pronotum punctation fine; punctures usually separated by a distance of two puncture widths or more... 11 - Pronotum punctation coarse; punctures very close to each other, usually separated by a distance of less than two puncture widths...... 10 10 Pronotal punctures separated by a distance of one puncture width or less. Known from Mexico (Fig. 80)...... Scol. howdeni sp. nov. - Pronotal punctures separated by a distance of one to two puncture widths. Known from Mexico and Gua- temala (Fig. 80)...... Scol. indecisus sp. nov. 11 Antennomeres of the antennal club completely fused to each other (Fig. 65C). Interstice between punctures finely microreticulate. Scutellum more than 0.13X the greatest elytral width. Known only from Pan- ama (Fig. 80)...... Scol. panamensis sp. nov. - Antennomeres of the antennal club separate, not fused (Fig. 63C). Interstice between punctures smooth, without microreticulation. Scutellum less than 0.13X the greatest elytral width. Known from Costa Rica (Fig. 80) ...... Scol. danielssoni sp. nov.

Identification key to the Indo-Malayan, Australasian and Oceanic species of Scolytocis Blair *

1 Each metatibia expanded to apex; spines of the outer edge separated by one spine width or more. Total length from 1.05 to 1.26mm ...... 4 - Each metatibia with outer margin broadly rounded, expanded to the middle; spines very close to each other at apical half. Total length from 1.26 to 2.05 mm ...... 2 2 Apical antennomere of the club longer than the preceding two antennomeres together. Total length more than 1.40 mm ...... 3 - Apical antennomere shorter than the preceding two antennomeres together. Total length less than 1.40 mm. Known from the Philippines (Fig. 134)...... Scol. philippinensis sp. nov. 3 Prosternum biconcave. Total length more than 1.7mm. Known from the Philippines (Fig. 134)...... Scol. thayerae sp. nov. - Prosternum concave. Total length less than 1.7mm. Known from Fiji (Fig. 135) ...... Scol. zimmermani sp. nov. 4 Antennal club near 1.6X longer than the funicle. Pronotum with a rugose border along the posterior margin. Total length more than 1.15mm...... 5 - Antennal club at least 1.8X longer than the funicle. Pronotum devoid of a rugose border at the posterior margin. Total length less than 1.15mm. Known from Malaysia (Southeastern Asia; Fig. 134) ...... Scol. malayanus sp. nov. 5 Rugose border (of the posterior pronotal margin) narrow. Metaventrite bearing a distinct, long discrimen. Known from the Philippines (Fig. 134)...... Scol. werneri sp. nov.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 15 - Rugose border comparatively broader. Discrimen short, not reaching the middle of the metaventrite. Known from Northern New Zealand (Fig. 135)...... Scol. novaezelandiae sp. nov.

* Scolytocis samoensis Blair not included

Northern Neotropical species of Scolytocis Blair

Scolytocis bouchardi Lopes-Andrade sp. nov. (Figs 27–29, 54, 63A, 66A, 69, 80)

Etymology The species is named after Dr. Patrice Bouchard, curator of the Coleoptera collection of CNCI. Among the specimens loaned by CNCI, I found the first specimens of two new Scolytocis species described here (Scol. bouchardi sp. nov. and Scol. howdeni sp. nov.). The study of the remaining specimens will certainly enhance the current knowledge on the Ciidae taxonomy, mainly of the Neotropical fauna.

Diagnosis The species belongs to the bouchardi species-group, due to the smooth posterior margin of pronotum and the triconcave prosternum. It differs from Scol. difficillimus sp. nov. by combination of the following characters: (i) width of eye usually more than 1.2X the basal width of scutellum; (ii) basal width of scutellum near 0.17X the greatest elytral width; (iii) Metatibiae less than 4X as long as broad.

Description Holotype. (Figs 27–29) Measurements in mm: TL 1.42; PL 0.47; PW 0.58; EL 0.89; EW 0.63; GD 0.58. Ratios: PL/PW 0.81; EL/EW 1.41; EL/PL 1.89; GD/EW 0.92; TL/EW 2.25. Body dark brown; head and pronotum darker than elytra, almost black; antennae, mouthparts and legs light yellowish brown. Head sparsely punctate, finely and inconspicuously microreticulate. Each antenna (Fig. 63A; paratype) with nine antennomeres (FL 0.098mm; CL 0.170mm; CL/FL 1.74); length of the antennomeres (in mm) as follows: 0.088; 0.043; 0.048; 0.018; 0.015; 0.018; 0.045; 0.045; 0.080. Eyes with greatest width 1.28X the basal width of scutellum. Pronotum sparsely punctate; puncture fine, shallow, separated by a distance of two to four puncture widths; interstice between punctures inconspicuously, finely microreticulate; anterolateral margins slightly arched inwards; posterolateral angles broadly rounded; posterior margin without rugose border. Scutellum bearing small punctures near the anterior margin, interstice smooth and shiny; basal width 0.17X the EW. Elytral punctation subseriate; puncture coarser, shallower and more irregular than those of pronotum; interstice between punctures shiny, rugose, but not microreticulate; elytral apex subacute, with some sparse and small cuticular globules; apical region bearing a concavity. Each hindwing (Fig. 54; paratype) with apical area bearing one vague, incomplete pigmented line near the anterior margin. Prosternum triconcave, without longitudinal carina at midline; surface shiny, rather smooth, with indistinct rugosity in high magnifications (>100X). Metaventrite bearing small and sparse setae; discrimen one-fourth the length of metaventrite at midline. Abdominal ventrites bearing small and sparse setae. Each metatibia (Fig. 66A, paratype) broadly rounded, near 3.5X as long as broad; outer edge with spines (around 20) regularly distributed at basal two thirds and then getting sparser. Male genitalia (in a paratype). (Fig. 69, paratype from Oaxaca, Mexico) Apex of tegmen (sclerotized apical two-thirds) subtriangular; median lobe extremely membranous, elongate, near 1.5X longer than tegmen.

Type series Holotype. (CNCI) Guatemala: /GUAT. Isabal [sic], 2Km N Las Escobas 300m, 15.XI.1986 M. Sharkey / Scolytocis bouchardi Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Guatemala: 4 speci-

16 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE mens (2 CNCI, 2 LAPC), same data as holotype. Mexico: 1 specimen (CMNC) /MEX: Veracruz, 33Km N. Catemaco, Los Tuxtlas Biology Stn. 8-15.VI.1984 D. H. Lindeman / Scolytocis sp. [handwritten] Det. J. F. Lawrence [printed]/; 2 specimens (1 ANIC,1 LAPC) /6mi.S Valle Nac. 2000’ Oaxaca MEX. July 30, 1971 A. Newton, coll. / Rigidoporus sp. / J. F. Lawrence Lot [printed] 3312 [handwritten]/; 12 specimens (1 ANIC, 7 FMNH, 4 LAPC) /Tezonapa Veracruz, MEX [printed] VIII-41 [handwritten] / Col. by Henry S. Dybas/; 11 specimens (7 FMNH, 4 LAPC) /El Fortin, Ver. MEX. VII-9.41 H.S. Dybas #23 / ex Polyporus [printed] lignosus [handwritten]/; 8 specimens (6 FMNH, 2 LAPC) /El Fortin, Ver. MEX. VII-9-41 H. Dybas # [printed] 23 [handwritten] / ex Polyporus lignosus/; 21 specimens (1 ANIC, 1 SMTD, 12 FMNH, 7 LAPC) /El Fortin Ver- acruz Mex. [printed] VII-9-41 [handwritten] / H.S. Dybas coll. [printed] 23 [handwritten] / ex Polyporus [printed] lignosus [handwritten]/. All the paratypes labeled /Scolytocis bouchardi Lopes-Andrade PARATY- PUS [printed on yellow paper]/

Variation Measurements in mm (n = 24, including the holotype): TL 1.37–1.58 (1.46 ± 0.05); PL 0.42–0.53 (0.48 ± 0.02); PW 0.58–0.63 (0.61 ± 0.03); EL 0.84–1.05 (0.93 ± 0.05); EW 0.63–0.74 (0.68 ± 0.03); GD 0.58–0.68 (0.63 ± 0.03). Ratios: PL/PW 0.73–0.84 (0.78 ± 0.04); EL/EW 1.29–1.54 (1.37 ± 0.05); EL/PL 1.70–2.25 (1.96 ± 0.13); GD/EW 0.85–1.08 (0.93 ± 0.05); TL/EW 2.07–2.33 (2.16 ± 0.09).

Distribution Known only from Guatemala and Mexico (Fig. 80).

Host fungi Rigidoporus sp. (Meripilaceae).

Comments The “triconcave” prosternum of Scol. bouchardi sp. nov. and Scol. difficillimus sp. nov. is unique among the Xylographellini, although that is a character not easy to observe (the prosternum itself is very short in Sco- lytocis). The “triconcave” prosternum is here defined as the prosternum with a slightly concave disc, with either lateral side concave but in different plane than the disc, appearing triconcave in frontal view. Scolytocis bouchardi sp. nov. is closely related to Scol. difficillimus sp. nov. (from Ecuador). Specimens of both Scol. bouchardi sp. nov. and Scol. difficillimus sp. nov. may be confused with the species of the lawrencei group, mainly with Scol. lawrencei sp. nov. However, the species of the bouchardi group have larger hindwings and smaller male genitalia.

Scolytocis cariborum Lopes-Andrade sp. nov. (Figs 30–32, 55, 63B, 66B, 70, 81)

Etymology The specific epithet refers to the Caribs, native habitants of the island of Dominica.

Diagnosis The species belongs to the lawrencei species-group, and so is distinguishable from the other Scolytocis by the smooth posterior margin of pronotum and the concave prosternum. It differs from the other species in the group by the elytral apex with concavity bearing several setae (best seen in lateral view) and distinct morphology of male genitalia.

Description Holotype. (Figs 30–32) Measurements in mm: TL 1.21; PL 0.42; PW 0.53; EL 0.74; EW 0.58; GD 0.47.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 17 Ratios: PL/PW 0.80; EL/EW 1.27; EL/PL 1.75; GD/EW 0.82; TL/EW 2.09. Body brown; basal antennomeres, mouthparts and legs yellowish brown. Head with shallow, coarse and sparse punctation; interstice between punctures smooth, shiny, barely microreticulate near margins. Each antenna (Fig. 63B, paratype) with nine antennomeres (FL 0.075mm; CL 0.140mm; CL/FL 1.87); length of the antennomeres (in mm) as follows: 0.068; 0.035; 0.038; 0.013; 0.013; 0.013; 0.043; 0.035; 0.063. Eyes with greatest width 1.33X the basal width of scutellum. Pronotum sparsely punctate; punctures shallow, irregular, separated by a distance of two to four puncture widths; interstice between punctures finely, inconspicuously microreticulate; anterolateral margins slightly arched inwards; posterolateral angles broadly rounded; posterior margin without rugose border. Scutellum impunctate, surface smooth; basal width 0.17X the EW. Elytra with coarse, shallow, subseriate punctation, confuse near apex; border of punctures somewhat indistinct from surrounding surface; interstice between punctures irregular, but not microreticulate; elytral apex subacute; apical region with concavity, which bears several setae (best seen in lateral view); cuticular globules very small, inconspicuous. Each hindwing (Fig. 55, paratype) with apical area bearing one vague, incomplete pigmented line near the anterior margin. Prosternum concave, rugose. Metaventrite glabrous, finely rugose; discrimen small, one-fifth the length of the metaventrite at midline. Abdominal ventrites bearing some sparse setae; surface finely rugose. Each metatibia (Fig. 66B, paratype) just slightly expanded near apex, more than 4X as long as broad; outer apical and outer lateral edges forming an obtuse rounded angle; spines (around 15) sparse. Male genitalia (in a paratype). (Fig. 70) Apex of tegmen (sclerotized three-fourths) subtriangular; median lobe around 1.5X longer than tegmen, with a conspicuous globular apex.

Type series Holotype. (ANIC) Dominica: /New Florida Estate 2000’ SW slope of Morne Anglais, DOMINICA VI- 11-73 / Cary P. Sheldon Collector / Rigidoporus sp. / J. F. Lawrence Lot [printed] 3451 [handwritten]/ Scoly- tocis cariborum Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Dominica: 4 specimens (2 ANIC, 2 LAPC), same data as holotype but distinguished labeled /Scolytocis cariborum Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 4, including the holotype): TL 1.11–1.21 (1.17 ± 0.05); PL 0.37–0.42 (0.41 ± 0.03); PW 0.47–0.53 (0.49 ± 0.03); EL 0.68–0.79 (0.72 ± 0.05); EW 0.53–0.58 (0.54 ± 0.03); GD 0.47–0.47 (0.47 ± 0.0). Ratios: PL/PW 0.78–0.89 (0.84 ± 0.06); EL/EW 1.27–1.50 (1.34 ± 0.11); EL/PL 1.63–1.88 (1.78 ± 0.12); GD/EW 0.82–0.90 (0.88 ± 0.04); TL/EW 2.09–2.30 (2.17 ± 0.10).

Distribution Known only from Dominica Island (Fig. 81) in the Lesser Antilles province (sensu Morrone 2006).

Host fungi Rigidoporus sp. (Meripilaceae).

Comments Scol. cariborum sp. nov. most resembles Scol. kiskeyensis sp. nov., and both comprise a subgroup of the lawrencei species-group due to the distinctly sclerotized median lobe that is globose (at least at apex). Both species are confined to the insular Northern Neotropical Region.

Etymology The species is named after Dr. Roy Danielsson, curator of the Entomological Collection of MZLU. The loan of a large number of ciids of MZLU has been very important to conclude a series of works on the Ciidae systematics, and two specimens of this new species were found among the specimens he sent me.

Diagnosis The species belongs to the danielssoni species-group, and so it is distinguishable from the other Scolytocis species by the biconcave prosternum with a narrow longitudinal carina at midline, the rugose border along the posterior pronotal margin and the comparatively large tibiae. It differs from Scol. howdeni sp. nov. and Scol. indecisus sp. nov. by the fine and sparse pronotal punctation, and the lack of microreticulation at the interstice between punctures. It differs from Scol. panamensis sp. nov. by the distinct separation of the three antennomeres of the club. Description Holotype. (Figs. 33–35) Measurements in mm: TL 1.68; PL 0.58; PW 0.74; EL 1.00; EW 0.79; GD 0.68. Ratios: PL/PW 0.78; EL/EW 1.27; EL/PL 1.72; GD/EW 0.86; TL/EW 2.13. Body shiny black; upper and ventral surfaces subglabrous. Head with dorsum smooth, shiny, sparsely punctate. Each antenna (Fig. 63C, paratype) with nine antennomeres (FL 0.143mm; CL 0.175mm; CL/FL 1.23); length of the antennomeres (in mm) as follows: 0.06; 0.050; 0.063; 0.028; 0.025; 0.028; 0.048; 0.050; 0.078. Eyes with greatest width 1.5X the basal width of scutellum. Pronotum punctate; punctures sparse, small, deep, uniform in size, separated by a distance of two to three puncture widths; interstice between punctures smooth, shiny; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin with a conspicuous, narrow rugose border along it. Scutellum glabrous, impunctate, shiny; basal width 0.11X the EW. Elytral punctures slightly coarser than those of pronotum; interstice between punctures smooth and shiny; elytral apex truncate, bearing a distinct concavity when seen from above; cuticular globules very small, inconspicuous. Each hindwing (Fig. 56) with apical area bearing two distinct pigmented lines, one near the anterior margin and the other almost reaching the posterior margin. Prosternum biconcave, with a narrow longitudinal carina at midline; surface rugose. Metaventrite disc delimited by a circular row of obsolescent punctures, microreticulate; discrimen inconspicuous, being just a small impression restricted to the very base of the metaventrite; either side bearing small and sparse setae, each arising from a large obsolescent puncture, with interstice microreticulate. Abdominal ventrites bearing some small and sparse setae. Each metatibia (Fig. 66C, paratype) slender, more than 4X as long as broad; outer apical and outer lateral edges forming an obtuse rounded angle; outer edge with spines (around 20) regularly distributed at apical two-thirds and then getting sparser. Male genitalia. Unknown.

Type series Holotype. (MZLU) Costa Rica: /Costa Rica, San Jose, Cerro de la Muerte 3200m, 20.III.1986 leg. M. Sörensson / Scolytocis danielssoni Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Costa Rica: 1 specimen (LAPC), same data as the holotype; 3 specimens (2 ANIC, 1 LAPC) labeled /2.5mi.S.W. Rincon Puntarenas, Costa Rica Mar. 1-7, ’67 / Robin Andrews Coll. / Phellinus sp. / J. F. Lawrence Lot [printed] 2185 [handwritten]/. All the paratypes with an additional label /Scolytocis danielssoni Lopes- Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 4, including the holotype): TL 1.53–1.79 (1.67 ± 0.11); PL 0.53–0.63 (0.58 ± 0.04); PW 0.63–0.74 (0.71 ± 0.05); EL 0.89–1.11 (1.01 ± 0.09); EW 0.74–0.84 (0.80 ± 0.05); GD 0.63–0.79 (0.71 ± 0.07). Ratios: PL/PW 0.78–0.86 (0.82 ± 0.04); EL/EW 1.20–1.31 (1.26 ± 0.05); EL/PL 1.67–1.91 (1.74 ± 0.11); GD/EW 0.85–0.94 (0.88 ± 0.04); TL/EW 2.00–2.13 (2.08 ± 0.06).

Host fungi Phellinus sp. (Hymenochaetaceae).

Comments Among the Scolytocis examined, this new species occurs at the highest known altitude (type locality, Cerro de la Muerte, 3200m over the sea).

Scolytocis difficillimus Lopes-Andrade sp. nov. (Figs 36–38, 57, 64A, 67A, 71, 80)

Etymology The specific epithet is a reference to the difficulty in recognizing the species, mainly in distinguishing it from Scol. bouchardi sp. nov.

Diagnosis The species belongs to the bouchardi species-group, so it differs from the other Scolytocis mainly by the triconcave prosternum. It differs from Scol. bouchardi sp. nov., the most closely related species, by the comparatively smaller body (total length less than 1.3 mm) and small eyes (1.0X the basal width of scutellum).

Description Holotype. (Figs. 36–38) Measurements in mm: TL 1.16; PL 0.42; PW 0.53; EL 0.74; EW 0.53; GD 0.53. Ratios: PL/PW 0.80; EL/EW 1.40; EL/PL 1.75; GD/EW 1.00; TL/EW 2.20. Body dark brown; antennae, mouthparts and legs yellowish brown. Head with shallow, sparse, inconspicuous punctation; interstice between punctures finely microreticulate. Each antenna (Fig. 64A, paratype) with nine antennomeres (FL 0.078mm; CL 0.150mm; CL/FL 1.94); length of the antennomeres (in mm) as follows: 0.088; 0.040; 0.040; 0.013; 0.013; 0.013; 0.040; 0.038; 0.073. Eyes with greatest width 1.0X the basal width of scutellum. Pronotal punctation coarse, sparse, shallow; punctures separated by a distance of two to four punctures widths; interstice microreticulate; anterolateral margins slightly arched inwards; posterolateral angles broadly rounded; posterior margin without rugose border. Scutellum with some fine, sparse punctures; basal width 0.21X the EW. Elytral punctation confuse at base and subseriate at disc; punctures coarse, shallow, irregular; punctures without conspicuous border; interstice between punctures finely rugose; elytral apex subacute, declivity with weak concavity; cuticular globules very small, inconspicuous; elytral suture weakly margined at apical third. Each hindwing (Fig. 57) with apical area bearing one vague, incomplete pigmented line near the anterior margin. Prosternum triconcave, without longitudinal carina at midline; surface shiny, rugose. Metaventrite bearing small and sparse setae, with interstice finely microreticulate; discrimen one-fifth the length of the metaventrite at midline. Abdominal ventrites bearing small and sparse setae; interstice finely microreticulate. Each metatibiae (Fig. 67A) broadly rounded, near 3.5X as long as broad; outer edge with spines (around 20) regularly distributed at apical two thirds and then getting sparser. Male genitalia (in a paratype). (Fig. 71) Apex of tegmen (sclerotized apical two-thirds) subtriangular; median lobe not observed, probably extremely membranous.

Type series Holotype. (ANIC) Ecuador: /ECUAD: Pichincha Rio Palenque, 47Km.S Sto Domingo JULY 20-30, 1975 / S. & J. Peck Collectors / Rigidoporus sp. / Scolytocis difficillimus Lopes-Andrade HOLOTYPUS

20 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE [printed on red paper]/. Paratypes. Ecuador: 4 specimens (2 ANIC, 2 LAPC), same data as holotype. All the paratypes with an additional label /Scolytocis difficillimus Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 2, including the holotype): TL 1.16–1.26 (1.21 ± 0.07); PL 0.42–0.42 (0.42 ± 0.00); PW 0.53–0.53 (0.53 ± 0.00); EL 0.74–0.84 (0.79 ± 0.07); EW 0.53–0.58 (0.55 ± 0.04); GD 0.53–0.53 (0.53 ± 0.00). Ratios: PL/PW 0.80–0.80 (0.80 ± 0.00); EL/EW 1.40–1.45 (1.43 ± 0.04); EL/PL 1.75–2.00 (1.88 ± 0.18); GD/EW 0.91–1.00 (0.95 ± 0.06); TL/EW 2.18–2.20 (2.19 ± 0.01).

Distribution Known only from the type locality in Ecuador (Fig. 80).

Host fungi Rigidoporus sp. (Meripilaceae).

Comments I was a little bit reluctant in describing this species, due to its similarities to Scol. bouchardi sp. nov. How- ever, after a careful comparison of morphological features, I concluded that some specimens from Ecuador, from a single field collection, are quite distinct and cannot be adequately included in any of the new northern Neotropical species of Scolytocis. After delimiting Scol. difficillimus sp. nov., I found out that it was very similar to Scol. bouchardi sp. nov. in the morphology of hindwings, tibiae, and prosternum (“triconcave”), which led me to define a morphological group for both species (the bouchardi species-group).

Scolytocis howdeni Lopes-Andrade sp. nov. (Figs 39–41, 58, 64B, 67B, 72, 80)

Etymology The species is named after Henry F. Howden, who collected six specimens of the type series.

Description Holotype. (Figs 39–41) Measurements in mm: TL 2.00; PL 0.79; PW 0.89; EL 1.21; EW 0.95; GD 0.89. Ratios: PL/PW 0.88; EL/EW 1.28; EL/PL 1.53; GD/EW 0.94; TL/EW 2.11. Body dark brown, head and pronotum slightly darker than elytra; basal antennomeres, mouthparts and tarsi yellowish brown; dorsal and ventral surfaces almost entirely glabrous. Head with dorsum coarsely punctate; punctures irregular, with interstice finely microreticulate. Each antenna (Fig. 64B, paratype) with nine antennomeres (FL 0.098mm; CL 0.160mm; CL/FL 1.64); length of the antennomeres (in mm) as follows: 0.118; 0.055; 0.053; 0.015; 0.015; 0.015; 0.050; 0.045; 0.065. Eyes with greatest width 1.33X the basal width of scutellum. Pronotum coarsely punctate; punctures irregular, limits distinct, usually separate by a distance of one puncture width or less, but not coalescent; interstice between punctures finely microreticulate; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin with a conspicuous rugose border along it. Scutellum rugose,

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 21 basal width 0.13X the EW. Elytra with punctation slightly coarser, denser and more irregular than that of pronotum; interstice between punctures smooth; elytral apex truncate, with a small concavity when seen from above; cuticular globules absent. Each hindwing (Fig. 58) with apical area bearing one vague, incomplete pigmented line near the anterior margin. Prosternum biconcave, with a narrow longitudinal carina at midline; surface microreticulate to striated. Metaventrite glabrous, surface microreticulate to striated; discrimen indistinct; either side bearing large obsolescent punctures. Abdominal ventrites glabrous; surface microreticulate. Each metatibia (Fig. 67B) near 3.0X as long as broad; outer apical and outer lateral edges forming an obtuse angle that is broadly rounded; outer edge with spines (around 20) regularly distributed and more close to each other at apical third (outer apical edge), then getting sparser. Male genitalia (in a paratype). (Fig. 72) Apex of tegmen (sclerotized apical half) subtriangular, base membranous; median lobe membranous, elongate, near 1.2X longer than tegmen.

Type series Holotype. (CNCI) Mexico: /3 mi. N.W. San Cristobal L.C., Chis. Mex. V. 29 1969 H.F. Howden / Scoly- tocis howdeni Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Mexico: 4 specimens (2 CNCI, 2 LAPC), same data as holotype; 1 specimen (ANIC) /6.4mi. S. Tenango de Doria Hidalgo MEX.July 24, 1969 / S. & J. Peck Collectors / Ganoderma sp. / J. F. Lawrence Lot [printed] 3079 [handwritten]/. All the paratypes with an additional label /Scolytocis howdeni Lopes-Andrade PARATYPUS [printed on yellow paper]/

Variation Measurements in mm (n = 6, including the holotype): TL 1.84–2.00 (1.92 ± 0.07); PL 0.68–0.79 (0.70 ± 0.04); PW 0.79–0.89 (0.85 ± 0.04); EL 1.16–1.26 (1.20 ± 0.04); EW 0.84–0.95 (0.92 ± 0.04); GD 0.84–0.89 (0.86 ± 0.03). Ratios: PL/PW 0.76–0.88 (0.83 ± 0.04); EL/EW 1.22–1.38 (1.31 ± 0.06); EL/PL 1.53–1.85 (1.72 ± 0.11); GD/EW 0.89–1.00 (0.93 ± 0.04); TL/EW 1.94–2.19 (2.09 ± 0.08).

Distribution Known from Chiapas and Hidalgo, Mexico (Fig. 80).

Host fungi Ganoderma sp. (Ganodermataceae).

Comments It resembles Scol. indecisus sp. nov. in the coarse and dense punctation of elytra and pronotum, but is easily distinguished by the features mentioned in the diagnosis.

Scolytocis indecisus Lopes-Andrade sp. nov. (Figs 42–44, 59, 64C, 67C, 73, 80)

Etymology The specific epithet refers to my indecision in choosing the name of this new species.

22 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE Description Holotype. (Figs 42–44) Measurements in mm: TL 1.42; PL 0.47; PW 0.58; EL 0.89; EW 0.68; GD 0.63. Ratios: PL/PW 0.82; EL/EW 1.31; EL/PL 1.89; GD/EW 0.92; TL/EW 2.08. Body shiny dark-brown, with head and pronotum darker than elytra; antennae, mouthparts and legs yellowish brown. Head coarsely punctate, with interstice between punctures microreticulate. Each antenna (Fig. 64C, paratype) with nine antennomeres (FL 0.095mm; CL 0.145mm; CL/FL 1.53); length of the antennomeres (in mm) as follows: 0.068; 0.038; 0.050; 0.015; 0.015; 0.015; 0.038; 0.040; 0.068. Eyes with greatest width 1.0X the basal width of scutellum; ommatidia with irregular surface. Pronotum with deep and coarse punctation; punctures separated by a distance of one to two puncture widths, some areas of the disc with sparser punctation; interstice between punctures inconspicuously microreticulate, shiny; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin with a rugose border along it. Scutellum glabrous, impunctate, shiny, basal width 0.13X the EW. Elytra with confuse punctation, slightly coarser than that of pronotum; punctures deep, with interstice smooth and shiny; elytral apex truncate, slightly concave when seen from above, bearing small, inconspicuous cuticular globules. Each hindwing (Fig.59) extremely membranous, probably not or barely functional for flight; apical area without conspicuous pigmented lines. Prosternum biconcave, with a narrow longitudinal carina at midline; surface rugose. Metaventrite with rugose surface; disc delimited by a circular row of punctures; discrimen absent; either side bearing small and sparse setae, each arising from large obsolescent punctures. Abdominal ventrites with rugose surface and bearing small and sparse setae. Each metatibia (Fig. 67C, paratype) near 3.2X as long as broad; outer apical and outer lateral edges forming an obtuse angle that is broadly rounded; outer edge with spines (around 20) regularly distributed and more close to each other at apical two-thirds, then getting sparser. Male genitalia (in a paratype). (Fig. 73) Tegmen with apex (sclerotized apical half) subpentagonal, basal half extremely membranous; median lobe as long as tegmen.

Type series Holotype. (ANIC) Mexico: /Tenango de Doria Hidalgo, MEXICO July 24, 1969 / S. & J. Peck Collectors / Phellinus sp. / J. F. Lawrence Lot [printed] 3075 [handwritten]/ Scolytocis indecisus Lopes-Andrade HOLO- TYPUS [printed on red paper]/. Paratypes. Mexico: 8 specimens (5 ANIC, 3 LAPC), same data as holotype; 2 specimens (ANIC) /Tenango de Doria Hidalgo, MEXICO July 24, 1969 / S. & J. Peck Collectors / Gano- derma sp. / J. F. Lawrence Lot [printed] 3074 [handwritten]/; 3 specimens (2 ANIC, 1 LAPC) /7mi. SW Ten- ango de Doria Hidalgo, MEX. VII-2-71 A. Newton / Phellinus sp. / J. F. Lawrence Lot [printed] 3258 [handwritten]/. Guatemala: 1 specimen /GUAT.: QUEZALTEN.[ANGO] 12Km.S.E. Zunil, N.W. face Cerro Zunil, Fuentes Georginas, 2700m 17.VI.1993, R. Anderson hardwood for.litter, 93-2C [sic]/; 9 specimens (6 FMNH, 3 LAPC) / Sierra de las Minas / Montana Piamont – 9000ft. / Dept. Progresso Guat. [printed] II:7:42 [handwritten] / Col. By A.E. Vatter, Jr. / F. Mus. Bot. Exp. to Guat. / Fungus # 436-6 [handwritten]/; 75 specimens (48 FMNH, 25 LAPC, 2 SMTD)/ Sierra de las Minas; Montana Piamont – 9000ft.; Dept. Progresso Guat. II:7:42; Col. By A.E. Vatter, Jr. / F. Mus. Bot. Exp. to Guat.; Fungus # 436-6/. All the paratypes with an additional label /Scolytocis indecisus Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 20, including the holotype): TL 1.26–1.68 (1.51 ± 0.15); PL 0.42–0.63 (0.52 ± 0.05); PW 0.53–0.74 (0.63 ± 0.06); EL 0.74–1.16 (0.95 ± 0.12); EW 0.63–0.84 (0.72 ± 0.07); GD 0.58–0.74 (0.66 ± 0.06). Ratios: PL/PW 0.73–0.92 (0.83 ± 0.05); EL/EW 1.13–1.50 (1.32 ± 0.10); EL/PL 1.40–2.20 (1.83 ± 0.20); GD/EW 0.85–1.00 (0.92 ± 0.04); TL/EW 1.92–2.29 (2.10 ± 0.10). Distribution Known from Mexico and Guatemala (Fig. 80).

Host fungi Ganoderma sp. (Ganodermataceae) and Phellinus sp. (Hymenochaetaceae).

Etymology The specific epithet refers to Kiskeya (also Quisqueya), an indigenous name for the island of Hispaniola (Haiti and Dominican Republic), terra typica of this species.

Diagnosis The species belongs to the lawrencei species-group, and so is distinguishable from the other Scolytocis by the smooth posterior margin of pronotum and the concave prosternum. Scol. cariborum sp. nov. has several setae at the elytral apex, and Scol. lawrencei sp. nov. has setae at the metaventrite and abdominal ventrites.

Description Holotype. (Figs 45–47) Measurements in mm: TL 1.00; PL 0.37; PW 0.42; EL 0.63; EW 0.47; GD 0.42. Ratios: PL/PW 0.88; EL/EW 1.33; EL/PL 1.71; GD/EW 0.89; TL/EW 2.11. Body reddish brown; antennae, mouthparts and legs slightly lighter than body. Head shiny, sparsely punctate; punctures fine, interstice smooth. Each antenna (Fig. 65A) with nine antennomeres (FL 0.073mm; CL 0.108mm; CL/FL 1.48); length of the antennomeres (in mm) as follows: 0.060; 0.035; 0.035; 0.013; 0.013; 0.013; 0.025; 0.033; 0.050. Eyes with greatest width 1.0X the basal width of scutellum. Pronotum sparsely, finely punctate; puncture deep, separated by a distance of three to five puncture widths; interstice between punctures microreticulate; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin without rugose border. Scutellum shiny, rugose but not microreticulate, bearing one distinct puncture; basal width 0.16X the EW. Elytra with coarse, shallow, subseriate punctation; limits of punctures not well distinct (when seen from above), interstice between punctures smooth; elytral apex rounded, slightly concave, bearing very small, indistinct cuticular globules (best seen in lateral view). Each hindwings (Fig. 60) with apical area without conspicuous pigmented lines. Prosternum concave, without longitudinal carina at midline; surface rugose. Metaventrite glabrous, rugose; discrimen indistinct. Abdominal ventrites glabrous, rugose. Each metatibiae (Fig. 68A) near 3X as long as broad; outer apical and outer lateral edges forming an almost perpendicular angle that is rounded; outer edge with spines (around 15) regularly distributed at apical two-thirds. Male genitalia (in paratypes). (Fig. 74–76) Both tegmen and median lobe completely and equally sclerotized, elongate but somewhat globose; median lobe broadly rounded, only 4X as long as broad and larger at middle.

Type series Holotype. (ANIC) Dominican Republic: /DOM. REP.: Polo, 690m July, 1974 P. Hertz / Fomes sclerodermeus / J. F. Lawrence Lot No. [printed] 3659 [handwritten]/ Scolytocis kiskeyensis Lopes-Andrade HOLO- TYPUS [printed on red paper]/. Paratypes. Dominican Republic: 2 specimens (1 ANIC, 1 LAPC), same data as holotype. Puerto Rico: 2 specimens (LAPC) labeled /Cerro Dona Juana Ponce, P.R. Dec. 28, 1966 / S. Peck Collector / Ganoderma sp. / J. F. Lawrence Lot. [printed] 1929 [handwritten]/; 1 specimen (ANIC) / Luquillo Exp. For. Humacao, P.R. Dec. 23, 1966 / S. Peck Collector / ex Fomes sclerodermeus / J. F. Lawrence Lot. [printed] 1926 [handwritten]/. All the paratypes with an additional label /Scolytocis kiskeyensis Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 3, including the holotype): TL 1.00–1.16 (1.07 ± 0.08); PL 0.32–0.42 (0.37 ± 0.05); PW 0.42–0.47 (0.44 ± 0.03); EL 0.63–0.74 (0.68 ± 0.05); EW 0.47–0.53 (0.49 ± 0.03); GD 0.42–0.47 (0.44 ± 0.03). Ratios: PL/PW 0.75–0.89 (0.84 ± 0.08); EL/EW 1.33–1.44 (1.39 ± 0.06); EL/PL 1.71–2.17 (1.88 ± 0.25); GD/EW 0.89–0.90 (0.89 ± 0.01); TL/EW 2.11–2.22 (2.18 ± 0.06).

Host fungi Fomes fasciatus (Sw.) Cooke (Polyporaceae) and Ganoderma sp. (Ganodermataceae).

Comments The morphology of the male genitalia in this species is unique among all the known Scolytocis. The genitalia of Scol. cariborum sp. nov. has a median lobe with a globose apex, somewhat resembling the one of Scol. kiskeyensis sp. nov.

Scolytocis lawrencei Lopes-Andrade sp. nov. (Figs 48–50, 61, 65B, 68B, 77–78, 80)

Etymology The species is named after John F. Lawrence, great coleopterist (and ciidologist) who arranged and organized most of the Xylographellini described in this work.

Diagnosis The species belongs to the lawrencei species-group, and so is distinguishable from the other Scolytocis by the smooth posterior margin of pronotum and the concave prosternum. It differs from the other Neotropical species in the group mainly by the glabrous elytral apex and the slightly setose metaventrite and abdominal ventrites.

Description Holotype. (Figs 48–50) Measurements in mm: TL 1.16; PL 0.42; PW 0.53; EL 0.74; EW 0.58; GD 0.53. Ratios: PL/PW 0.80; EL/EW 1.27; EL/PL 1.75; GD/EW 0.91; TL/EW 2.00. Body brown; basal antennomeres, mouthparts and legs light yellowish brown. Head finely and sparsely punctate; interstice between puncture inconspicuously microreticulate. Each antenna (Fig. 65B, paratype) with nine antennomeres (FL 0.073mm; CL 0.130mm; CL/FL 1.79); length of the antennomeres (in mm) as follows: 0.073; 0.033; 0.035; 0.013; 0.013; 0.013; 0.038; 0.030; 0.063. Eyes with greatest width 1.0X the basal width of scutellum. Prono- tum with shallow, coarse, sparse punctation; punctures separate by a distance of two to three puncture widths, interstice microreticulate; anterolateral margins slighted arched inwards; posterolateral angles broadly rounded; posterior margin without rugose border. Scutellum impunctate, microreticulate; basal width 0.19X the EW. Elytra with subseriate punctation; punctures coarser and shallower than those of pronotum, border indistinct from elytral surface (at least when seen from above); interstice between punctures rugose, but without microreticulation; elytral apex subacute, declivity without conspicuous concavity but bearing some small, inconspicuous cuticular globules. Each hindwing (Fig. 61) with apical area bearing one vague, incomplete pigmented line near the anterior margin. Prosternum concave, without longitudinal carina at midline; surface shiny, slightly rugose. Metaventrite slightly microreticulate; disc delimited by a circular row of obsolescent punctures; discrimen one-fourth the length of the metaventrite at midline; either side bearing small and sparse setae, each arising from a large obsolescent punctures. Abdominal ventrites bearing small and sparse setae, best seen in lateral view. Each metatibia (Fig. 68B) just slightly expanded near apex, around 4X as long as broad; outer edge broadly rounded, bearing spines (around 20) regularly distributed at apical two thirds and then getting sparser. Male genitalia (in paratypes). (Figs 77–78) Tegmen and median lobe barely sclerotized; tegmen with apex (apical three-fourths) subtriangular; median lobe extremely elongate, more than 10X longer than broad.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 25 Type series Holotype. (ANIC) Costa Rica: /C. RICA: Heredia Finca la Selva 1974 [printed] July [handwritten]/ J. F. Lawrence Lot [printed] 3643 [handwritten]/ S. Klein-Feldt [handwritten]/ Rigidoporus sp./ 636 [handwritten]/ Scolytocis lawrencei Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Costa Rica: 7 specimens (4 ANIC, 3 LAPC), same data as holotype; 1 specimen (ANIC) /Puerto Viejo Costa Rica VIII-4-65 / A. Raske Collector / ex Polyporus zonalis / J. F. Lawrence Lot [printed] 1613 [handwritten]/; 1 specimen (ANIC) /Turrialba Costa Rica VIII-31-66/ Robin Andrews Collector / Fomes (Rigidoporus) auberianus ? / J. F. Lawrence Lot [printed] 1848 [handwritten]/; 1 specimen (CMNC) "COSTA RICA: Punt.[arenas] S. Vito, Las Cruces July 1982 B. Gill 1200m/. Panama: (Total of 51 paratypes: 19 ANIC, 16 FMNH, 15 LAPC, 1 SMTD) 1 specimen /PANAMA: Almirante Bocas del Toro Prov. March 27, 1959 [handwritten] / H. Dybas # [printed] 59-138 [handwritten] / Rigidoporus lignosus/; 5 specimens /Barro Colorado Is. CANAL ZONE [printed] 1.26.59 [handwritten] / H. Dybas [printed] #59-82 [handwritten] / ex Polyporus lignosus/; 17 specimens / Barro Colorado Is. CANAL ZONE [printed] 1.26.59 [handwritten] / H. Dybas [printed] #59-83 [handwritten] / ex Polyporus lignosus/; 11 specimens labeled /Canal Zone: Is. Barro Colorado II.19-III.9-75 Lawrence, Erwin / Rigidoporus sp. /, 5 distinguished labeled /J. F. Lawrence Lot [printed] 3777 [handwritten]/ and 6 /J. F. Lawrence Lot [printed] 3811 [handwritten]/; 4 specimens /Barro Colorado Is. Canal Zone Feb. 15,1968 / J. F. Lawrence Lot [printed] 2419 [handwritten]/, 3 specimens distinguished labeled /Polyporus lignosus/ and 1 / Trametes corrugata/; 3 specimens /Barro Colorado Is. Canal Zone Panama, IV-3-67 / ex Polyporus lignosus / J. F. Lawrence Lot [printed] 2100 [handwritten]/; 2 specimens /Panama: Canal Zone Barro Colorado Is. Feb- ruary 6 1976 A. Newton / litter under rotting logs /; 4 specimens /Barro Colorado Is. Canal Zone July 11 1969 / J. F. Lawrence Lot [printed] 2847 [handwritten]/; 2 specimens /Barro Colorado Is. Canal Zone July 11 1969 / Rigidoporus sp. / J. F. Lawrence Lot [printed] 2838 [handwritten]/; 2 specimens /Barro Colorado Is. Canal Zone Aug. 13 1969 / Rigidoporus sp. / J. F. Lawrence Lot [printed] 3042 [handwritten]/. Mexico: 1 specimen (CNCI) /MEX. Chis., Palenque 2-30.VII.1983, rain forest, S&J Peck & R. Anderson, 100m/; 16 specimens (10 FMNH, 6 LAPC) /Palenque, MEX. Chiapas VII-49 C.J. Goodnight/. All the paratypes with an additional label /Scolytocis lawrencei Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 28, including the holotype): TL 1.00–1.37 (1.19 ± 0.09); PL 0.37–0.47 (0.41 ± 0.03); PW 0.42–0.58 (0.50 ± 0.04); EL 0.58–0.89 (0.75 ± 0.08); EW 0.47–0.63 (0.55 ± 0.05); GD 0.42–0.63 (0.51 ± 0.05). Ratios: PL/PW 0.73–0.90 (0.82 ± 0.05); EL/EW 1.10–1.67 (1.35 ± 0.10); EL/PL 1.38–2.14 (1.83 ± 0.18); GD/EW 0.82–1.00 (0.92 ± 0.05); TL/EW 2.00–2.44 (2.15 ± 0.09).

Distribution Known from Mexico, Costa Rica and Panama (Fig. 80).

Host fungi Rigidoporus sp., Rigidoporus lineatus (Pers.) Ryvarden and Rigidoporus microsporus (Sw.) Overeem (Meripilaceae); Earliella scabrosa (Pers.) Gilb. & Ryvarden (Polyporaceae). Comments A common and widespread continental Mesoamerican Scolytocis. It co-occurs with Scol. panamensis sp. nov. in Panama, and large specimens of Scol. lawrencei sp. nov. may be confused with it in dorsal view. How- ever, Scol. panamensis sp. nov. belongs to the danielssoni species-group, and is unique among the Scolytocis in the fused antennomeres of the club.

Etymology The specific epithet refers to Panama, terra typica of this new species.

Diagnosis The species belong to the danielssoni species-group, and so it is distinguishable from the other Scolytocis species by the biconcave prosternum with a narrow longitudinal carina at midline, the rugose border along the posterior pronotal margin and the comparatively large metatibia (Fig. 68C). The species is easily distinguished from all the other Central American Scolytocis by the fused antennomeres of the club, which appear as one large antennomere even in slide preparations (Fig. 65C). Description Holotype. (Figs51–53) Measurements in mm: TL 1.53; PL 0.53; PW 0.68; EL 0.95; EW 0.74; GD 0.68. Ratios: PL/PW 0.77; EL/EW 1.29; EL/PL 1.80; GD/EW 0.93; TL/EW 2.07. Body black; basal antennomeres, mouthparts and legs light yellowish brown. Head sparsely and finely punctate; interstice between puncture smooth, shiny. Each antenna (Fig. 65C) with seven antennomeres (FL 0.098mm; CL 0.145mm; CL/FL 1.49); funicle with four antennomeres and club with only one distinguishable large antennomere; length of the antennomeres (in mm) as follows: 0.085; 0.045; 0.050; 0.018; 0.015; 0.015; 0.145 (= fused antennomeres of the club). Eyes with greatest width 1.28X the basal width of scutellum. Pronotum with fine, shallow, sparse punctation; punctures separate by a distance of three to five punctures, interstice microreticulate; anterolateral margins of pronotum slightly arched inwards; posterolateral angles broadly rounded; posterior margin bearing a rugose border along it. Scutellum punctate, rugose; basal width 0.15X the EW. Elytra with confuse punctation; punctures coarser and shallower than that of pronotum; punctures without conspicuous limits (at least when seen from above), separated by a distance of one to two puncture widths, interstice smooth, shiny; elytral apex truncate; apical declivity concave, bearing small cuticular globules. Each hindwing (Fig. 62) with apical area bearing one vague, incomplete pigmented line near the anterior margin. Prosternum biconcave, with a narrow longitudinal carina at midline; surface striated. Metaventrite glabrous, shiny, rugose; discrimen one-fourth the length of the metaventrite at midline; either side with some large obsolescent punctures. Abdominal ventrites glabrous, rugose. Each metatibia (Fig. 68C) broadly rounded, near 3X as long as broad; outer apical and outer lateral edges forming an obtuse rounded angle; spines (less than 20) close to each other at the outer apical edge, then getting sparser. Male genitalia (in a paratype). (Fig. 79) Genitalia very small (around 0.25mm, including the ninth segment), barely sclerotized. Apical portion of tegmen (apical four- fifths) triangular, apex conspicuously acute; median lobe extremely membranous, indistinct, as long as tegmen.

Type series Holotype. Panama: /Barro Colorado Is. Canal Zone July 14 1969 Phellinus sp. / J. F. Lawrence Lot [printed] 2889 [handwritten]/ Scolytocis panamensis Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Panama: 2 specimens, same data as holotype (1 ANIC, 1 LAPC); 4 specimens (2 ANIC, 2 LAPC) /Barro Colorado Is. Canal Zone Feb. 5, 1968 / Fomes pseudosenex / J. F. Lawrence Lot [printed] 2310 [handwritten]/; 1 specimen (LAPC) /Barro Colorado Is. Canal Zone July 6 1969 / Phellinus sp. / J. F. Lawrence Lot [printed] 2787 [handwritten]/. All paratypes with an additional label /Scolytocis panamensis Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 5, including the holotype): TL 1.53–1.63 (1.56 ± 0.05); PL 0.53–0.58 (0.56 ± 0.03); PW 0.63–0.68 (0.67 ± 0.02); EL 0.89–1.00 (0.95 ± 0.04); EW 0.68–0.79 (0.74 ± 0.04); GD 0.63–0.68

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 27 (0.66 ± 0.03). Ratios: PL/PW 0.77–0.92 (0.83 ± 0.06); EL/EW 1.20–1.36 (1.29 ± 0.06); EL/PL 1.55–1.90 (1.70 ± 0.14); GD/EW 0.86–0.93 (0.90 ± 0.04); TL/EW 2.07–2.23 (2.12 ± 0.07).

Distribution Known only from Panama (Fig. 80).

Host fungi Phellinus sp. (Hymenochaetaceae) and Fomes pseudosenex (Murrill) Sacc. & Trotter (Polyporaceae).

Comments This species is unique among all the other Ciidae due to the fused antennomeres of the antennal club (so each antenna has seven antennomeres). Besides the fused club, the gross antennal morphology of Scol. panamensis sp. nov. is similar to that of other species in the danielssoni group.

Southern Neotropical species of Scolytocis

Scolytocis fritzplaumanni Lopes-Andrade sp. nov. (Figs 82–84, 91, 94A, 95A, 96–97, 101)

Etymology The specific epithet is in honour of the great entomologist Fritz Plaumann, who first collected the species. A German immigrant, Fritz Plaumann lived in Brazil from 1924 until his death in 1994. His collection was considered the biggest private insect collection of Latin America, and the ciids he collected are an unique source of knowledge on the Ciidae fauna of southern Brazil.

Diagnosis The species belongs to the fritzplaumanni species-group and so it is distinguishable from the other Scoly- tocis by the biconcave prosternum without longitudinal carina at midline, and the smooth border of the posterior pronotal margin. It differs from the other species in the group by the coarse and dense pronotal punctation, with punctures separated by a distance of one puncture width of less.

Description Holotype. (Figs 82–84) Measurements in mm: TL 1.74; PL 0.68; PW 0.79; EL 0.95; EW 0.84; GD 0.79. Ratios: PL/PW 0.86; EL/EW 1.13; EL/PL 1.40; GD/EW 0.94; TL/EW 2.07. Body dark brown, including antennae, mouthparts and legs; tarsi a little bit lighter than legs. Head with dorsum punctate; puncture coarse, with interstice finely microreticulate. Each antenna (Fig. 94A) with nine antennomeres (FL 0.095mm; CL 0.148mm; CL/FL 1.55); length of the antennomeres (in mm) as follows: 0.095; 0.038; 0.053; 0.015; 0.015; 0.013; 0.038; 0.043; 0.068. Eyes with greatest width 1.22X the basal width of scutellum. Pronotum coarsely and densely punctate; punctures irregular, shallow, with limits well-distinct, separate by a distance of one puncture width or less; interstice between punctures finely microreticulate; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin without a rugose border. Scutellum smooth, shiny, with sparse punctures, basal width 0.15X the EW. Elytra with coarse and confused punctation; punctures irregular, deeper than those of pronotum, sometimes without distinct limits; interstice between punctures smooth; elytral apex truncate, slightly concave when seen from above, bearing sparse but conspicuous cuticular globules (Figs 82–84, small arrows). Each hindwing (Fig. 91) with apical area bearing two pigmented lines, one conspicuous line near the anterior margin and the other a vague line almost reaching the posterior margin. Pros- ternum biconcave, bearing a very narrow longitudinal carina at midline; surface shiny, slightly rugose. Metaventrite finely microreticulate at disc to striated near sides; disc delimited by a circular row of obsoles-

28 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE cent punctures; discrimen one-third the length of the metaventrite at midline; either side bearing some small and sparse setae, each arising from large obsolescent punctures. Abdominal ventrites bearing small and sparse setae. Each metatibia (Fig. 94A) broadly rounded, less than 3X as long as broad; outer edge with spines (around 25) closer to each other at apical one-fourth, then getting sparser. Male genitalia (in paratypes). (Figs 96–97) Genitalia elongate, equally sclerotized; apex of tegmen subtriangular; median lobe around 1.25X longer than tegmen. Type series Holotype. (MZSP) Brazil: /Nova Teutonia Santa Catarina Brazil Fritz Plaumann [printed] XI-64 [handwritten]/ Scolytocis fritzplaumanni Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Brazil: 24 specimens (16 ANIC, 8 LAPC) labeled /Nova Teutonia Santa Catarina BRAZIL March F. Plaumann/; 3 specimens (2 ANIC, 1 LAPC) /Brasilien Nova Teutonia 27º11' B 52º23' L Fritz Plaumann 300-500m v. 1960/ ; 3 specimens (2 ANIC, 1 LAPC) /Serra do Mar, 1200m 25' 46, 49' 06 BRAZIL May, 1964 Plaumann/; 1 specimen (ANIC) /Serra do Mar Brazil 1000m May, 1964 [handwritten]/ Fritz Plaumann Collector [printed]/; 4 specimens (3 ANIC, 1 LAPC) /Nova Teutonia Santa Catarina BRAZIL Fritz Plaumann [printed] June 1972 [handwritten]/; 2 specimens (LAPC) /Boqueirão, 29'16, 52' 32 Rio Grande do Sul Brazil ix.66 / Fritz Plau- mann Collector/; 2 specimens (ANIC) /Boqueirão 29'16 52' 32 600m / F. Plaumann Sept. 1960/; 121 specimens (2 CAMB, 10 CNCI, 2 GOPC, 2 ISNB, 10 LUND, 2 MCZ, 6 MHNG, 4 RRPC, 10 SMTD) /BRASIL: PR Guaratuba; “Pontal do Rio Itararé”; 900m iii.2005 leg. P. Coelho Grossi / Ganoderma sp./; 85 specimens (49 LAPC, 2 LUND, 30 MCNZ, 4 MHNG) /BRASIL: RS São Francisco de Paula "FLONA SFP"; 2006 leg. L.V. Graf/. All paratypes with an additional label /Scolytocis fritzplaumanni Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 25): TL 1.58–2.00 (1.75 ± 0.11); PL 0.63–0.79 (0.68 ± 0.05); PW 0.68–0.89 (0.79 ± 0.04); EL 0.89–1.21 (1.03 ± 0.09); EW 0.74–0.95 (0.85 ± 0.05); GD 0.74–0.89 (0.79 ± 0.04). Ratios: PL/PW 0.8–0.94 (0.86 ± 0.05); EL/EW 1.07–1.44 (1.2 ± 0.08); EL/PL 1.28–1.92 (1.51 ± 0.14); GD/EW 0.88– 1 (0.92 ± 0.04); TL/EW 1.89–2.19 (2.05 ± 0.08).

Distribution Known from the South Region of Brazil, from various localities in the states of Paraná, Santa Catarina and Rio Grande do Sul (Fig. 101).

Host fungi Ganodermataceae, in Ganoderma sp. (specimens from Guaratuba), and Ganoderma australe (Fr.) Pat. (specimens from São Francisco de Paula; L.V. Graf pers. comm.).

Comments This is a common Scolytocis from the South Region of Brazil, being easily collected.

Scolytocis furieriae Lopes-Andrade sp. nov. (Figs 10, 85–87, 92, 94B, 95B, 98, 101)

Etymology The species is named after the ecologist, entomologist and great collector of Ciidae, Karina Schimdt Furieri.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 29 Diagnosis The species belongs to the fritzplaumanni species-group and so it is distinguishable from the other Scoly- tocis by the biconcave prosternum without longitudinal carina at midline, and the smooth border of the posterior pronotal margin. It differs from Scol. fritzplaumanni sp. nov. by the fine and sparse pronotal punctation, and from Scol. paschoali sp. nov. by the smaller body (usually less than 1.5mm) and smaller antennal-club (around 1.6X longer than funicle).

Description Holotype. (Figs 85–87) Measurements in mm: TL 1.32; PL 0.53; PW 0.63; EL 0.74; EW 0.68; GD 0.63. Ratios: PL/PW 0.84; EL/EW 1.09; EL/PL 1.40; GD/EW 0.93; TL/EW 1.94. Body dark brown; antennae, mouthparts and legs dark yellowish brown. Dorsum of head with small, shallow and sparse punctures; interstice between punctures distinctly microreticulate. Each antenna (Fig. 94B) with nine antennomeres (FL 0.088mm; CL 0.125mm; CL/FL 1.43); length of the antennomeres (in mm) as follows: 0.080; 0.035; 0.043; 0.015; 0.015; 0.015; 0.035; 0.035; 0.055. Eyes with greatest width 1.25X the basal width of scutellum. Prono- tum finely and sparsely punctate; puncture small, shallow, separate by a distance of two to three puncture widths; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin without a rugose border. Scutellum slightly rugose, basal width 0.17X the EW. Elytra with punctation confused, coarser and denser than that of pronotum; punctures irregular, deep, coalescent near the margin; interstice between punctures smooth to finely rugose; elytral apex truncate, bearing sparse and small cuticular globules, with weak concavity when seen from above. Each hindwing (Fig. 92) with apical area bearing two distinct pigmented lines, one near the anterior margin and the other almost reaching the posterior margin. Prosternum biconcave, without conspicuous longitudinal carina at midline; surface shiny, slightly rugose. Metaventrite glabrous, finely granulose at disc; either side bearing some large obsolescent punctures, with rugose to striated interstice; discrimen half the length of the metaventrite at midline. Abdominal ventrites bearing some small and sparse setae; surface finely granulose. Each metatibia (Fig. 95B) broadly rounded, around 3X as long as broad; outer apical and outer lateral edges forming an obtuse rounded angle; outer edge with spines (around 20) closer to each other at apical one-fourth, then getting sparser. Male genitalia (in paratypes). (Figs 98) Api- cal portion of tegmen subpentagonal; median lobe around 1.3X longer than tegmen.

Type series Holotype. (MZSP) Brazil: /BRASIL: BA Jussari; “RPPN Serra do Teimoso” 23.iii a 17.iv.2005 leg. K. S. Furieri / Scolytocis furieriae Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Brazil: 7 specimens (6 LAPC, 1 UFES), same data as holotype; 70 specimens (5 ANIC, 6 CNCI, 2 GOPC, 2 ISNB, 5 LUND, 3 MHNG, 5 SMTD) /BRASIL: BA Eunápolis xi.2004 JNC Louzada leg./; 63 specimens (5 ANIC, 2 CAMB, 6 CNCI, 1 UFES, 2 MCZ, 5 LUND, 3 MHNG, 5 SMTD) /BRASIL: ES Linhares 11-21.x.2004 P. C. Grossi leg./; 12 specimens (LAPC) /BRASIL: PA Marabá; “Reserva Biológica de Tapirapé; Amazônia Legal” 19.xi.2003 leg. F. Gumier-Costa/; 1 specimen (LAPC) /BRASIL: PA Marabá 03.xii.2003 leg. F. Gumier- Costa/; 3 specimens /BRASIL: ES Sooretama 27.i.2005 legs KS Furieri, R Kawada & JLA Campos/. All paratypes with an additional label /Scolytocis furieriae Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 38, including the holotype): TL 1.16–1.58 (1.39 ± 0.09); PL 0.47–0.58 (0.55 ± 0.04); PW 0.53–0.68 (0.62 ± 0.04); EL 0.68–1.00 (0.80 ± 0.07); EW 0.58–0.74 (0.67 ± 0.04); GD 0.53–0.68 (0.62 ± 0.04). Ratios: PL/PW 0.75–1.00 (0.88 ± 0.06); EL/EW 1.00–1.35 (1.19 ± 0.09); EL/PL 1.25–1.89 (1.48 ± 0.16); GD/EW 0.78–1.00 (0.92 ± 0.04); TL/EW 1.78–2.25 (2.07 ± 0.10).

30 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE Distribution Known from various localities in the states of Pará (North Region), Bahia (Northeast Region), and Espírito Santo (Southeast Region) (Fig. 101).

Host fungi Unknown.

Comments Scol. furieriae sp. nov. was first collected by Gumier-Costa (2004), who found it only in a primary forest of the Amazon Basin. The type locality is Serra do Teimoso (Jussari, BA), a small forest remnant (around 200ha) maintained as a “Reserva Particular do Patrimônio Natural” (RPPN, Private Reserve of Natural Patri- mony). The species is sympatric to Falsocis brasiliensis Lopes-Andrade (an endangered ciid species; Lopes- Andrade 2007b).

Scolytocis paschoali Lopes-Andrade sp. nov. (Figs 1–4, 88–90, 93, 94C, 95C, 99–101)

Etymology The species is named after Paschoal Coelho Grossi, entomologist, good friend, and great collector of ciids.

Diagnosis The species belongs to the fritzplaumanni species-group and so it is distinguishable from the other Scoly- tocis by the biconcave prosternum without longitudinal carina at midline, and the smooth border of the posterior pronotal margin. It differs from the other species in the group by the combination of a fine and sparse pronotal punctation and a large antennal club (2X longer than funicle).

Description Holotype. (Figs 88–90) Measurements in mm: TL 1.70; PL 0.60; PW 0.75; EL 1.10; EW 0.85; GD 0.75. Ratios: PL/PW 0.80; EL/EW 1.29; EL/PL 1.83; GD/EW 0.88; TL/EW 2.00. Body black, almost entirely glabrous; legs brown; antennae, mouthparts and tarsi yellowish brown. Head coarsely punctate; punctures small, with interstice microreticulate. Each antenna (Fig. 94C) with nine antennomeres (FL 0.093mm; CL 0.190mm; CL/FL 2.05); length of the antennomeres (in mm) as follows: 0.105; 0.045; 0.050; 0.013; 0.015; 0.015; 0.053; 0.053; 0.085. Eyes with greatest width 1.5X the basal width of scutellum. Pronotal punctation sparse; punctures shallow and narrow, separated by a distance of one to three puncture widths; interstice between punctures finely and conspicuously microreticulate; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin without a rugose border. Scutellum impunctate, but microreticulate; basal width 0.14X the EW. Elytra with confuse punctation, coarser and denser than that of pronotum; punctures deep, usually separated by a distance of two puncture widths or less; interstice between punctures smooth, shiny; elytral apex truncate, apical border slightly concave when seen from above, bearing small, sparse cuticular globules. Each hindwing (Fig. 93) with apical area bearing two distinct pigmented lines, one near the anterior margin and the other almost reaching the posterior margin. Prosternum biconcave, with very narrow longitudinal carina at midline; surface shiny, inconspicuously rugose. Metaventrite glabrous; disc finely microreticulate, with discrimen half of its length at midline; either side bearing some large obsolescent punctures, with interstice microreticulate to striated. Abdominal ventrites glabrous, microreticulate. Each metatibia (Fig. 95C) broadly rounded, around 3X as long as broad; outer apical and outer lateral edges forming an obtuse rounded angle; outer edge with spines (around 20) close to each other at the apex and getting sparser until its base. Male genitalia (in paratypes). (Fig. 99–100) Apical portion of tegmen with sides subparallel; median lobe slightly longer than tegmen.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 31 Type series Holotype. (MZSP) Brazil: /BRASIL: RJ Macaé de Cima 03.xi.2003 leg. P.C. Grossi / Scolytocis paschoali Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Brazil: 42 specimens, same data as holotype (5 ANIC, 1 CAMB, 2 CNCI, 1 GOPC, 24 LAPC, 2 LUND, 2 MHNG, 4 RRPC, 2 SMTD); 1& (LAPC) /Brasil: Venda Nova do Imigrante, ES 13-16.xi.1999 leg. F. Z. Vaz-de-Mello/; 2 specimens /BRASIL: ES St. Teresa; “ESBL” 25-26.xi.2003; legs K.S. Furieri, G.R. Loiola/. All paratypes with an additional label / Scolytocis paschoali Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 11, including the holotype): TL 1.53–1.74 (1.66 ± 0.09); PL 0.58–0.68 (0.62 ± 0.04); PW 0.68–0.79 (0.72 ± 0.04); EL 0.89–1.10 (0.99 ± 0.07); EW 0.74–0.85 (0.80 ± 0.04); GD 0.68–0.79 (0.72 ± 0.04). Ratios: PL/PW 0.80–0.92 (0.85 ± 0.04); EL/EW 1.13–1.33 (1.24 ± 0.06); EL/PL 1.47–1.83 (1.60 ± 0.13); GD/EW 0.86–0.94 (0.90 ± 0.03); TL/EW 1.94–2.20 (2.09 ± 0.08). Distribution Known from the type locality in Nova Friburgo (in the state of Rio de Janeiro), Santa Teresa and Venda Nova do Imigrante (in the state of Espírito Santo), Southeast Region of Brazil (Fig. 101).

Host fungi Unknown.

Comments The species seems to be closest related to Scol. furieriae sp. nov.

Indo-Malayan, Australasian and Oceanic species

Scolytocis malayanus Lopes-Andrade sp. nov. (Figs 102–104, 120, 126A, 128A, 130, 134)

Etymology The specific epithet refers to Malaysia, terra typica of this species.

Diagnosis The species belongs to the lawrencei species-group, and so is distinguishable from the other Scolytocis by the smooth posterior margin of pronotum and the concave prosternum. It differs from all the other species in this group by the surfaces of the prosternum, metaventrite and abdominal ventrites, which have a distinct microreticulation in high relief.

Description Holotype. (Fig. 102–104) Measurements in mm: TL 1.05; PL 0.42; PW 0.42; EL 0.63; EW 0.47; GD 0.47. Ratios: PL/PW 1.00; EL/EW 1.33; EL/PL 1.50; GD/EW 1.00; TL/EW 2.22. Body black; antennae, mouthparts and legs dark yellowish brown. Head sparsely punctate; punctures shallow, interstice finely microreticulate. Each antenna (Fig. 126A) with nine antennomeres (FL 0.068mm; CL 0.125mm; CL/FL 1.85); length of the antennomeres (in mm) as follows: 0.060; 0.028; 0.030; 0.013; 0.013; 0.013; 0.033; 0.038; 0.055. Eyes with greatest width 1.2X the basal width of scutellum. Pronotum with coarse, sparse, shallow punctation; puncture irregular, separated by a distance of two to three puncture widths, interstice conspicuously microreticulate; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin without a rugose border. Scutellum impunctate; basal width 0.16X the EW. Elytra with subseriate punctation; punctures

32 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE coarser and shallower than those of pronotum; border of punctures indistinct, interstice slightly rugose; elytral apex rounded, glabrous, its declivity with a weak concavity, cuticular globules absent. Each hindwing (Fig. 120) with apical area without conspicuous pigmented lines. Prosternum concave, without longitudinal carina; surface with distinct rugosity. Metaventrite glabrous, finely rugose; discrimen one-fourth the length of the metaventrite at midline. Abdominal ventrites glabrous, rugosity similar to that of prosternum and metaventrite. Each metatibia (Fig. 128A) near 3X as long as broad; outer apical and outer lateral edges forming an almost perpendicular angle that is rounded; outer edge with spines (around 15) regularly distributed at apical two-thirds, then getting sparser. Male genitalia (in a paratype). (Fig. 130) Genitalia barely sclerotized; apex of tegmen acute; median lobe extremely membranous and elongate, around 1.8X longer than tegmen.

Type series Holotype. (ANIC) Malaysia: /Malaysia, Pahang Temerloh Distr. Krau Game Res. Feb. 18, 1975 J. Flea- gle, coll. / Ganoderma sp./ J. F. Lawrence Lot [printed] 3911 [handwritten]/ Scolytocis malayanus Lopes- Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Malaysia: 2 specimens (ANIC), same locality data as holotype; 2 specimens (LAPC), /Malaysia, Pahang Temerloh Distr. Krau Game Res. Feb. 18, 1975 J. Fleagle, coll. / Phellinus sp. / J. F. Lawrence Lot [printed] 3906 [handwritten]/; 2 specimens (1 ANIC, 1 LAPC) /Malaysia: Perak N. slopes Gunong Bubu, K. Kangsar Dist. Feb. 2, 1975 J. Fleagle, coll. / Phellinus sp./, one additionally labeled /J. F. Lawrence Lot [printed] 3765 [handwritten]/, and the other /J. F. Lawrence Lot [printed] 3768 [handwritten]/. All paratypes with an additional label /Scolytocis malayanus Lopes- Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 5, including the holotype): TL 1.00–1.16 (1.08 ± 0.06); PL 0.37–0.42 (0.40 ± 0.03); PW 0.42–0.47 (0.43 ± 0.02); EL 0.63–0.74 (0.66 ± 0.05); EW 0.42–0.53 (0.47 ± 0.04); GD 0.42–0.47 (0.44 ± 0.03). Ratios: PL/PW 0.88–1.00 (0.93 ± 0.07); EL/EW 1.20–1.56 (1.41 ± 0.14); EL/PL 1.50–1.86 (1.66 ± 0.16); GD/EW 0.89–1.00 (0.94 ± 0.06); TL/EW 2.10–2.44 (2.30 ± 0.14).

Distribution Known only from Malaysia (Fig. 134).

Host fungi Ganoderma sp. (Ganodermataceae), Phellinus sp. (Hymenochaetaceae).

Comments The placement of this species in the lawrencei species-group is justifiable, since it resembles other species of the group not only by external morphological features but also by the morphology of male genitalia (com- pare Figs 78 and 130).

Scolytocis novaezelandiae Lopes-Andrade sp. nov. (Figs 105–107, 121, 126B, 128B, 135)

Etymology The specific epithet refers to New Zealand, terra typica of this species.

Diagnosis The species belongs to the werneri species-group, and so is distinguishable from the other Scolytocis by the combination of a rugose border along the posterior pronotal margin and a concave prosternum. It lacks the

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 33 long metaventrite discrimen of Scol. werneri sp. nov. and has a broader rugose border along the posterior pronotal margin. Besides the species in this group, the other Scolytocis with such rugose border all have a biconcave prosternum with a narrow longitudinal carina at midline (the danielssoni group). Description Holotype. (Figs 105–107) Measurements in mm: TL 1.21; PL 0.42; PW 0.53; EL 0.74; EW 0.58; GD 0.53. Ratios: PL/PW 0.80; EL/EW 1.27; EL/PL 1.75; GD/EW 0.91; TL/EW 2.09. Body dark brown; antennae and mouthparts yellowish brown; legs reddish brown. Head with coarse, sparse, shallow punctation; interstice between punctures inconspicuously microreticulate, shiny. Each antenna (Fig. 126B) with nine antennomeres (FL 0.083mm; CL 0.133mm; CL/FL 1.61); length of the antennomeres (in mm) as follows: 0.088; 0.038; 0.038; 0.015; 0.015; 0.015; 0.038; 0.038; 0.058. Eyes with greatest width 1.0X the basal width of scutellum. Pronotum with coarse, shallow, confuse punctation; punctures irregular, separated by a distance of one to three puncture widths; interstice between punctures microreticulate, being much more conspicuous near the lateral margin; anterolateral margins slightly arched inwards; posterolateral angles broadly rounded; posterior margin bearing a conspicuous rugose border along it. Scutellum impunctate, rugose; basal width 0.16X the EW. Elytra with confuse punctation, coarser and denser than that of pronotum; interstice between puncture finely microreticulate near the anterior margin, then getting rugose posteriorly; elytral apex truncate, declivity conspicuously concave and bearing very small cuticular globules. Each hindwing (Fig. 121) with apical area without conspicuous pigmented lines. Prosternum concave, without longitudinal carina at midline; surface regularly granulose. Metaventrite glabrous, regularly granulose; discrimen one-sixth the length of the metaventrite at midline. Abdominal ventrites glabrous, regularly granulose. Each metatibia (Fig. 128B) near 3.25X as long as broad; outer apical and outer lateral edges forming an obtuse angle that is broadly rounded; outer edge with spines (around 15) regularly distributed at apical two-thirds. Male genitalia. Unknown.

Type series Holotype. (ANIC) New Zealand: /New Zealand: ND: Waipoua SF, Wairau Summit 387m 11-14 iv.1980 podocarp-mixed broadleaf A. Newton, M. Thayer/ Ganoderma sp. / Scolytocis novaezelandiae Lopes- Andrade HOLOTYPUS [printed on red paper]/. Paratypes. New Zealand: 1 specimen, same data as holotype (ANIC); 1 specimen (LAPC) /New Zealand, CL Kirikiri Saddle, 835m Coromandel Range 1 Feb. 1986 J. F. Lawrence / J. F. Lawrence Lot Ganoderma sp./. The paratypes with an additional label /Scolytocis novaezelandiae Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 3, including the holotype): TL 1.16–1.26 (1.21 ± 0.05); PL 0.42–0.47 (0.44 ± 0.03); PW 0.47–0.58 (0.53 ± 0.05); EL 0.68–0.79 (0.74 ± 0.05); EW 0.53–0.63 (0.58 ± 0.05); GD 0.47–0.58 (0.53 ± 0.05). Ratios: PL/PW 0.80–0.89 (0.84 ± 0.05); EL/EW 1.25–1.30 (1.27 ± 0.03); EL/PL 1.63–1.75 (1.68 ± 0.06); GD/EW 0.90–0.92 (0.91 ± 0.01); TL/EW 2.00–2.20 (2.10 ± 0.10).

Distribution Known only from New Zealand (Fig. 135).

Host fungi Ganoderma sp. (Ganodermataceae).

Comments This species was cited by Leschen et al. (2003) as Scolytocis sp.

Etymology The specific epithet refers to the Philippines, terra typica of this species.

Description Holotype. (Figs 108–110) Measurements in mm: TL 1.37; PL 0.47; PW 0.58; EL 0.84; EW 0.63; GD 0.63. Ratios: PL/PW 0.82; EL/EW 1.33; EL/PL 1.78; GD/EW 1.00; TL/EW 2.17. Body with dorsal surface dark reddish brown, ventral surface light reddish brown; basal antennomeres and funicle yellowish brown; labial palpi, maxillary palpi and club brown. Head finely and sparsely punctate on dorsum; interstice between punctures microreticulate, shiny. Each antenna (Fig. 126C) with nine antennomeres (FL 0.08mm; CL 0.17mm; CL/FL 2.04); length of the antennomeres (in mm) as follows: 0.107; 0.033; 0.040; 0.017; 0.013; 0.013; 0.040; 0.050; 0.080. Eyes with greatest width 1.14X the basal width of scutellum. Pronotum with shallow punctation; punctures fine, separated by a distance of three to five puncture widths; interstice between punctures finely microreticulate; anterolateral margins slightly arched inwards; posterolateral angles broadly rounded; posterior margin without rugose border. Scutellum impunctate, inconspicuously microreticulate; basal width 0.19X the EW. Elytra with confuse and dense punctation; punctures coarser and shallower than those of pronotum, each one with the border indistinct from surrounding surface (when seen from above); interstice between punctures rugose, but without microreticulation; elytral apex rounded, declivity with shallow concavity and bearing small cuticular globules. Each hindwing (Fig. 122) with apical area bearing one incomplete pigmented line near the anterior margin. Prosternum biconcave, with conspicuous longitudinal carina at midline; rugose. Metaventrite glabrous; disc smooth, delimited by a circular row of obsolescent punctures; either side rugose, bearing some large obsolescent punctures; discrimen half the length of the metaventrite at midline. Abdominal ventrites almost glabrous, finely rugose. Each metatibia (Fig. 128C) conspicuously expanded at middle, near 3.5X as long as broad; outer margin broadly rounded, equally spinulose from apex to middle, with spines separated by a distance of one spine width, and spines then getting sparse to the base. Male genitalia (in a paratype). (Fig. 131) Median lobe subcylindrical, membranous, elongate, as long as tegmen; tegmen with lateral margins gradually converging from base to apex.

Type series Holotype. (FMNH) Philippines: /PHILIPPINES: Mt. McKiling, Laguna Prov. 4Km SE Los Banos 12-IV- 1977 / ex fungi LEWatrous [sic] / Scolytocis philippinensis Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Philippines: 39 specimens (2 ANIC, 22 FMNH, 14 LAPC, 1 SMTD), same data as holotype. All paratypes additionally labeled /Scolytocis philippinensis Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 10, including the holotype): TL 1.26–1.37 (1.32 ± 0.05); PL 0.42–0.47 (0.47 ± 0.02); PW 0.58–0.63 (0.58 ± 0.02); EL 0.74–0.89 (0.83 ± 0.04); EW 0.58–0.68 (0.62 ± 0.03); GD 0.58–0.63 (0.59 ± 0.03). Ratios: PL/PW 0.73–0.82 (0.80 ± 0.03); EL/EW 1.23–1.45 (1.33 ± 0.07); EL/PL 1.56–1.89 (1.77 ± 0.10); GD/EW 0.85–1.00 (0.96 ± 0.06); TL/EW 2.00–2.27 (2.13 ± 0.09).

Host fungi Unknown.

Scolytocis samoensis Blair, 1928

Diagnosis and comments Scol. samoensis was described based on just one specimen, probably a female, deposited in BPBM (Hawaii, USA). Blair (1928) mentioned that the holotype is very defective. From the description provided by Blair (1928), Scol. samoensis may be distinguished from the other Scolytocis described here by the completely rounded humeri and the first abdominal ventrite longer than the next two ventrites together. In the other Scolytocis the first abdominal ventrite is as long as or shorter than the next two ventrites together, and the humeri is always angulate. As mentioned by Blair, the tarsi of Scol. samoensis appear to have three tarsomeres. The basal tarsomere of all Scolytocis examined in this study is extremely small and barely visible, even at high magnifications (over 100X under stereomicroscope). However, in slide preparations, I could evaluate that all species have four tarsomeres in each tarsus. The same is probably true for Scol. samoensis. A small discussion on the hypo- thetical presence of three tarsomeres (rather than four) in each tarsus of some ciid species is provided by Lopes-Andrade (2007a). Due to its shorter length, the seriate elytral punctation, and the antennal club that is almost twice as long as the funicle, Scol. samoensis may be tentatively included in the lawrencei species-group.

Distribution Known only from the type locality: Tutuila, Pago Pago, Samoa (Polynesia; Fig. 135).

Scolytocis thayerae Lopes-Andrade sp. nov. (Figs 111–113, 123, 127A, 129A, 132, 134)

Etymology The species is named after the great entomologist Margaret Thayer (Field Museum of Natural History), who kindly authorized the loan of the specimens from the Philippines and many Mesoamerican Scolytocis.

Description Holotype. (Figs 111–113) Measurements in mm: TL 1.89; PL 0.68; PW 0.79; EL 1.16; EW 0.84; GD 0.84. Ratios: PL/PW 0.87; EL/EW 1.38; EL/PL 1.69; GD/EW 1.00; TL/EW 2.25. Body with dorsal and ventral surfaces dark reddish brown; basal antennomeres, labial and maxillary palpi, and tarsi light reddish brown; legs reddish brown. Head sparsely punctate on dorsum; interstice between punctures microreticulate. Each antenna (Fig. 127A) with 9 antennomeres (FL 0.10mm; CL 0.21mm; CL/FL 2.10); length of the antennomeres (in

36 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE mm) as follows: 0.127; 0.040; 0.047; 0.020; 0.020; 0.013; 0.033; 0.043; 0.133. Eyes with greatest width 1.07X the basal width of scutellum. Pronotum with shallow punctation; punctures fine at disc, separated by a distance of two to four puncture widths; near the lateral margins, the punctures are conspicuously coarser and closer than those at disc, being separated by one puncture width or less, but not coalescent; interstice between punctures conspicuously microreticulate; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin without a rugose border. Scutellum impunctate, but microreticulate; basal width 0.19X the EW. Elytra with confuse punctation, which is shallower, denser and coarser than that of pronotum; interstice between punctures rugose, shiny; elytral apex truncate; apical declivity concave, with inconspicuous cuticular globules. Each hindwing (Fig. 123) with apical area bearing two distinct pigmented lines, one near the anterior margin and the other almost reaching the posterior margin. Prosternum biconcave, bearing a very thin, inconspicuous longitudinal carina at midline; surface rugose. Metaventrite glabrous; surface finely microreticulate; discrimen absent; either side bearing some large obsolescent punctures. Abdominal ventrites microreticulate; bearing some small and sparse setae. Each metatibia (Fig. 129A) conspicuously expanded at middle, near 3X as long as broad; outer margin broadly rounded, equally spinulose from apex to middle, with spines very close to each other and then getting sparse to the base. Male genitalia (in a paratype). (Fig. 132) Median lobe subcylindrical, membranous, elongate, as long as tegmen; tegmen with lateral margins gradually converging from base to apex.

Type series Holotype. (FMNH) Philippines: /CAMP MERAN E [handwritten] slope Mt. Apo, Davo Prov. Mindanao, P.I. [printed] XI-46 [handwritten] / F.G. Werner Collector / Scolytocis thayerae Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Philippines: 10 specimens (5 FMNH, 5 LAPC), same data as holotype; 1 specimen (FMNH) / CAMP MERAN E [handwritten] slope Mt. Apo, Davo Prov. Mindanao, P.I. [printed] XI- 5-46 [handwritten] / F.G. Werner Collector / ex Fomes applanatum on [sic]/; 1 specimen (FMNH) / LAKE LINAU N [handwritten] slope Mt. Apo, Davo Prov. Mindanao, P.I. [printed] X-XI-46 [handwritten] / F.G. Werner Collector/; 9 (1 ANIC, 6 FMNH, 3 LAPC) specimens /E slope Mt.Mckinley Davao Prov, Mindanao P.I. [printed] IX.3.46 6500’ [handwritten] / F.G. Werner Collector / Lot 43 [handwritten]/; 3 specimens (2 FMNH, 1 LAPC) /Lake Linau, N. slope Mt. Apo, Davao Prov., MINDANAO. El. 7900ft. mossy forest; ex rotten fungus X-25 to XI-4, 1946 / CNHM. Philippine Zool. Exped. (1946-47) F.G. Werner leg./. All paratypes additionally labeled /Scolytocis thayerae Lopes-Andrade PARATYPUS [printed on yellow paper]/. Variation Measurements in mm (n = 10, including the holotype): TL 1.79–2.05 (1.94 ± 0.09); PL 0.63–0.74 (0.71 ± 0.04); PW 0.79–0.89 (0.83 ± 0.04); EL 1.05–1.26 (1.17 ± 0.06); EW 0.84–0.95 (0.90 ± 0.05); GD 0.79–0.89 (0.86 ± 0.04). Ratios: PL/PW 0.80–0.88 (0.85 ± 0.03); EL/EW 1.25–1.38 (0.30 ± 0.04); EL/PL 1.54–1.75 (1.67 ± 0.06); GD/EW 0.94–1.00 (0.95 ± 0.02); TL/EW 2.11–2.25 (2.15 ± 0.05).

Distribution Known from the islands of Mindanao and Luzon in the Philippines, at the Mounts Apo and Makiling, respectively (Fig. 134).

Host fungi Ganoderma applanatum (Pers.) Pat. (Ganodermataceae).

Scolytocis werneri Lopes-Andrade sp. nov. (Figs 114–116, 124, 127B, 129B, 133, 134)

Etymology The specific epithet is in honour of the great entomologist Floyd Gerald Werner (1921-1992), who col-

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 37 lected the known specimens of the species. He also collected the known specimens of Scol. thayerae sp. nov. and Xylographella speciosa sp. nov., all during the Philippine Zoological Expedition (1946 to 1947), spon- sored by the Chicago Museum of Natural History (now the Field Museum of Natural History).

Diagnosis The species belongs to the werneri species-group, and so is distinguishable from the other Scolytocis by the combination of a concave prosternum and the presence of a rugose border along the posterior margin of pronotum. It differs from Scol. novaezelandiae, the other species of the group, by its narrower rugose border of pronotum and the metaventrite bearing a long discrimen.

Description Holotype. (Figs 114–116) Measurements in mm: TL 1.26; PL 0.42; PW 0.53; EL 0.84; EW 0.58; GD 0.58. Ratios: PL/PW 0.80; EL/EW 1.45; EL/PL 2.00; GD/EW 1.00; TL/EW 2.18. Body dark brown; antennae, mouthparts and legs yellowish brown. Head sparsely punctate; interstice between punctures inconspicuously microreticulate, shiny. Each antenna (Fig. 127B) with nine antennomeres (FL 0.07mm; CL 0.12mm; CL/FL 1.59); length of the antennomeres (in mm) as follows: 0.067; 0.037; 0.033; 0.013; 0.013; 0.013; 0.033; 0.030; 0.053. Eyes with greatest width 1.3X the basal width of scutellum. Pronotum with coarse, sparse punctation; punctures irregular, separated by a distance of one to three puncture widths; interstice between punctures finely microreticulate. Scutellum impunctate, rugose; basal width 0.14X the EW. Elytra with confuse punctation, coarser and denser than that of pronotum; interstice between punctures smooth to slightly rugose, shiny, but not microreticulate; elytral apex truncate, declivity slightly concave and bearing small, inconspicuous cuticular globules. Each hindwing (Fig. 124) with apical area without conspicuous pigmented line, but one vague, inconspicuous line near the posterior margin. Prosternum concave, without longitudinal carina at midline; surface finely rugose. Metaventrite bearing a distinct discrimen along its midline, beginning at the posterior margin and almost reaching the anterior; surface finely granulose at disc and rugose on either side. Abdominal ventrite glabrous, finely microreticulate. Each metatibia (Fig. 129B) near 3X as long as broad; outer apical and outer lateral edges forming an obtuse angle that is broadly rounded; apex with outer edge bearing almost 15 spines very close to each other, outer lateral edge with near five spines regularly distributed. Male genitalia (in a paratype). (Fig. 133) Median lobe subcylindrical, membranous, elongate, as long as tegmen; tegmen with lateral margins converging from the middle to the apex. Type series Holotype. (FMNH) Philippines: / CAMP MERAN E [handwritten] slope Mt. Apo, Davo Prov. Mind- anao, P.I. [printed] XI-5-46 [handwritten] / F.G. Werner Collector / Scolytocis werneri Lopes-Andrade HOLO- TYPUS [printed on red paper]/. Paratypes. Philippines: 3 specimens (1 FMNH, 2 LAPC), same data as holotype, but one additionally labeled /ex Fomes applanatus on [sic]/. All paratypes additionally labeled /Sco- lytocis werneri Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 4, including the holotype): TL 1.26–1.32 (1.29 ± 0.03); PL 0.42–0.42 (0.42 ± 0); PW 0.47–0.53 (0.51 ± 0.03); EL 0.79–0.84 (0.83 ± 0.03); EW 0.53–0.58 (0.57 ± 0.03); GD 0.47–0.58 (0.54 ± 0.05). Ratios: PL/PW 0.80–0.89 (0.82 ± 0.04); EL/EW 1.45–1.50 (1.47 ± 0.02); EL/PL 1.88–2.00 (1.97 ± 0.06); GD/EW 0.90–1.00 (0.95 ± 0.06); TL/EW 2.18–2.40 (2.28 ± 0.09).

Distribution Known from a single collection at the Mount Apo, in the island of Mindanao, Philippines (Fig. 134).

Host fungi Ganoderma applanatum (Ganodermataceae).

Etymology The species is named after the outstanding entomologist Elwood Curtin Zimmerman (1912-2004), who collected the three known specimens of the species.

Diagnosis The species belongs to the zimmermani species-group, in which the species have each tibia expanded to the middle, with its outer margin broadly rounded and bearing several spines very close to each other at the apical half. It differs from Scol. thayerae sp. nov. mainly by its smaller body (less than 1.7mm) and from Scol. philippinensis sp. nov. by the conspicuous larger apical antennomere. Description Holotype. (Figs 117–119) Measurements in mm: TL 1.58; PL 0.63; PW 0.68; EL 0.89; EW 0.68; GD 0.68. Ratios: PL/PW 0.93; EL/EW 1.31; EL/PL 1.41; GD/EW 1.00; TL/EW 2.32. Body with dorsum dark brown, ventral surface reddish brown; basal antennomeres, mouthparts and tarsi yellowish brown. Head finely and sparsely punctate on dorsum; interstice between punctures smooth, shiny, rugose near the anterior margin. Each antenna (Fig. 127A) with nine antennomeres (FL 0.113mm; CL 0.195mm; CL/FL 1.73); length of the antennomeres (in mm) as follows: 0.100; 0.043; 0.065; 0.020; 0.015; 0.013; 0.045; 0.050; 0.100. Eyes with greatest width 1.11X the basal width of scutellum. Pronotum with shallow punctation; punctures fine at disc, separated by a distance of two to three puncture widths; near the lateral margins, the punctures are coarse and somewhat elongate, separated by one puncture width or less, but not coalescent; interstice between punctures finely and inconspicuously microreticulate; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin without a rugose border. Scutellum impunctate, shiny; basal width 0.21X the EW. Elytra with confuse punctation, which is shallower, denser and coarser than that of pronotum; interstice between punctures smooth, shiny; elytral apex truncate; apical declivity concave, with inconspicuous cuticular globules. Each hindwing (Fig. 125) with apical area bearing two distinct pigmented lines, one near the anterior margin and the other almost reaching the posterior margin. Prosternum concave, without longitudinal carina at midline; surface finely rugose. Metaventrite glabrous, with disc delimited by a circular row of obsolescent punctures, finely microreticulate; either side microreticulate, and bearing some obsolescent punctures; discrimen one-sixth the length of metaventrite at midline. Abdominal ventrites glabrous, finely microreticulate. Each metatibia (Fig. 129C) conspicuously expanded at middle, 3.5X as long as broad; outer margin broadly rounded, equally spinulose from apex to middle, with spines very close to each other and then getting sparse to the base. Male genitalia. Unknown.

Type series Holotype. (ANIC) Fiji: /Wainiloka Ovalau, Fiji [printed] VII-II-38 [handwritten]/ Shelf Fungi / ECZim- merman Collector / Scolytocis zimmermani Lopes-Andrade HOLOTYPUS [printed on red paper]/. Paratypes. Fiji: 2 specimens (1 ANIC, 1 LAPC), same data as holotype (1 ANIC, 1 LAPC). The paratypes with an additional label /Scolytocis zimmermani Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 3): TL 1.42–1.58 (1.53 ± 0.09); PL 0.58–0.63 (0.60 ± 0.03); PW 0.63–0.68 (0.66 ± 0.03); EL 0.79–0.95 (0.88 ± 0.08); EW 0.63–0.68 (0.66 ± 0.03); GD 0.63–0.68 (0.65 ± 0.03). Ratios: PL/PW 0.85–0.93 (0.90 ± 0.04); EL/EW 1.25–1.40 (1.32 ± 0.08); EL/PL 1.36–1.64 (1.47 ± 0.15); GD/EW 0.93–1.00 (0.98 ± 0.04); TL/EW 2.25–2.32 (2.30 ± 0.04).

Host fungi Unknown.

Comments Although the morphology of male genitalia is not provided for this species, it is sufficiently different to allow the description at this moment.

Syncosmetina Lopes-Andrade subtr. nov. (Figs 7–8, 11–12, 136–159)

Included genera Syncosmetus Sharp, 1891 (type genus) Tropicis Scott, 1926

Diagnosis This new subtribe is characterized by the combination of the following characters: (i) lateral pronotal carina always crenulated; (ii) pronotal and elytral punctation very coarse and dense, punctures coalescent in some portions; (iii) scutellum inconspicuous, its basal width less than 0.10X the greatest elytral width; (iv) elytra bearing longitudinal keels in most species; (v) prosternal process broad; (vi) procoxae cylindrical, at the same level of the prosternal process; (vii) apex of prementum rounded.

Distribution Syncosmetina subtrib. nov. is known to occur in parts of the Afrotropical (restricted to the western Indian Ocean; Tropicis) and Palearctic (restricted to Japan; Syncosmetus) Regions.

Syncosmetus Sharp, 1891 (Figs 11, 113–122) Omogocis Miyatake, 1954 (synonymised by Miyatake 1970)

Included species Syncosmetus japonicus Sharp, 1891 (type species) Syncosmetus reticulatus Miyatake, 1985

Diagnosis and comments Syncosmetus is distinguishable from Tropicis by the combination of the following features: (i) body suboval to oval; (ii) humeri rounded; (iii) hindwings rudimentary; (iv) female terminalia bearing gonostyli. The genus has two described species, both from Japan, and an identification key in Japanese was provided by Kawanabe (2005).

Identification key to the species of Syncosmetus Sharp

1 Lateral margins of pronotum broad, explanate (Fig. 136). Elytra with longitudinal keels prominent, extending from base to apex (Figs 136–137); megapunctures of the interstice between keels at least twice as large as pronotal punctures (Fig. 136)...... Syncosmetus japonicus Sharp - Lateral margins of pronotum narrow, not explanate (Fig. 139). Elytra with longitudinal keels weak, being more conspicuous at the apical two-thirds (Figs 139–140); punctures of the interstice between keels as

Syncosmetus japonicus Sharp, 1891 (Figs 11, 136–138, 142A, 143A, 144–145) Omogocis tuberculifrons Miyatake, 1954 (synonymised by Miyatake 1970)

Diagnosis and comments The species differs from Syn. reticulatus by the more convex and oval body (Figs 136–138), with more marked keels and larger punctation on elytra. The male genitalia (Fig. 144) is well sclerotized, with a cylindrical median lobe that is longer than the tegmen and as long as the ninth segment. The basal piece is distinct, with almost half the length of tegmen, the anterior margin broadly rounded and the posterior margin truncate. Specimens examined Japan. Three specimens (LAPC) labeled /JAPAN, Shikoku Ishizuchi Mt. N.P. M. Tsutsujo, 1600m 14.VIII.80, S. Peck / Betula-Fagus logs, stump & moss litter/; one specimen /JAPAN, Shikoku Ishizuchi Mt. N.P. Omogo Valley 18-25.VIII.1980 S. Peck, 700m / fungi on logs & moss, mixed warm temp for/. All specimens with an identification label /Syncosmetus japonicus Sharp, 1891 C. Lopes-Andrade det. 2007/.

Host-fungi Pleurotus sp. (Pleurotaceae) (Kawanabe 1995, doubtful report), Trametes versicolor (L.) Lloyd (Polypo- raceae) (cited as Coriolus versicolor in Kawanabe 1996, as a doubtful report); Fomes fomentarius (L.) J.J. Kickx (Polyporaceae), Pyrrhoderma sendaiense (Yasuda) Imazeki (Hymenochaetaceae) and Elfvingia applanata (Pers.) P. Karst. (Ganodermataceae) (Kawanabe 1998); Inonotus mikadoi (Lloyd) Gilb. & Ryvarden (Hymenochaetaceae) (Kawanabe 1999).

Distribution Known from Japan (Fig. 145).

Syncosmetus reticulatus Miyatake, 1985 (Figs 7, 139–141, 142B, 143B, 145)

Diagnosis and comments The species differs from Syn. japonicus by the more parallel-sided body (Figs 139–141) and less marked keels on elytra. No males were examined, so it was not possible to evaluate whether some of the distinct characters of the male genitalia of Syn. japonicus are restricted to the species or diagnostic of the genus.

Specimen examined Japan: One && labeled / JAPAN NARA Pref. Mt. Obako-dake, 1300m near Nosegawa-mura vill. 25 III 2006, leg. P. Jaloszynski [printed] / Syncosmetus reticulatus [handwritten] det. Rafal RUTA 2005 [sic] [printed]/

Host-fungi Fomes fomentarius (L.) J.J. Kickx (Polyporaceae) and Elfvingia applanata (Pers.) P. Karst. (Ganoder- mataceae) (Kawanabe 1998). Regarded as belonging to the Ganoderma host-use group (Orledge & Reynolds 2005).

Distribution Known from Japan (Fig. 145).

Included species Tropicis brevicarinatus Scott, 1926 Tropicis cuccodoroi Lopes-Andrade sp. nov. Tropicis flexicarinatus Scott, 1926 Tropicis sexcarinatus (Waterhouse, 1876) (type species)

Diagnosis and comments Very similar to Syncosmetus, but all the known species are macropterous. Consequently, the body of Trop- icis is subparallel-sided (rather than suboval to oval) with the humeri angulate (rather than rounded).

Identification key to the species of Tropicis Scott

1 Pronotal punctures irregularly polygonal or elongate, with narrow interstice or with interstice microreticulate. Elytral keels extending from the apex to at least the apical two-thirds of elytra...... 3 - Pronotal punctures circular, with interstice smooth and shining. Elytral keels restricted to the apical half of elytra ...... 2 2 Elytral keels distinct, prominent; elytral apex truncate ...... Trop. brevicarinatus Scott - Elytral keels indistinct, restricted to the apical declivity (Figs 146–147); elytral apex broadly rounded (Fig. 146) ...... Trop. cuccodoroi sp. nov. 3 Pronotal punctures shallow, very close to each other, giving an appearance of an irregular honeycomb to the pronotal surface (Fig. 149). Elytral keels obsolescent near base (Figs 149–150) ...... Trop. sexcarinatus (Waterhouse) - Pronotal punctures elongate, separate, with interstice excessively reticulate-striolate. Elytral keels sharply elevated, sinuous, distinct almost to the very base of the elytra...... Trop. flexicarinatus Scott

Tropicis brevicarinatus Scott, 1926

Diagnosis and comments As mentioned by Scott (1926), it differs from the other Tropicis mainly by the short elytral keels. In Trop. cuccodoroi sp. nov the elytral keels are almost indistinct.

Specimen examined No specimen examined.

Host-fungi Unknown.

Distribution Western Indian Ocean, Seychelles (Fig. 158).

Tropicis cuccodoroi Lopes-Andrade sp. nov. (Figs. 146–148, 152, 154A, 155A, 156, 158)

Etymology This species is named after Giulio Cuccodoro, who kindly authorized the loan of the Tropicis from the collection of MHNG.

42 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE Diagnosis The species close resembles Trop. brevicarinatus Scott, and it differs from it and the other described Trop- icis by the indistinct elytral keels.

Description Holotype. (Figs 146–148) Measurements in mm: TL 1.32; PL 0.42; PW 0.53; EL 0.79; EW 0.58; GD 0.53. Ratios: PL/PW 0.80; EL/EW 1.36; EL/PL 1.88; GD/EW 0.91; TL/EW 2.27. Body dark brown; head and pronotum darker than elytra; basal antennomeres, mouthparts and legs yellowish brown; abdominal ventrite brown. Head with conspicuous and coarse punctation; interstice between punctures smooth, shiny. Each antenna (Fig. 154A) with ten antennomeres (FL 0.103mm; CL 0.135mm; CL/FL 1.32); length of the antennomeres (in mm) as follows: 0.07; 0.038; 0.035; 0.025; 0.013; 0.015; 0.015; 0.035; 0.038; 0.063. Eyes with greatest width 1.67X the basal width of scutellum. Pronotum deeply and coarsely punctate; punctures separated by a distance of one puncture width or less; interstice between punctures smooth, shiny; anterolateral margins straight; posterolateral angles broadly rounded; posterior margin without rugose border. Scutellum very small, impunctate, glabrous, shiny; basal width 0.075X the EW. Elytral punctation and interstice similar to that of pronotum; elytral apex rounded, without longitudinal keels; apical declivity with very weak concavity, cuticular globules inconspicuous; elytral suture margined at apical third. Each hindwing (Fig.152) with apical area bearing two distinct pigmented lines, one near the anterior margin and the other almost reaching the posterior margin. Prosternum with fine, regular, circular microreticulation in high relief. Metaventrite and abdominal ventrites glabrous, bearing microreticulation similar to that of prosternum. Each metatibia (Fig. 155A) slender, subtriangular, just slightly expanded to apex; outer lateral edge straight, bearing around 5 sparse spines; outer apical edge bearing around 10 spines, close to each other. Male genitalia (in paratypes). (Fig. 156) Genitalia elongate, subcylindrical, sclerotized; median lobe nearly 1.3X longer than tegmen, around 15X as long as wide.

Type series Holotype. (MHNG) Réunion: /La Réunion Y. Gomy [printed] Pl. des Chicots 21.4.73 - 1860m Tamisage [handwritten]/ Tropicis cuccodoroi Lopes-Andrade HOLOTYPUS [printed on red paper]. Paratypes. Réunion: Total of 26 specimens (7 LAPC; 19 MHNG). 7 specimens /La Réunion Y. Gomy [printed] Cilaos Grand Matarum 1.xi.69 tamisage [handwritten]/; 1 specimen /La Réunion J. David [printed] Foret Bebour Aout 1973 [handwritten]/; 3 specimens /La Réunion Y. Gomy [printed] Brûlé de Mare Longue 23.2.72 tamisage tronc [handwritten]”; 3 specimens /La Réunion Y. Gomy [printed] Takamaka 26.i.72 chemin du barrage Tamisage souche très humide [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Plaine des Makes 1500m-19.xi.70 s/ ecorces tamarin [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Piste Forestier de la Pl. des Affouches 19.11.67 P.K5 [sic] [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Pl. des Makes 14km - 15.xi.70 tamisage troncset humus [handwritten]/; 4 specimens /La Réunion Y. Gomy [printed] Pl. d’Affouches 1200m-1.xi.71 tamisage souche et humus Gite [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] ND de la Paix 31.xii.70-1500m tamisage [handwritten]/; 2 specimens /La Réunion P. Schauenberg [printed] Forêt endémique Ravine Grand Chaloupe -590m 11.i.75 [handwritten]/; 1 specimen / La Réunion Y. Gomy [printed] Plaine des Cafres ND de la Paix 9.i.72 tamisage [handwritten]/; 1 specimen / La Réunion Y. Gomy [printed] Gite Forestiere de Bebour 22.4.72 tamisage humus [handwritten]/. Mauritius: Total of 76 specimens (2 ANIC, 19 LAPC, 55 MHNG). 11 specimens labeled /Ile Maurice Y. Gomy [printed] Macabe Forest 13.i.70 tronc mort [handwritten]/; 5 specimens /Ile Maurice Y. Gomy [printed] Macabe 11.4.70 [handwritten]/; 6 specimens /Ile Maurice Y. Gomy [printed] Magenta 11.4.70 [handwritten]/; 50 specimens / Ile Maurice Y. Gomy [printed] Mont Cocotte 600m 28.i.71 tamisage [handwritten]/; 1 specimen /Ile Maurice Y. Gomy [printed] Mare Longue 19.i.71 tamisage tronc [handwritten]/; 1 specimen /Ile Maurice Y. Gomy [printed] Mont Cocotte 19.i.1970 tronc mort [handwritten]/; 1 specimen /Ile Maurice Y. Gomy [printed]

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 43 Riviere Patate 19.i.71 tamisage tronc [handwritten]/. All paratypes with an additional label / Tropicis cuccodoroi Lopes-Andrade PARATYPUS [printed on yellow paper]/.

Variation Measurements in mm (n = 20, including the holotype): TL 0.95–1.58 (1.19 ± 0.15); PL 0.32–0.53 (0.39 ± 0.05); PW 0.42–0.68 (0.51 ± 0.06); EL 0.63–0.95 (0.74 ± 0.08); EW 0.47–0.74 (0.55 ± 0.07); GD 0.42–0.58 (0.50 ± 0.04). Ratios: PL/PW 0.70–0.89 (0.77 ± 0.04); EL/EW 1.18–1.50 (1.35 ± 0.09); EL/PL 1.63–2.14 (1.90 ± 0.14); GD/EW 0.77–1.00 (0.90 ± 0.06); TL/EW 1.91–2.30 (2.15 ± 0.10).

Distribution Mascarene Islands (La Reunion, Mauritius; Fig. 158). Host-fungi Unknown.

Comments This species resembles Trop. brevicarinatus Scott from Seychelles, but differs in lacking longitudinal keels at the elytral apex.

Tropicis flexicarinatus Scott, 1926

Diagnosis and comments As mentioned by Scott (1926), it differs from Tropicis sexcarinatus mainly by the sinuous elytral keels.

Specimen examined No specimen examined.

Host-fungi Unknown.

Distribution Western Indian Ocean, Seychelles (Fig. 158).

Tropicis sexcarinatus (Waterhouse, 1876) (Figs 8, 12, 149–151, 153, 154B, 155B, 157–158)

Diagnosis and comments It differs from the other Tropicis by the six longitudinal keels at elytra, most conspicuous at the apical half, and the coarse pronotal punctation with coalescent punctures. It most resembles Trop. flexicarinatus, but do not co-occur with it (see Fig. 158). The species is variable morphologically and may be a complex of close related species. Lesne (1917) reported Cis sexcarinatus Waterhouse from Mauritius, La Reunión, Madagascar and Sey- chelles, before Scott (1926) erected the genus Tropicis based on this species and other two species from Sey- chelles. The specimens of Lesne (1917) from La Reunión were collected in Microporus affinis (cited as Polyporus microloma), being the unique record of host-fungus for Trop. sexcarinatus. The record of the species from Seychelles should be considered with care: it may refer to Trop. flexicarinatus, a morphologically similar species described by Scott (1926). The record from Madagascar may be correct, as that is very close to Mauritius and La Reunión, but it should be checked.

44 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE Specimens examined Mauritius: Total of 40 specimens (5 LAPC; 35 MHNG). 12 specimens labeled /Ile Maurice Y. Gomy [printed] Macabe Forest 13.i.70 tronc mort [handwritten]/; 7 specimens /Ile Maurice Y. Gomy [printed] Macabe Forest 22.i.66 Champignon [handwritten]/; 12 specimens labeled /Ile Maurice Y. Gomy [printed] Maré Lonque 19.i.71 tamisage tronc [handwritten]/; 9 specimens /Ile Maurice Y. Gomy [printed] Macabe For- est 19.i.71 tamisage tronc [handwritten]/. Rodrigues: 1 specimen (MHNG) labeled /Ile Rodrigue Y. Gomy [printed] Mont Limon 396m-8.5.72 tamisage dans hampe d’Alôes [handwritten]/. Réunion: Total of 48 specimens (5 LAPC; 43 MHNG). 9 specimens labeled /La Réunion Y. Gomy [printed] Cilaos Grand Matarum tamisage i.xi.69 [handwritten]/; 12 specimens /La Réunion Y. Gomy [printed] Bras de la Plaine 22.8.65 dans champignon [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Roche Ecrite 8.10.65 – 2200m au fauchoir [handwritten]/; 2 specimens /Cilaos Grand Matarum 6.8.67 dans champignon Y. Gomy [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Hell Bourq 6.11.65 s. ecorces [handwritten]/; 1 specimen / La Réunion Y. Gomy [printed] Cirque de Salazie Piton Marmite 1800m 14.i.72 tamisage tronc pourri [handwritten]/; 2 specimens /La Réunion J. David [printed] Foret Bebour aout 1973 [handwritten]/; 3 specimens /La Réunion Y. Gomy [printed] La Petite Plaine 20.3.66 champignon arborie [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] La Petite Plaine Pl. des Palmistes 25.v.69-1300m [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Ste Philippe-Alt.100m 15.2.71 tamisage souche [handwritten]; 3 specimens /La Réunion Y. Gomy [printed] Plaine des Chicots 1730m-10.10.66 Champignon [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Cilaos Grand Matarum 7.xi.69 tamisage [handwritten]/; 5 specimens /La Réunion Y. Gomy [printed] Pl. des Chicots Gite 1850m-19.4.73 tamisage terreau [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Pl. des Chicots Gite 21.4.73 1800m fauchoir [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Pl. des Chicots 21.4.73-1800m fauchage [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Pl. des Chicots Gite 1800m-21.4.73 s. calumets mort [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Hts. de Sta. Rose 25.10.70 tamisage [handwritten]/; 1 specimen /La Réunion Y. Gomy [printed] Pl. des Makes 15.xi.70 tamisage troncset humus [handwritten]/.

Host-fungi Microporus affinis (Polyporaceae), based in a record by Lesne (1917; see comments above).

Distribution Mascarene Islands (La Réunion, Mauritius and Rodrigues; Fig. 158) and probably Madagascar (Fig. 158, question mark).

Discussion

Morphological remarks Several external morphological characters of ciids, such as head and pronotal sculpture, pattern of colora- tion, pilosity, rugosity, and punctation may vary in a genus or even between closely related species. However, there are some morphological features that are shared by most of the species in each major group of Ciidae. For instance, Ceracis and Xylographus are usually glabrous, Orophiini usually has weak or no secondary sexual characters and spinose outer apical edge of protibiae, Ciini usually has broad prosternal process, and so on. There are exceptions for all these cases: some Xylographus are pilose; some male Orophiini (several Octotem- nus and Xylographus species) have modified mandibles; some genera currently in Ciini have laminate prosternal process; and the protibial spines of Paratrichapus (Orophiini) are restricted to their apices. All these characters in combination are currently being used to define genera and suprageneric taxa of Ciidae, but it is advisable to search for other robust characters to better delimit these taxa. Good characters may be found in

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 45 structures such as mouthparts, male pregenital segments, and female terminalia. However, these structures are not described for most ciid taxa. The next advances in the higher classification of Ciidae depend on the adequate comparison of these and other structures between its known genera and major taxa. The morphology of some characters, and its importance to the classification of the family, is briefly discussed below. The small cuticular protuberances on the dorsal surface of several Xylographellini species (see Figs 82– 84, small arrows) may be a character unique to tribe. These prominences have been called here “cuticular globules”, and they were more easily seen in some Scolytocis species. Each cuticular globule is a small protu- berance usually bearing a seta at its apex. Such globules may be present on the surface of the head and pronotum. However, they are more frequent on the elytra, where they are usually arranged in longitudinal rows on the apical declivity. These globules were also observed in species of the other three genera, but they were somewhat concealed by the strongly sculptured elytra of the species. In Tropicis and in Xyl. speciosa sp. nov. each row of cuticular globules is exactly on each longitudinal elytral keel. Indeed, these elytral keels may prove to be the alignment of several cuticular globules in a way that forms longitudinal lines in high relief. The antennae of Ciidae are peculiar due to the presence of “sensillifers” (group of modified and organized sensillae) in all the antennomeres of the club. These structures are absent only in Sphindocis denticollis Fall and in the genus Nipponapterocis Miyatake. When present, they are usually observed in number of four in each antennomere of the club. Scott (1926) was probably the first to observe more than four sensillifers (“sen- sory pores” in his words) in each apical antennomere of Tropicis, citing it as a character of the genus. Blair (1928) wrote that the antennae of Scolytocis resemble those of Tropicis, but he did not mentioned the sensillifers. Such variation in the number of antennal sensillifers was not used again in Ciidae taxonomy until Kawanabe and Miyatake (1996) cited it as a diagnostic character of Xylographellini. Besides the number, the position and configuration of the sensillifers may be of great taxonomic value (Lopes-Andrade & Zacaro 2002, 2003). It has already been speculated that these sensillifers may be hygroreceptors (Lawrence 1973) or even chemoreceptors (Lopes-Andrade & Zacaro 2002). Lawrence (1973) also proposed that these sensillifers would be homologous with the hygroreceptor organs of Tribolium, which were first studied by Roth and Wil- lis (1951). However, it should be noted here that the function of insect sensillae is hardly identifiable by their external appearance (Altner & Loftus 1985), and thus the function of the antennal sensillifers of ciids remains unknown. The mouthparts have not been adequately described for most ciid genera, but these structures certainly have characters of great taxonomic and phylogenetic importance. For instance, conspicuous morphological variations between some genera were already found in the prementum and apical labial palpomeres (e.g. Kawanabe & Miyatake 1996), galea (e.g. Kawanabe 2002), lacinia and basal labial palpomeres (e.g. Lopes- Andrade 2007b). The distinct configurations of the prementum and apical labial palpomeres of Xyl. punctata was used in the original delimitation of Xylographellini (Kawanabe & Miyatake 1996). Here, I have shown that their basic morphology is stable throughout the tribe, and can continue to be used as diagnostic characters to separate Xylographellini from the other major groups of ciids. Secondary sexual characters have already been observed in male mandibles of some Orophiini. Several male Octotemnus have the mandibles developed and projected forward, while the left mandible of some male Xylographus have a prominent upright tooth (Mellié 1849; Scott 1926). The absence of conspicuous secondary sexual characters in male head, pronotum and protibiae of Xylographellini is worth mentioning. Many species of the other major groups of Ciidae have tubercles, teeth, horns or other modifications on male head and/or pronotum, and in some species each protibia of male has a more produced outer apical angle. Even after careful observations and comparisons, I could not distinguish male and female of most Xylographellini prior to dissection. The similarity between male and female is more evident in the subtribe Xylographellina (Xylographella and Scolytocis), in which males are devoid of the abdominal sex patch. In Syncosmetina (Syncosmetus and Tropicis), the males usually have a small sex patch on the first abdominal ventrite. This structure is probably an external opening of a gland, and may be involved

46 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE in the secretion of pheromone (Faustini & Halstead 1982), probably sexual pheromone (Lopes-Andrade et al. 2003b). The secretion of aggregation pheromone was not observed in male abdominal sex patch of Xylogra- phus contractus Mellié (Lopes-Andrade et al. 2003b), but it remains a possibility for the other ciid species. The loss of the male sex patch in Xylographellina would be related to changes in the sexual behaviour and/or in the dynamic of fungi occupation and exploitation. The shape of the prosternal process is a useful character in Ciidae taxonomy, but should be considered with care. The genera previously included in Orophiini (Octotemnus, Paratrichapus, Xylographus, Ropal- odontus, Hyalocis) and Xylographellini (Scolytocis and Xylographella) are all characterized by a laminate prosternal process. But there are exceptions. For instance, Octotemnus omogensis Miyatake has a slightly broader prosternal process, but there is no doubt that it is a true Octotemnus as it fits the combination of the other diagnostic character of the genus. On the other hand, a laminate prosternal process occurs in genera not included in, and probably not close related to, Orophiini or Xylographellini. The latter is the case of Acantho- cis, Falsocis and Porculus, which possess all the characters of a true Ciini, but have laminate prosternal process and subconical and projected procoxae (Lopes-Andrade 2007b). Therefore, the configuration of the prosternal process and procoxae alone are weak arguments against the inclusion of Syncosmetus and Tropicis in Xylographellini. Other features are more robust and are unique to the genera here included in the tribe: the singular compact antennal club with their outnumbered sensillifers; prementum with a longitudinal sulcus at its external face; labial palpi inserted at the middle of prementum; and the Y-shaped ninth segment of the male abdomen (pregenital segment). Other characters are shared by Xylographellini and the Orophiini genera, such as the simplified female terminalia (Lopes-Andrade 2007a, b). Indeed, the morphology of the female terminalia deserves special atten- tion, and it may provide cues to solve several incongruities in the classification of the family. The basic configuration of the female terminalia observed in Xylographellini and Orophiini was already observed in several genera currently included in Ciini (for instance, Ceracis and Phellinocis), showing that the monophyly of Ciini is hardly sustainable (Lopes-Andrade 2007a, b). In the other hand, the female terminalia of the “true” Ciini genera (as Cis, Ennearthron, and Falsocis; Lopes-Andrade 2007a, b) all have the basic configuration found in many tenebrionoid families (Tanner 1927; Tschinkel & Doyen 1980; Banaszkiewicz 2006).

Host-use The pin label data of the specimens examined allowed the recognition of five host fungi genera of Scoly- tocis: Earliella and Fomes (Polyporaceae), Ganoderma (Ganodermataceae), Phellinus (Hymenochaetaceae) and Rigidoporus (Meripilaceae). Ganoderma, Rigidoporus, Fomes and Earliella are host fungi grouped under the Ganoderma host-use group (sensu Orledge & Reynolds 2005). Four Scolytocis species were found in Phellinus fungi: Scol. danielssoni sp. nov.; Scol. indecisus sp. nov., Scol. panamensis sp. nov.; and Scol. malayanus sp. nov. However, the latter three species were also collected in fungi of the Ganoderma host-use group. Orledge & Reynolds (2005) placed other Xylographellini species (Syn. reticulatus and Xyl. punctata) in the Ganoderma host-use group. Therefore, the use of fungi of the Phellinus host-use group by Xylographellini species may be incidental or alternative. The unique record of host-use for Tropicis (Lesne 1917) should be checked.

Geographic distribution Most of the species described here were collected in one or few localities. Some are even known from a single field collection. However, it cannot be said that they are rare species, since no attempt or little effort was made to recollect ciids in most of the localities. That is the case for the Indo-Malayan, Australasian and Oceanic species described here. In the regions where ciids were more frequently collected, the Xylographell- ini species are known from several field collections and localities. That applies to the Afrotropical, Japanese and Neotropical species. Considering the material I have examined, Tropicis is quite common in the Mas-

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 47 carene Islands. Among the Japanese species, both Syncosmetus species seem to be less frequently found, a pattern expected in the case of apterous forms. Most Neotropical Xylographellini were collected more than once, excepting Scol. cariborum sp. nov. and Scol. difficillimus sp. nov. However, the latter is very similar to Scol. bouchardi sp. nov. and may prove to be a synonym in the future. Some Neotropical species, such as Scol. bouchardi sp. nov., Scol. lawrencei sp. nov. and the three Brazilian Scolytocis are quite common and have wide distributions. The Brazilian species, for which I have the most detailed field data, are frequently collected. However, they are known only from forest remnants or conservation units. None of them were collected in open areas, urban areas or Cerrado vegetation (Brazilian Savana). Indeed, there seem to be three separate groups of ciids: one restricted to open areas (urban areas or other anthropized environments, and natural open vegetations); the other occurring only in forests; and a group of species which are able to occupy both enviornments. The Xylographellini species seem to better fit the second group. In Brazil, several ciids are known to occupy only open areas, as Cis gumiercostai Almeida & Lopes-Andrade and Ennearthron victori Lopes-Andrade & Zacaro (Almeida & Lopes-Andrade 2004; Lopes-Andrade 2008). Others are known only from forests, such as Falsocis brasiliensis Lopes-Andrade and the Brazilian Scolytocis (Gumier-Costa 2004; Lopes-Andrade 2007b). However, the ciids from forest are much more diversified (Gumier-Costa 2004; Lopes-Andrade pers. obs.), and they are certainly the most threatened (Lopes-Andrade 2007b). There are two major taxonomic groups in the tribe Xylographellini, formally treated here as separate subtribes (Fig. 159): (i) the Xylographellina, occurring in the Southeastern Asia (five species, one at Malaysia and four at the Philippines), Japan (one species), New Zealand (one species), Polynesia (Samoa, one species), Melanesia (Fiji, one species) and throughout the Neotropical Region (twelve species); (ii) the Syncosmetina subtr. nov., with two species from Japan (genus Syncosmetus) and four species from the Western Indian Ocean (in the Seychelles and Mascarene Islands; genus Tropicis). Therefore, both subtribes are distributed through distinct areas and overlap only on Japan (Figs 21, 122, 136). Xylographellini does not occur in the Nearctic Region, Europe and northern Asia, all with well-known Ciidae faunas. The tribe is known neither from the Andean Region nor the South American transition zone (sensu Morrone 2006), but both have barely studied Ciidae faunas. Both subtribes of Xylographellini occur in Japan, but they are represented by only three species (one Xylographellina and two Syncosmetina subtrib. nov.). The occurrence of the tribe throughout the Indo-Malayan Region is plausible, and already confirmed in Malaysia and the Philippines. The case of the Afrotropical Region is also of great interest. The islands where Tropicis (Syncosmetina subtr. nov.) occur are all in the western Indian Ocean (Seychelles and the Mascarene Islands), and most Tropicis species seem to be quite common there. Other ciids described from these islands (or close related species) are already known from the African continent (Ferrer 1997; Lopes-Andrade pers. obs.), showing that there may be great affinities between the continental African and the western Indian Ocean ciid faunas. Indeed, there may be a true Afrotropical Ciidae fauna, with many species shared by the continent and part of the surrounding islands. It is quite expected that Syncosmetina subtr. nov. occurs on the continental sub-Saharan Africa. However, I have not found any species resembling it even after examining a good number of ciids (more than 5,000 specimens from the KMMA) collected in central Africa and Madagascar during the 20th century. The subtribe Xylographellina is most diversified in the Neotropical Region, being well represented by the genus Scolytocis. The Neotropical fauna of Scolytocis has three basic components: (i) a Northern continental fauna, comprising species of three morphological species-groups (bouchardi, danielssoni and lawrencei groups); (ii) an insular fauna (Greater and Lesser Antilles), with one species-group (lawrencei group); (iii) a Southern fauna, including only one morphological species-group (fritzplaumanni group). The continental Northern and Southern Scolytocis faunas seem to be well separated by the South American transition zone (sensu Morrone 2006). The species from Antilles, although included in the lawrencei species-group, have the male genitalia with features not yet observed in any other Scolytocis. Based on the data provided above, the following biogeographical hypotheses can be proposed: (i) a Gond-

48 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE wanan origin of Xylographellini; (ii) an origin of the subtribe Xylographellina on western Gondwana; (iii) an origin of Syncosmetina subtr. nov. on eastern Gondwana; (iv) a subsequent expansion of the South-American Xylographellina fauna to the Australasian, Oceanic and Indo-Malayan Regions; (v) isolation of the western Indian-Ocean and the Indo-Malayan faunas of Syncosmetina subtr. nov., seen today as two distinct genera (Tropicis and Syncosmetus, respectively). However, without a robust phylogenetic scenario for Ciidae, all these propositions remain speculative. Alternative explanations would be recent events of oceanic dispersion, and/or higher persistence of species (lower extinction rate) in islands than in continental lands. Indeed, molec- ular dating analyses have supported oceanic dispersal over tectonic vicariance for many taxa (De Queiroz 2005). However, it should be emphasized here that vicariance and dispersal hypotheses are not mutually exclusive, as they may explain the geographic expansion of Xylographellini in different evolutive times. The study of the insular Xylographellini fauna will certainly provide key-evidences on the evolution and biogeography of the tribe.

Acknowledgements

I'm very grateful to John F. Lawrence for all the information and specimens provided, and also for indicating some museum collections containing Xylographellini specimens. This work would not be possible without his help. I’m very grateful to Rafal Ruta and an anonymous referee for reviewing the manuscript, and also for the Coleoptera editor Darren Pollock for all the assistance. Adilson A. Zacaro, Carlos.F. Sperber, Fernando Z. Vaz-de-Mello, Marcos G. Lhano, Og F. F. de Souza and Rodrigo F. Krüger carefully read a preliminary version of this paper and made valuable corrections and suggestions. Fabiano Gumier-Costa, Fernando Vaz-de- Mello, Karina Furieri, Júlio Louzada, Letícia Graf and Paschoal Grossi collected and provided me with specimens of the Brazilian Scolytocis. Fernando Vaz-de-Mello searched for Ciidae in the collections of CMNC and CNCI, and kindly translated the abstract to Spanish. Rafal Ruta provided me with specimens of Xylographella punctata and Syncosmetus reticulatus. Makoto Kawanabe kindly sent me the original article on Xylographell- ini and other of his articles on Ciidae. Fernando Vaz-de-Mello, Glenda Orledge, Klaus-Dieter Klass and Mar- cos Lhano sent me hard copies or PDF files of several articles. My wife Teresinha Nolasco performed the automontage of most photographs used in the plates of dorsal, ventral and lateral views of the species. Equip- ment facilities were provided by “Laboratório de Orthopterologia” and “Laboratório de Biologia Estrutural, Histofisiologia Reprodutiva e Disgestiva” (DBG/UFV), and also by Alexandre Nolasco. CLA was granted by CNPq for his doctorate degree (processo número 140463/2004–6) and also for his post-doctoral research (PDJ, processo número 151800/2007-3). John F. Lawrence first recognized some of the new Scolytocis, Trop- icis and Xylographella described here, and organized them for the loans provided by ANIC, FMNH and MHNG. However, all conclusions provided here are of my full responsibility. Adam Slipinski (ANIC), Giulio Cuccodoro (MHNG), Margaret Thayer and James Boone (FMNH), Robert Anderson (CMNC), Pat Bouchard (CNCI) and Roy Danielsson (MZLU) arranged and/or authorized loans from their institutions. SEM photographs were made at NAP/MEPA (ESALQ/USP) with the assistance of Dr. Elliot W. Kitajima.

References

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50 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE Miyatake, M. (1954) Studies on the Japanese Ciidae, I (Coleoptera). Scientific Reports of the Matsuyama Agricultural College, 14, 40–67, pls. 1–11. Miyatake, M. (1985) Ciidae, pp. 278–285. IN: Y. Kurasawa, S. Hisamatsu and H. Sasaji (eds.), The Coleoptera of Japan in Color, 3. Hoikusha, Osaka (in Japanese). Morrone, J.J. (2006) Biogeographic areas and transition zones of Latin America and the Caribbean Islands based on pan- biogeographic and cladistic analyses of the entomofauna. Annual Review of Entomology, 51, 467–494. Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D’Amico, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wetten- gel, W.W., Hedao, P. & Kassem, K.R. (2001) Terrestrial ecoregions of the world: a new map of life on Earth. Bio- science, 51, 933–938. Orledge, G.M. & Reynolds, S.E. (2005) Fungivore host-use groups from cluster analysis: patterns of utilization of fungal fruiting bodies by ciid beetles. Ecological Entomology, 30, 620–641. Roth, L.M. & Willis, E.R. (1951) Hygroreceptors in adults of Tribolium (Coleoptera, Tenebrionidae). Journal of Experi- mental Zoology, 116, 527–570. Scott, H. (1926) Coleoptera, Ciidae. Reports of the Percy Sladen Trust Expedition to the Indian Ocean in 1905. Vol. 8, No. 1. Transactions of the Linnean Society of London, Ser. 2. Zoology, 19(1), 1–41. Sharp, D. (1891) Descriptions of two new species and a new genus of Japanese Coleoptera. Entomologist, 24, 6–7. Tanner, V.M. (1927) A preliminary study of the genitalia of female Coleoptera. Transactions of the American Entomo- logical Society, 53, 5–50, pls 2–15. Tschinkel, W.R. & Doyen, J.T. (1980) Comparative anatomy of the defensive glands, ovipositors and female genital tubes of tenebrionid beetles (Coleoptera). International Journal of Insect Morphology and Embryology, 9, 321–368. Waterhouse, C. O. (1876) New species of Coleoptera from the island of Rodriguez, collected by the naturalists accompa- nying the Transit of Venus expedition. Annals and Magazine of Natural History, 18(4), 105–121.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 51 FIGURES 1–4. Scolytocis paschoali sp. nov. 1. Part of the head, frontal view showing the eye. 2. Antennal club, showing the sensillifers (arrows). Note that the club is compact, with antennomeres not freely articulated. 3. Head, ventral view showing the mouthparts. Note the longitudinal sulcus of prementum (arrow). 4. Ventral view showing the metaventrite.

52 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 5–8. Prementum (pm) with labial palpi (l.p). Note that the apical labial palpomeres are enlarged (small arrows), and that the apex of prementum is acute in Xylographella and Scolytocis, and rounded in Syncosmetus and Trop- icis (see big arrows). 5. Xyl. punctata Miyatake, lateral view. 6. Scol. paschoali sp. nov. 7. Sync. reticulatus Miyatake. 8. Trop. sexcarinatus (Waterhouse).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 53 FIGURES 9–12. Female terminalia, showing the gonostyli (gs), gonocoxites (gc), baculi of basal gonocoxites (small arrows), paraprocts (pp), baculi of paraprocts (b.pp), proctiger (pt), and spiculum ventrale (sv). The baculi of proctiger are barely sclerotized, so they are not indicated. 9. Xylographella punctata Miyatake. 10. Scolytocis furieriae sp. nov. 11. Syncosmetus japonicus Sharp. 12. Tropicis sexcarinatus (Waterhouse).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 57 FIGURES 21–22. Xylographella Miyatake. 21. Antennae of Xyl. punctata Miyatake (A) and Xyl. speciosa sp. nov. (B). Note that the first antennomere is not shown in the latter (asterisk). 22. Metatibiae of Xyl. punctata Miyatake (A) and Xyl. speciosa sp. nov. (B).

58 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 23–25. Xylographella Miyatake. 23. Tegmen (teg) and penis (pen) of Xyl. punctata. 24. Y-Shaped ninth segment (broken, see asterisks) of Xyl. punctata. 25. Male genitalia of Xyl. speciosa sp. nov. showing the Y-shaped ninth- segment (ix), tegmen (teg) and penis (pen).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 59 FIGURE 26. Distribution map of Xylographella Miyatake, showing the known distribution of Xyl. punctata Miyatake (triangles) in Japan and Xyl. speciosa sp. nov. (stars) in the Philippines.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 69 FIGURES 54–56. Hindwings of Northern Neotropical Scolytocis Blair. 54. Scol. bouchardi sp. nov. 55. Scol. cariborum sp. nov. 56. Scol. danielssoni sp. nov.

70 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 57–59. Hindwings of Northern Neotropical Scolytocis Blair. 57. Scol. difficillimus sp. nov. 58. Scol. howdeni sp. nov. 59. Scol. indecisus sp. nov.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 71 FIGURES 60–62. Hindwings of Northern Neotropical Scolytocis Blair. 60. Scol. kiskeyensis sp. nov. 61. Scol. lawrencei sp. nov. 62. Scol. panamensis sp. nov.

72 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 63–65. Antennae of the Northern Neotropical Scolytocis Blair. 63. Scol. bouchardi sp. nov. (A), Scol. cariborum sp. nov. (B), Scol. danielssoni sp. nov. (C). 64. Scol. difficillimus sp. nov. (A), Scol. howdeni sp. nov. (B), Scol. indecisus sp. nov. (C). 65. Scol. kiskeyensis sp. nov. (A), Scol. lawrencei sp. nov. (B), Scol. panamensis sp. nov. (C). First antennomere not shown in Scol. danielssoni sp. nov. (63C, asterisk).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 73 FIGURES 66–68. Metatibiae of the Northern Neotropical Scolytocis Blair. 66. Scol. bouchardi sp. nov. (A), Scol. cariborum sp. nov. (B), Scol. danielssoni sp. nov. (C). 67. Scol. difficillimus sp. nov. (A), Scol. howdeni sp. nov. (B), Scol. indecisus sp. nov. (C). 68. Scol. kiskeyensis sp. nov. (A), Scol. lawrencei sp. nov. (B), Scol. panamensis sp. nov. (C).

74 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 69–71. Male genitalia of Scolytocis Blair showing the Y-shaped ninth segment (ix), basal piece (b.p), tegmen (teg) and penis (pen). 69. Scol. bouchardi sp. nov. 70. Scol. cariborum sp. nov. (ninth segment broken; see asterisks). 71. Scol. difficillimus sp. nov.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 75 FIGURES 72–73. Male genitalia of Scolytocis Blair showing the Y-shaped ninth segment (ix), tegmen (teg) and penis (pen). 72. Scol. howdeni sp. nov. 73. Scol. indecisus sp. nov. The dashed line indicates the membranous basal portion of tegmen. Note that the apical portion of tegmen is extremely sclerotized.

76 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 74–76. Male genitalia of Scolytocis kiskeyensis sp. nov. showing the Y-shaped ninth segment (ix), tegmen (teg) and penis (pen). 74. Tegmen and penis of specimen from the Dominican Republic (type locality). 75. Tegmen and penis of specimen from Puerto Rico. 76. Ninth segment (pregenital segment) of specimen from Puerto Rico.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 77 FIGURES 77–79. Male genitalia of Scolytocis Blair showing the Y-shaped ninth segment (ix), basal piece (b.p), tegmen (teg) and penis (pen). 77. Scol. lawrencei sp. nov., specimen from Costa Rica. The dashed line indicates the membranous basal portion of tegmen. 78. Scol. lawrencei sp. nov., specimen from Panama. 79. Scol. panamensis sp. nov.

78 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURE 80. Distribution map of the continental Northern Neotropical species of Scolytocis Blair, showing the known distributions of Scol. bouchardi sp. nov. (stars), Scol. danielssoni sp. nov. (double circles), Scol. difficillimus sp. nov. (fulfilled square), Scol. howdeni sp. nov. (open squares), Scol. indecisus sp. nov. (plus symbols), Scol. lawrencei sp. nov. (fulfilled triangles), Scol. panamensis sp. nov. (open circle).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 79 FIGURE 81. Distribution map of the insular Northern Neotropical species of Scolytocis Blair, showing the known distributions of Scol. cariborum sp. nov. (plus symbol) and Scolytocis kiskeyensis sp. nov. (stars).

80 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 82–84. Scolytocis fritzplaumanni sp. nov, holotype. Note the presence of small cuticular globules (protuberances) on the head, pronotum and elytra (small arrows). 82. Dorsal view. 83. Lateral view. 84. Ventral view.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 83 FIGURES 91–93. Hindwings of the Southern Neotropical Scolytocis Blair. 91. Scol. fritzplaumanni sp. nov. 92. Scol. furieriae sp. nov. 93. Scol. paschoali sp. nov.

84 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 94–95. Southern Neotropical Scolytocis Blair. 94. Antennae of Scol. fritzplaumanni sp. nov. (A), Scol. furieriae sp. nov. (B) and Scol. paschoali sp. nov. (C). 95. Metatibiae of Scol. fritzplaumanni sp. nov. (A), Scol. furieriae sp. nov. (B) and Scol. paschoali sp. nov. (C). First antennomere not shown in Scol. furieriae sp. nov. (94B, asterisk).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 85 FIGURES 96–97. Male genitalia of Scolytocis fritzplaumanni sp. nov. showing the Y-shaped ninth segment (ix), basal piece (b.p), tegmen (teg) and penis (pen). 96. Specimen from Nova Teutônia, Santa Catarina, Brazil (type locality). 97. Specimen from Guaratuba, Rio Grande do Sul, Brazil.

86 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 98–100. Male genitalia of Scolytocis Blair showing the Y-shaped ninth segment (ix), basal piece (b.p), tegmen (teg) and penis (pen). 98. Scol. furieriae sp. nov. from Jussari, Bahia, Brazil (type locality). 99–100. Scol. paschoali sp. nov., specimens from Macaé de Cima, Rio de Janeiro, Brazil (type locality). 99. Dorsal-ventral view. 100. Lateral view.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 87 FIGURE 101. Map showing the distribution of the Southern Neotropical species of Scolytocis. Scol. fritzplaumanni sp. nov. (circles), Scol. furieriae sp. nov. (triangles), Scol. paschoali sp. nov. (squares).

94 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 120–122. Hindwings of the new Indo-Malayan (in part) and Australasian Scolytocis Blair. 120. Scol. malayanus sp. nov. 121. Scol. novaezelandiae sp. nov. 122. Scol. philippinensis sp. nov.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 95 FIGURES 123–125. Hindwings of the new Indo-Malayan (in part) and Oceanic Scolytocis Blair. 123. Scol. thayerae sp. nov. 124. Scol. werneri sp. nov. 125. Scol. zimmermani sp. nov.

96 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 126–127. Antennae of the Indo-Malayan, Australasian and Oceanic species of Scolytocis Blair. 126. Scol. malayanus sp. nov. (A), Scol. novaezelandiae sp. nov. (B) and Scol. philippinensis sp. nov. (C). 127. Scol. thayerae sp. nov. (A), Scol. werneri sp. nov. (B) and Scol. zimmermani sp. nov. (C).

FIGURES 128–129. Metatibiae of the Indo-Malayan, Australasian and Oceanic species of Scolytocis Blair. 128. Scol. malayanus sp. nov. (A), Scol. novaezelandiae sp. nov. (B) and Scol. philippinensis sp. nov. (C). 129. Scol. thayerae sp. nov. (A), Scol. werneri sp. nov. (B) and Scol. zimmermani sp. nov. (C).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 97 FIGURES 130–133. Male genitalia of the Indo-Malayan species of Scolytocis Blair showing the Y-shaped ninth segment (ix), tegmen (teg) and penis (pen). 130. Scol. malayanus sp. nov. 131. Scol. philippinensis sp. nov. 132. Scol. thayerae sp. nov. 133. Scol. werneri sp. nov.

98 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURE 134. Distributional map of the Indo-Malayan species of Scolytocis Blair, showing the distribution of Scol. malayanus sp. nov. (triangles), Scol. philippinensis sp. nov. (double circle), Scol. thayerae sp. nov. (letters “X”) and Scol. werneri sp. nov. (plus symbol).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 99 FIGURE 135. Distributional map of the Australasian and Oceanic species of Scolytocis Blair, showing the distribution of Scol. novaezelandiae sp. nov. (triangles), Scol. samoensis Blair (plus symbol) and Scol. zimmermani sp. nov. (square).

102 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURES 142–144. Syncosmetus Sharp. 142. Antennae of Sync. japonicus Sharp (A) and Sync. reticulatus Miyatake (B). Note that the first antennomere is not shown in the latter (asterisk). 143. Metatibiae of Sync. japonicus Sharp (A) and Sync. reticulatus Miyatake (B). Note the distribution of small spines in the outer margin of each tibia (small arrows). 144. Male genitalia of Sync. japonicus Sharp showing the Y-shaped ninth segment (ix), basal piece (b.p), tegmen (teg) and penis (pen).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 103 FIGURE 145. Distribution map of Syncosmetus Sharp in Japan, showing the known distributions of Sync. japonicus Sharp (triangles) and Sync. reticulatus Miyatake (hexagon).

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 107 FIGURES 154–157. Tropicis Scott. 154. Antennae of Trop. cuccodoroi sp. nov. (A) and Trop. sexcarinatus (B). Note that the first antennomere is not shown in the former (asterisk). 155. Metatibiae of Trop. cuccodoroi sp. nov. (A) and Trop. sexcarinatus (B). Note the distribution of small spines in the outer margin of each tibia (small arrows). 156–157. Male genitalia, showing the ninth segment (ix-seg), tegmen (teg) and penis (pen). 156. Trop. cuccodoroi sp. nov. 157. Trop. sexcarinatus (Waterhouse) showing the tegmen and penis (A) and the ninth segment (B).

108 · Zootaxa 1832 © 2008 Magnolia Press LOPES-ANDRADE FIGURE 158. Distribution map of Tropicis Scott in the Western Indian Ocean, showing the known distributions of Trop- icis brevicarinatus Scott (double circles), Trop. cuccodoroi sp. nov. (open stars), Trop. flexicarinatus Scott (“X” symbols) and Trop. sexcarinatus (Waterhouse) (plus symbol). The question mark indicates a doubtful record of Trop. sexcarinatus from Madagascar.

THE TRIBE XYLOGRAPHELLINI Zootaxa 1832 © 2008 Magnolia Press · 109 FIGURE 159. Distribution map of Xylographellini, showing the known distributions of the subtribes Xylographellina Kawanabe & Miyatake (stars) and Syncosmetina subtrib. nov. (triangles).

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