A monograph of the Xyleborini (Coleoptera, Curculionidae, Scolytinae) of the Indochinese Peninsula (except Malaysia) and China

Sarah M. Smith; Roger A. Beaver; Anthony I. Cognato

doi:10.3897/zookeys.983.52630

Monograph

A monograph of the Xyleborini (Coleoptera, Curculionidae, Scolytinae) of the Indochinese Peninsula (except Malaysia) and China

Sarah M. Smith^‡, Roger A. Beaver^§, Anthony I. Cognato^‡

‡ Michigan State University, East Lansing, United States of America

§ Unaffiliated, Chiangmai, Thailand

Corresponding author: Sarah M. Smith ( camptocerus@gmail.com )

Academic editor: Miguel Alonso-Zarazaga

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Smith SM, Beaver RA, Cognato AI (2020) A monograph of the Xyleborini (Coleoptera, Curculionidae, Scolytinae) of the Indochinese Peninsula (except Malaysia) and China. ZooKeys 983: 1-442. https://doi.org/10.3897/zookeys.983.52630

ZooBank: urn:lsid:zoobank.org:pub:7DED4CE2-934C-4539-945F-758930C927F9

Abstract

The Southeast Asian xyleborine ambrosia beetle fauna is reviewed for the first time. Thirty-four genera and 315 species are reviewed, illustrated, and keyed to genera and species. Sixty-three new species are described: Amasa cycloxyster sp. nov., Amasa galeoderma sp. nov., Amasa gibbosa sp. nov., Amasa lini sp. nov., Amasa tropidacron sp. nov., Amasa youlii sp. nov., Ambrosiophilus caliginestris sp. nov., Ambrosiophilus indicus sp. nov., Ambrosiophilus lannaensis sp. nov., Ambrosiophilus papilliferus sp. nov., Ambrosiophilus wantaneeae sp. nov., Anisandrus achaete sp. nov., Anisandrus auco sp. nov., Anisandrus auratipilus sp. nov., Anisandrus congruens sp. nov., Anisandrus cryphaloides sp. nov., Anisandrus feronia sp. nov., Anisandrus hera sp. nov., Anisandrus paragogus sp. nov., Anisandrus sinivali sp. nov., Anisandrus venustus sp. nov., Anisandrus xuannu sp. nov., Arixyleborus crassior sp. nov., Arixyleborus phiaoacensis sp. nov., Arixyleborus setosus sp. nov., Arixyleborus silvanus sp. nov., Arixyleborus sittichayai sp. nov., Arixyleborus titanus sp. nov., Coptodryas amydra sp. nov., Coptodryas carinata sp. nov., Coptodryas inornata sp. nov., Cyclorhipidion amasoides sp. nov., Cyclorhipidion amputatum sp. nov., Cyclorhipidion denticauda sp. nov., Cyclorhipidion muticum sp. nov., Cyclorhipidion obesulum sp. nov., Cyclorhipidion petrosum sp. nov., Cyclorhipidion truncaudinum sp. nov., Cyclorhipidion xeniolum sp. nov., Euwallacea geminus sp. nov., Euwallacea neptis sp. nov., Euwallacea subalpinus sp. nov., Euwallacea testudinatus sp. nov., Heteroborips fastigatus sp. nov., Heteroborips indicus sp. nov., Microperus latesalebrinus sp. nov., Microperus minax sp. nov., Microperus sagmatus sp. nov., Streptocranus petilus sp. nov., Truncaudum bullatum sp. nov., Xyleborinus cuneatus sp. nov., Xyleborinus disgregus sp. nov., Xyleborinus echinopterus sp. nov., Xyleborinus ephialtodes sp. nov., Xyleborinus huifenyinae sp. nov., Xyleborinus jianghuansuni sp. nov., Xyleborinus thaiphami sp. nov., Xyleborinus tritus sp. nov., Xyleborus opacus sp. nov., Xyleborus sunisae sp. nov., Xyleborus yunnanensis sp. nov., Xylosandrus bellinsulanus sp. nov., Xylosandrus spinifer sp. nov.. Thirteen new combinations are given: Ambrosiophilus consimilis (Eggers) comb. nov., Anisandrus carinensis (Eggers) comb. nov., Anisandrus cristatus (Hagedorn) comb. nov., Anisandrus klapperichi (Schedl) comb. nov., Anisandrus percristatus (Eggers) comb. nov., Arixyleborus resecans (Eggers) comb. nov., Cyclorhipidion armiger (Schedl) comb. nov., Debus quadrispinus (Motschulsky) comb. nov., Heteroborips tristis (Eggers) comb. nov., Leptoxyleborus machili (Niisima) comb. nov., Microperus cruralis (Schedl) comb. nov., Planiculus shiva (Maiti & Saha) comb. nov., Xylosandrus formosae (Wood) comb. nov. Twenty-four new synonyms are proposed: Ambrosiophilus osumiensis (Murayama, 1934) (= Xyleborus nodulosus Eggers, 1941 syn. nov.); Ambrosiophilus subnepotulus (Eggers, 1930) (= Xyleborus cristatuloides Schedl, 1971 syn. nov.); Ambrosiophilus sulcatus (Eggers, 1930) (= Xyleborus sinensis Eggers, 1941 syn. nov.; = Xyleborus sulcatulus Eggers, 1939 syn. nov.); Anisandrus hirtus (Hagedorn, 1904) (= Xyleborus hirtipes Schedl, 1969 syn. nov.); Cnestus protensus (Eggers, 1930) (= Cnestus rostratus Schedl, 1977 syn. nov.); Cyclorhipidion bodoanum (Reitter, 1913) (= Xyleborus misatoensis Nobuchi, 1981 syn. nov.); Cyclorhipidion distinguendum (Eggers, 1930) (= Xyleborus fukiensis Eggers, 1941 syn. nov.; = Xyleborus ganshoensis Murayama, 1952 syn. nov.); Cyclorhipidion inarmatum (Eggers, 1923) (= Xyleborus vagans Schedl, 1977 syn. nov.); Debus quadrispinus (Motschulsky, 1863) (= Xyleborus fallax Eichhoff, 1878 syn. nov.); Euwallacea gravelyi (Wichmann, 1914) (= Xyleborus barbatomorphus Schedl, 1951 syn. nov.); Euwallacea perbrevis (Schedl, 1951) (= Xyleborus molestulus Wood, 1975 syn. nov.; Euwallacea semirudis (Blandford, 1896) (= Xyleborus neohybridus Schedl, 1942 syn. nov.); Euwallacea sibsagaricus (Eggers, 1930) (= Xyleborus tonkinensis Schedl, 1934 syn. nov.); Euwallacea velatus (Sampson, 1913) (= Xyleborus rudis Eggers, 1930 syn. nov.); Microperus kadoyamaensis (Murayama, 1934) (= Xyleborus pubipennis Schedl, 1974 syn. nov.; =Xyleborus denseseriatus Eggers, 1941 syn. nov.); Stictodex dimidiatus (Eggers, 1927) (=Xyleborus dorsosulcatus Beeson, 1930 syn. nov.); Webbia trigintispinata Sampson, 1922 (= Webbia mucronatus Eggers, 1927 syn. nov.); Xyleborinus artestriatus (Eichhoff, 1878) (= Xyelborus angustior [sic] Eggers, 1925 syn. nov.; = Xyleborus undatus Schedl, 1974 syn. nov.); Xyleborinus exiguus (Walker, 1859) (= Xyleborus diversus Schedl, 1954 syn. nov.); Xyleborus muticus Blandford, 1894 (= Xyleborus conditus Schedl, 1971 syn. nov.; = Xyleborus lignographus Schedl, 1953 syn. nov.). Seven species are removed from synonymy and reinstated as valid species: Anisandrus cristatus (Hagedorn, 1908), Cyclorhipidion tenuigraphum (Schedl, 1953), Diuncus ciliatoformis (Schedl, 1953), Euwallacea gravelyi (Wichmann, 1914), Euwallacea semirudis (Blandford, 1896), Microperus fulvulus (Schedl, 1942), Xyleborinus subspinosus (Eggers, 1930).

Keywords

ambrosia beetles, biodiversity, new combinations, new species, new synonymy, Oriental region, Scolytidae, taxonomy

Introduction

Xyleborine ambrosia beetles (Curculionidae: Scolytinae) occur throughout the forested regions of the world with the highest diversity occurring in the tropical and subtropical regions (Hulcr et al. 2015). It is hypothesized that xyleborines originated in the Orient given the region’s high species and generic diversity (Hulcr et al. 2015; Cognato et al. 2018). Since their origin 20 million years ago, xyleborines have successfully dispersed across the world, sparking radiations of species wherever colonists landed (especially the Neotropics) (Jordal and Cognato 2012; Cognato et al. 2018). There are approximately 1200 species currently recognized and they comprise the largest scolytine tribe, representing approximately 20% of total diversity. However, this total diversity has yet to be fully realized with an estimated 25–75% awaiting discovery and description (Hulcr et al. 2015; Smith et al. 2017a) and approximately 30% in tropical Asia. The biology of these beetles makes them extremely well-suited for colonization (Jordal et al. 2001; Gohli et al. 2016). They have a strongly female-skewed haplodiploid mating system with extreme inbreeding (Kirkendall 1993; Kirkendall et al. 2015). Usually, females mate with a brother before leaving the natal gallery. If unmated, a female lays haploid eggs, which develop into males. The adult male, which is dwarfed and flightless, may mate with his mother who then produces diploid eggs which develop into females. These beetles also cultivate symbiotic fungal gardens within tunnels they bore into trees. The beetles have specialized body parts (mycangia) which fill with fungi and provide secure transport of the fungi to new habitats. Mycangia are invaginated pouches which occur in the head near the mandibles, pronotum/mesonotum, and in the elytral bases (Beaver 1989). The type of mycangium tends to be taxon specific and several fungal genera form specific symbiotic relationships with xyleborine genera (Beaver 1989; Hulcr and Cognato 2010; Hulcr and Stelinski 2017). Thus, upon arrival at a new location, even an unmated female provisioned with symbiotic fungi can produce a fungal garden and a family which can eventually grow into a population of beetles. This great colonizing potential has led to the accidental introduction through global trade of 31 and 12 species to North America and Europe, respectively (Kirkendall and Faccoli 2010; Garonna et al. 2012; Terekhova and Skrylnik 2012; Dodelin 2018; Rabaglia et al. 2019, 2020a). Most of these introduced species were native to SE Asia (Haack and Rabaglia 2013). In North America, three SE Asian species Euwallacea fornicatus (Eichhoff, 1868), E. kuroshio (Gomez & Hulcr, 2018), and Xyleborus glabratus Eichhoff, 1877, have caused major economic and ecological damage to trees in urban/suburban and natural areas (Eskalen et al. 2013; Boland 2016; Carillo et al. 2016; Hughes et al. 2017; Coleman et al. 2019).

Taxonomic knowledge of xyleborines is mostly limited to alpha-level taxonomy that began in earnest with the description of Xyleborus by Eichhoff (1864), and progressed with major contributions from Eichhoff, Blandford, Eggers, Schedl, Browne, Murayama, Nobuchi and Wood (Wood and Bright 1992). Given the unique aspects of xyleborine biology (as described above), morphological aberrations that occur within a single foundress can rapidly propagate among progeny which may ultimately grow to population levels. This intraspecific variation has historically been problematic and confounded the delineation of species limits. This has led to numerous subjective synonyms for many species, especially widespread taxa (e.g., Xyleborus affinis Eichhoff, 1868, X. perforans (Wollaston, 1857), Xyleborinus exiguus (Walker, 1859). Many species were described from short series or singletons which insufficiently assessed intraspecific variation (e.g., Euwallacea fornicatus complex). Single individuals of multiple species from a variety of locations often seemingly formed a continuous spectrum of variation which has led to their synonymization (Hulcr and Cognato 2013). Generic taxonomy began with the description of Eccoptopterus (Motschulsky, 1863), Xyleborus (Eichhoff, 1864), and Amasa (Lea, 1894) and by 1990, 24 genera had been described through the efforts of Blandford, Hagedorn, Hopkins, Reitter, and Sampson (Wood and Bright 1992). The 2000’s brought the use of molecular phylogenies to identify monophyletic groups and elucidate taxon limits (Gomez et al. 2018b; Cognato et al. 2019, 2020a; Smith et al. 2020). Currently, there are 42 recognized xyleborine genera with the likely recognition of additional genera given the extensive morphological variation observed in the polyphyletic Xyleborus (Cognato et al. 2020a). Comprehensive species reviews and identification keys are limited to generic level studies (e.g., Beaver and Hulcr 2008; Beaver 2010; Dole and Cognato 2010; Smith 2017; Beaver et al. 2019) and faunal reviews of geographic regions: North and Central America (Wood 1982), China (Yin et al. 1984), Europe (Pfeffer 1994), South America (Wood 2007), India (Maiti and Saha 2004), Papua New Guinea (Hulcr and Cognato 2010), Taiwan (Beaver and Liu 2010), Thailand (Beaver et al. 2014) and the West Indies (Bright 2019). These geographic reviews and monographs provide a necessary foundation for understanding the xyleborine fauna, but quickly become outdated as new species are found and taxonomic changes made. Nevertheless, the keys provide a gateway into identifying this economically important group of beetles. A comprehensive publication for SE Asia is conspicuously absent and lack of this resource has caused delays in identifying non-native species or mistaken identities (Smith and Cognato 2015; Smith et al. 2017b; Hoebeke et al. 2018).

Given that SE Asia species are intercepted at US and other ports every year and have proven pestiferous (Haack and Rabaglia 2013), a review and key for the xyleborine fauna of SE Asia is critically needed (Smith and Cognato 2015; Smith et al. 2017b; Rabaglia et al. 2019). In 2016, AIC was funded to create identification tools including DNA barcodes and a Lucid key of this fauna (Smith et al. 2019a; Cognato et al. 2020b). As indicated by the title, the geographic region of study is awkward; it focuses on the Indochinese Peninsula (Cambodia, Myanmar, Laos, Thailand, Vietnam) excluding Malaysia and insular SE Asian countries, and includes subalpine Himalayan areas (Northern India, Nepal, Bhutan), Bangladesh, China, and Taiwan. This was intentional in order to focus the study on the region of greatest potential for harboring future pests in non-native regions outside the equatorial tropical rain forest belt (McCullough et al. 2006; Haack and Rabaglia 2013). As a result of creating these identification tools, a review of the fauna was accomplished, which is detailed in this publication.

Materials and methods

Examined specimens came from our own collections, fieldwork and through loans from several institutions. All descriptions, keys and diagnoses are based on females as males are largely unknown, rarely encountered, and not often present without a female of the same species. Type material was examined by all authors. Specimens were assembled and examined from the following entomological collections by one or more authors:

BPBM Bernice P. Bishop Museum, Honolulu, USA;

CASC California Academy of Sciences, San Francisco, USA;

CSLC Ching-Shan Lin collection, Chang Hua, Taiwan;

FRI Forest Research Institute, Dehra Dun, India;

HNHM Hungarian Natural History Museum, Budapest, Hungary;

IRSNB Institut Royale des Sciences Naturelles, Brussels, Belgium;

IZAS Institute of Zoology, Chinese Academy of Sciences, Beijing, China;

FSCA Florida State Collection of Arthropods, Gainesville, USA;

LYLC Lan-Yu Liu collection, Yilan, Taiwan;

MCG Museo Civico di Storia Naturale “Giacomo Doria”, Genova, Italy;

MCZ Museum of Comparative Zoology, Cambridge, USA;

MFNB Museum für Naturkunde, Berlin, Germany;

MIZ Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw, Poland;

MNHN Muséum National d'Histoire Naturelle, Paris, France;

MNHP Museum of Natural History, Prague, Cechia;

MSUC Michigan State University Arthropod Research Collection, East Lansing, USA;

NHMB Natural History Museum, Basel, Switzerland;

NHMUK Natural History Museum, London, UK;

NHMW Naturhistorisches Museum Wien, Austria;

NIAES National Institute for Agro-Environmental Sciences, Tsukuba, Japan;

NKME Naturkunde Museum, Erfurt, Germany;

NMNH National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA;

OMNH Sam Noble Oklahoma Museum of Natural History, University of Oklahoma, Norman, USA;

PPST Plant Protection Station, Tokyo, Japan;

QDAFB Queensland Department of Agriculture and Fisheries, Brisbane, Australia;

QSBG Queen Sirikit Botanical Garden, Chiang Mai, Thailand;

RABC Roger A. Beaver collection, Chiang Mai, Thailand;

RIFID Research Institute of Forest Insect Diversity, Namyangju, South Korea;

RJRC Robert J. Rabaglia collection, Annapolis, USA;

RMNH Naturalis Biodiversity Centre, Leiden, Netherlands;

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany;

SEMC University of Kansas Biodiversity Institute, Manhattan, USA;

SMNH Swedish Museum of Natural History, Stockholm, Sweden;

SSC Sunisa Sanguansub collection, Khampaengsaen, Thailand;

TARI Taiwan Agricultural Research Institute, Taichung, Taiwan;

UFFE University of Florida, Forest Entomology Laboratory, Gainesville, USA;

UHZM Universität Hamburg – Zoological Museum, Hamburg, Germany;

VNMN Vietnam National Museum of Nature, Hanoi, Vietnam;

ZFMK Zoological Research Museum Alexander Koenig, Bonn, Germany;

ZIN Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia;

ZMMU Zoological Museum at Moscow State University, Moscow, Russia;

ZSI Zoological Survey of India, Calcutta, India.

All the primary literature as well as types of nearly all 280 species and many of their synonyms known prior to this study were obtained so to assure correct identity of examined specimens. We employed a species concept sensu Hey (2006) and Yeates (2011), that is, species are hypotheses of evolutionary lineages, which are tested with available data. For most species, combinations of morphologically diagnostic characters were taken as evidence for species. In other cases, monophyly based on phylogenies derived from mitochondrial cytochrome oxidase I (COI) and nuclear CAD DNA sequences provided direct evidence of a species equating with an evolutionary lineage (Cognato et al. 2019, 2020b). Decisions to recognize monophyletic groups as species was based on the presence of morphological diagnostic characters and the demonstration of > 10% COI and > 2% CAD average pairwise uncorrected “p” distance between sister clades (Cognato et al. 2020b).

Specimens were primarily photographed by SMS with some by Rachel Osborn (MSU) with a Visionary Digital Passport II system (Dun Inc., Palmyra, VA) using a Canon EOS 5D Mark II, 65.0 mm Canon Macro photo lens, two Dynalite (Union, NJ) MH2015 road flash heads, Dynalite RoadMax MP8 power pack and a Stack Shot (Cognisys, Inc, Traverse City, MI). Montage images were assembled using Helicon Focus Mac Pro 6.7.1 (Helicon Soft, Kharkov, Ukraine). Additional photos were contributed by Wisut Sittichaya (Prince of Songkhla University) and AIC (methods detailed in Smith et al. 2019a).

Specimens were examined using Leica (Wetzlar, Germany) MZ6 and MZ16 stereomicroscopes and illuminated with an Ikea Jansjö LED work lamp (Delft, Netherlands). Length was measured from pronotum apex to the apex of the declivity and a maximum of five specimens per species were measured. Pedicel is not included in the number of funicle segments, following Hulcr and Smith (2010). Taxa are listed alphabetically by genus and then by species within each genus. Unless stated as examined, the location of type species is not given but can be found in Wood and Bright (1992), Bright and Skidmore (1997, 2002), or Bright (2014). This catalog and its supplements contain additional references on the biology of many of the included species. Distribution data were collected from: Wood and Bright 1992; Beaver and Liu 2010; Knížek 2011; Beaver et al. 2014; Zheng et al. 2017; Mandelshtam et al. 2018; Smith et al. 2018b, c; Lin et al. 2019; Sittichaya et al. 2019; Park et al. 2020; Li et al. 2020; Rabaglia et al. 2020b) and other sources are given for each species. New distribution records are denoted with an asterisk.

Terminology

Anatomical terminology is illustrated in Figure 1. Antennal club types (Figs 2, 3) and pronotal types from dorsal (Fig. 4) and lateral (Fig. 5) views follow those in Hulcr et al. (2007). The following commonly used terms are here defined:

alutaceous with fine, leather-like reticulation;

asperity(-ies) small flat denticle-like structures frequently arranged in rows or confined to specific areas;

carina a sharply elevated ridge or keel, not necessarily high or acute (Fig. 6A);

costa a more gradually elevated ridge that is rounded at its crest, without a sharp appearance (Fig. 6B);

declivity/declivital the downward slope of the elytra/pertaining to the declivity;

denticle a small tooth, the sides of which are equal, and the tip is above the middle of the base (Fig. 6C);

glabrous devoid of vestiture;

granule a small rounded protuberance, like a grain of sand (Fig. 6D);

opalescent showing varying colors, like an opal;

serrations row of asperities (flat denticles), a saw-like structure;

shagreened with a rough surface of closely set granules;

spine an elongate projection of the exoskeleton that is longer than its basal width (Fig. 6E);

summit highest point, used for pronotum and elytra, denotes the peak between pronotal frontal slope and disc, and between elytral disc and declivity;

tubercle a small knob-like or rounded protuberance of the exoskeleton (Fig. 6F);

unarmed without cuticular protuberances, e.g., granules, denticles, tubercles or spines;

vermiculate tortuous; marked by repeated twists, like worm tracks.

Figure 1.

Anatomical terminology illustrated on Euwallacea sibsagaricus A lateral habitus B dorsal habitus C ventral habitus.

Figure 2.

Obliquely truncate antennal clubs, types 1 and 2 (Hulcr et al. 2007). Type 1, Anisandrus percristatus; typical type 2, Xyleborus affinis; variation of type 2, Hadrodemius comans.

Figure 3.

Flattened antennal clubs, types 3, 4, and 5 (Hulcr et al. 2007). Type 3, Euwallacea interjectus; typical type 4, Amasa schlichii; variation type 4 (anterior), Fortiborus major; variation type 4 (posterior), Schedlia sumatrana; type 5, Amasa beesoni.

Figure 4.

Dorsal pronotal types. Type 0, Heteroborips seriatus; type 1 rounded, Cnestus gravidus; type 2 basic and parallel-sided, Amasa gibbosa; type 3 subquadrate with anterolateral corners slightly prominent, Cyclorhipidion amasoides; type 4 quadrate with anterolateral corners conspicuous and sides almost parallel, Euwallacea destruens; type 5 conical and elongate, Leptoxyleborus sordicauda; type 6 strongly conical, Anisandrus cryphaloides; type 7 rounded frontally and long, Tricosa cattienensis; type 8 elongate and subquadrate or quadrate, Euwallacea piceus; type 9 long and rounded frontally, Debus amphicranoides; type a long and quadrate frontally, Webbia duodecimspinata; type c conspicuously elongate and quadrate frontally, Streptocranus bicuspis. Drawings modified from Hulcr et al. 2007.

Figure 5.

Lateral pronotal types. Type 0 basic, Xylosandrus mancus; type 1 uniformly rounded without distinct summit, Ambrosiodmus rubricollis; type 2 taller than basic, Euwallacea perbrevis; type 3 short and tall, Anisandrus percristatus; type 4 robust with summit moved anteriad, Schedlia sumatrana; type 5 robust, subquadrate or rounded, Diuncus haberkorni; type 7 disc as long or slightly longer than anterior slope, Tricosa cattienensis; type 8 disc much longer than anterior slope, Cryptoxyleborus stenographus; type 9 anterior slope much longer than disc, Debus amphicranoides; type a very long ‘hooded frontally’, Streptocranus mirabilis; type b long flattened and bulging frontally, Webbia duodecimspinata. Drawings modified from Hulcr et al. 2007.

Figure 6.

Illustrated glossary of terms A carina B costa C denticles D granules E spine F tubercles.

Results and discussion

We identified 34 genera and 315 species as occurring in the study region. Sixty-three new species, 24 new synonyms and 13 new combinations were identified. Previously published records of two additional species were not confirmed as occurring in the region and are therefore considered dubious:

1. Cnestus bicornis (Eggers, 1923) is listed as occurring in India (Assam) (Wood and Bright 1992: 802) but a published Indian record was not found and no Indian specimens could be located. Occurrence in India is therefore doubtful and probably represents a misidentification of the morphologically similar C. bicornioides which does occur in India.

2. Xyleborus aquilus Blandford, 1894 was described from Japan and was previously reported from China (Fujian, Hunan, Sichuan), South Korea and Taiwan. Images of syntypes from NHMUK were compared to the description and diagnosis of Yin et al. (1984) in which the species is reported from China. The syntypes are of a Xyleborus species closely related to X. festivus Eichhoff, 1866 while Yin et al.’s description, illustration, and diagnosis represent an Euwallacea which we were unable to determine to species. Examination of Yin’s specimens in IZAS did not reveal any specimens bearing this name (You Li, pers. comm.). Xyleborus aquilus was reported from Korea by Murayama (1930) but no vouchered specimens have been found or since collected (Park et al. 2020). Beaver and Liu (2010) considered the Taiwan record dubious. It is very likely that X. aquilus is distributed only in Japan.

In part, this study relied on DNA based phylogenies to help resolve generic and species identities and designate species limits (Cognato et al. 2020b). The rubric of monophyly, a sequence difference threshold, and morphological diagnostic characters provided guidance for recognizing species given sufficient specimens, for example, Xyleborus glabratus (Cognato et al. 2019). These DNA based phylogenies and our examination of specimens revealed additional taxonomic problems. 1. Coptodryas Hopkins, 1915, Cryptoxyleborus Wood & Bright, 1992, Microperus Wood, 1980, Xyleborus and Xylosandrus Reitter, 1913 are likely not monophyletic (Cognato et al. 2020b). A more robust dataset including more genes and taxa will likely help resolve the monophyly for some genera. In other cases, reexamination of aberrant species may lead to the stabilization of genera (i.e. reassignment of generic placement for certain species) and recognition of new genera as with Fraudatrix Cognato, Smith & Beaver, 2020 and Tricosa Cognato, Smith & Beaver, 2020 (Cognato et al. 2020a). 2. Several Euwallacea species were para- or polyphyletic (Cognato et al. 2020b). These species (e.g., E. andamanensis Blandford, 1896) exhibit little morphological difference yet demonstrate > 12% COI nucleotide difference (Cognato et al. 2020b). Morphometric analyses as with the E. fornicatus complex may be necessary to help tease out cryptic species (Gomez et al. 2018b). 3. Some species are morphologically variable. For example, Ambrosiophilus osumiensis (Murayama, 1934), demonstrates < 7.5% COI nucleotide difference yet the morphological variation is associated with three species which we synonymized. These morphological characters, mainly the size, position, and number of declivital granules have been traditionally used to recognize species. Given these observations, other suspect variable species should be re-evaluated and caution given to future recognition of new species based on subtle morphological differences.

We discovered a total of 75 new species reported in this and associated publications (Smith et al. 2018c; Cognato et al. 2019, 2020a; Park et al. 2020) and additional new records for southern Thailand (Sittichaya et al. 2019). Given the wide range of some species distributions, examination of the broader region was necessary and as a result we identified new species and nomenclatural changes for Japan and Korea (Smith et al. 2018b; Park et al. 2020), and clarified species limits for some Indo-Malayan species that were incorrectly placed in synonymy (Smith et al. 2020). We also recognized a new species from insular SE Asia (Wang et al. 2020). Thus, this publication is the foundation for a future monograph of all the SE Asian fauna. Based on this study we estimate that 30% of the species are undescribed, but given that scolytine taxonomists have not collected in many areas especially Myanmar, Laos, Cambodia, and Philippines, the number of undiscovered species is likely greater. Also, generic taxonomy will likely continue to improve with delimitation and descriptions of new genera identified among Xyleborus species (Cognato et al. 2020b).

This study provides the first taxonomic review of xyleborine species occurring in mainland SE Asia and adjacent areas. The associated taxonomic tools, Lucid key, DNA sequences, and images complement this monograph and provide additional resources for species and generic identifications (Smith et al. 2019a; Cognato et al. 2020b). We consider the Lucid key and DNA database living “documents”, and as we continue to treat the Asian fauna, we will amend these tools with the goal of taxon completion.

Checklist of the Xyleborini of Southeast Asia

Amasa Lea, 1894

Pseudoxyleborus Eggers, 1930

Anaxyleborus Wood, 1980

Amasa aspersa (Sampson, 1921)

Amasa beesoni (Eggers, 1930)

Amasa concitata (Schedl, 1969a)

Amasa cycloxyster sp. nov.

Amasa cylindrotomica (Schedl, 1939b)

Xyleborus semitruncatus Schedl, 1942c

Xyleborus truncatellus Schedl, 1951a

Xyleborus jucundus Schedl, 1954

Amasa eugeniae (Eggers, 1930)

Amasa galeoderma sp. nov.

Amasa gibbosa sp. nov.

Amasa lini sp. nov.

Amasa opalescens (Schedl, 1937a)

Amasa resecta (Eggers, 1923)

Xyleborus abruptus Eggers, 1923

Xyleborus opacicauda Eggers, 1940

Amasa schlichii (Stebbing, 1907)

Acanthotomicus truncatus Stebbing, 1907

Xyleborus glaber Eggers, 1930

Xyleborus uniseriatus Eggers, 1936b

Xyleborus verax Schedl, 1939b

Amasa tropidacron sp. nov.

Amasa versicolor (Sampson, 1921)

Amasa youlii sp. nov.

Ambrosiodmus Hopkins, 1915a

Phloeotrogus Motschulsky, 1863

Brownia Nunberg, 1963

Ambrosiodmus asperatus (Blandford, 1895)

Xyleborus nepotulus Eggers, 1923

Xyleborus citri Beeson, 1930

Xyleborus nepotulomorphus Eggers, 1936b

Ambrosiodmus brunneipes (Eggers, 1940)

Ambrosiodmus conspectus (Schedl, 1964b)

Ambrosiodmus lewisi (Blandford, 1894b)

Ozopemon tuberculatus Strohmeyer, 1912

Xyleborus lewekianus Eggers, 1923

Xyleborus tegalensis Eggers, 1923

Ambrosiodmus minor (Stebbing, 1907)

Xyleborus crassus Hagedorn, 1910a

Ambrosiodmus rubricollis (Eichhoff, 1876a)

Xyleborus taboensis Schedl, 1952b

Xyleborus strohmeyeri Schedl, 1975b

Ambrosiophilus Hulcr & Cognato, 2009

Ambrosiophilus atratus (Eichhoff, 1876a)

Xyleborus collis Niisima, 1910

Ambrosiophilus caliginestris sp. nov.

Ambrosiophilus consimilis (Eggers, 1923), comb. nov.

Ambrosiophilus cristatulus (Schedl, 1953b)

Ambrosiophilus indicus sp. nov.

Ambrosiophilus lannaensis sp. nov.

Ambrosiophilus latisulcatus (Eggers, 1940)

Ambrosiophilus osumiensis (Murayama, 1934)

Xyleborus metanepotulus Eggers, 1939b

Xyleborus nodulosus Eggers, 1941b, syn. nov.

Xyleborus pernodulus Schedl, 1957

Xyleborus hunanensis Browne, 1983b

Ambrosiophilus peregrinus Smith & Cognato, 2015

Ambrosiophilus papilliferus sp. nov.

Ambrosiophilus satoi (Schedl, 1966b)

Ambrosiophilus sexdentatus (Eggers, 1940)

Ambrosiophilus subnepotulus (Eggers, 1930)

Xyleborus cristatuloides Schedl, 1971a, syn. nov.

Ambrosiophilus sulcatus (Eggers, 1930)

Xyleborus sulcatulus Eggers, 1939a, syn. nov.

Xyleborus sinensis Eggers, 1941b, syn. nov.

Ambrosiophilus wantaneeae sp. nov.

Ancipitis Hulcr & Cognato, 2013

Ancipitis puer (Eggers, 1923)

Xyleborus ceramensis Schedl, 1937a

Ancipitis punctatissimus (Eichhoff), 1880

Xyleborus spatulatus Blandford, 1896b

Anisandrus Ferrari, 1867

Anisandrus achaete sp. nov.

Anisandrus apicalis (Blandford, 1894b)

Anisandrus auco sp. nov.

Anisandrus auratipilus sp. nov.

Anisandrus carinensis (Eggers, 1923), comb. nov.

Anisandrus congruens sp. nov.

Anisandrus cristatus (Hagedorn, 1908), comb. nov., stat. res.

Xyleborus fabricii Schedl, 1964c

Anisandrus cryphaloides sp. nov.

Anisandrus dispar (Fabricius, 1792)

Bostrichus brevis Panzer, 1793

Bostrichus thoracicus Panzer, 1793

Scolytus pyri Peck, 1817

Bostrichus tachygraphus Sahlberg, 1836

Bostrichus ratzeburgi Kolenati, 1846

Xyleborus ishidai Niisima, 1909

Anisandrus aequalis Reitter, 1913

Anisandrus swainei Drake, 1921

Xyleborus dispar rugulosus Eggers, 1922

Xyleborus cerasi Eggers, 1937

Xyleborus khinganensis Murayama, 1943

Anisandrus eggersi (Beeson, 1930)

Anisandrus feronia sp. nov.

Anisandrus geminatus (Hagedorn, 1904)

Anisandrus hera sp. nov.

Anisandrus hirtus (Hagedorn, 1904)

Xyleborus hagedorni Stebbing, 1914

Xyleborus hirtuosus Beeson, 1930

Xyleborus hagedornianus Schedl, 1952d

Xyleborus tectonae Nunberg, 1956

Xyleborus hirtipes Schedl, 1969b, syn. nov.

Xyleborus taiwanensis Browne, 1980b

Anisandrus improbus (Sampson, 1913)

Anisandrus klapperichi (Schedl, 1955b), comb. nov.

Anisandrus lineatus (Eggers, 1930)

Xyleborus melancranis Beeson, 1930

Anisandrus longidens (Eggers, 1930)

Anisandrus maiche (Kurentzov, 1941)

Xyleborus maiche Eggers, 1942

Anisandrus mussooriensis (Eggers, 1930)

Anisandrus niger (Sampson, 1912)

Anisandrus paragogus sp. nov.

Anisandrus percristatus (Eggers, 1939a), comb. nov.

Anisandrus sinivali sp. nov.

Anisandrus ursulus (Eggers, 1923)

Anisandrus venustus sp. nov.

Anisandrus xuannu sp. nov.

Arixyleborus Hopkins, 1915a

Xyleboricus Eggers, 1923

Arixyleborus crassior sp. nov.

Arixyleborus grandis (Schedl, 1942c)

Arixyleborus granifer (Eichhoff, 1878a)

Xyleborus granifer borneensis Schedl, 1965

Arixyleborus granulifer (Eggers, 1923)

Arixyleborus hirsutulus Schedl, 1969a

Arixyleborus leprosulus Schedl, 1953b

Arixyleborus aralidii Nunberg, 1961

Arixyleborus malayensis (Schedl, 1954)

Arixyleborus mediosectus (Eggers, 1923)

Arixyleborus angulatus Schedl, 1942a

Arixyleborus minor (Eggers, 1940)

Arixyleborus trux Schedl, 1975c

Arixyleborus moestus (Eggers, 1930)

Arixyleborus nudulus Smith, Rabaglia & Cognato, 2018 (in Smith et al. 2018c)

Arixyleborus phiaoacensis sp. nov.

Arixyleborus puberulus (Blandford, 1896b)

Xyleborus hirtipennis Eggers, 1940

Arixyleborus resecans (Eggers, 1930), comb. nov.

Arixyleborus rugosipes Hopkins, 1915a

Webbia medius Eggers, 1927b

Webbia camphorae Eggers, 1936a

Arixyleborus scabripennis (Blandford, 1896b)

Arixyleborus setosus sp. nov.

Arixyleborus silvanus sp. nov.

Arixyleborus sittichayai sp. nov.

Arixyleborus suturalis (Eggers, 1936b)

Arixyleborus titanus sp. nov.

Arixyleborus tuberculatus (Eggers, 1940)

Arixyleborus yakushimanus (Murayama, 1958)

Beaverium Hulcr & Cognato, 2009

Beaverium lantanae (Eggers, 1930)

Beaverium latus (Eggers, 1923)

Beaverium magnus (Niisima, 1910)

Xyleborus rufobrunneus var. dihingensis Eggers, 1930

Xyleborus chujoi Schedl, 1951a

Cnestus Sampson, 1911

Tosaxyleborus Murayama, 1950

Cnestus ater (Eggers, 1923)

Xyleborus retusiformis Schedl, 1936d

Cnestus aterrimus (Eggers, 1927a)

Xyleborus glabripennis Schedl, 1942a

Tosaxyleborus pallidipennis Murayama, 1950

Cnestus nitens Browne, 1955

Cnestus murayamai Schedl, 1962a

Cnestus murayamai Browne, 1963

Cnestus pseudosuturalis Schedl, 1964c

Cnestus maculatus Browne, 1983b

Cnestus bicornioides (Schedl, 1952a)

Cnestus gravidus (Blandford, 1898)

Cnestus improcerus (Sampson, 1921)

Cnestus mutilatus (Blandford, 1894b)

Xyleborus sampsoni Eggers, 1930

Xyleborus banjoewangi Schedl, 1939b

Xyleborus taitonus Eggers, 1939b

Cnestus nitidipennis (Schedl, 1951a)

Cnestus protensus (Eggers, 1930)

Cnestus rostratus Schedl, 1977, syn. nov.

Cnestus quadrispinosus Sittichaya & Beaver, 2018

Cnestus suturalis (Eggers, 1930)

Cnestus testudo (Eggers, 1939b)

Coptodryas Hopkins, 1915a

Coptodryas amydra sp. nov.

Coptodryas bella (Sampson, 1921)

Coptodryas carinata sp. nov.

Coptodryas concinna (Beeson, 1930)

Xyleborus flexicostatus Schedl, 1942c

Coptodryas confusa Hopkins, 1915a

Xyleborus cryphaloides Schedl, 1942a

Coptodryas elegans (Sampson, 1923)

Coptodryas inornata sp. nov.

Coptodryas mus (Eggers, 1930)

Coptodryas nudipennis (Schedl, 1951a)

Coptodryas quadricostata (Schedl, 1942c)

Cryptoxyleborus Wood & Bright, 1992

Cryptoxyleborus Schedl, 1937a

Cryptoxyleborus barbieri Schedl, 1953a

Cryptoxyleborus confusus Browne, 1950

Cryptoxyleborus eggersi Schedl, 1936c

Cryptoxyleborus dryobalanopsis Schedl, 1942a

Xyleborus eggersianus Schedl, 1960b

Cryptoxyleborus percuneolus (Schedl, 1951a)

Cryptoxyleborus quadriporus Beaver, 1990

Cryptoxyleborus stenographus (Schedl, 1971b)

Cryptoxyleborus subnaevus Schedl, 1937a

Cryptoxyleborus turbineus (Sampson, 1923)

Cyclorhipidion Hagedorn, 1912b

Terminalinus Hopkins, 1915a

Notoxyleborus Schedl, 1934b

Kelantanius Nunberg, 1961

Cyclorhipidion amasoides sp. nov.

Cyclorhipidion amputatum sp. nov.

Cyclorhipidion armiger (Schedl, 1953c), comb. nov.

Cyclorhipidion bodoanum (Reitter, 1913)

Xyleborus punctulatus Kurentzov, 1948

Xyleborus californicus Wood, 1975b

Xyleborus misatoensis Nobuchi, 1981a, syn. nov.

Cyclorhipidion circumcisum (Sampson, 1921)

Xyleborus obtusus Eggers, 1923

Xyleborus subobtusus Schedl, 1942a

Cyclorhipidion denticauda sp. nov.

Cyclorhipidion distinguendum (Eggers, 1930)

Xyleborus fukiensis Eggers, 1941b, syn. nov.

Xyleborus ganshoensis Murayama, 1952, syn. nov.

Cyclorhipidion fouqueti (Schedl, 1937b)

Cyclorhipidion inarmatum (Eggers, 1923)

Xyleborus vagans Schedl, 1977, syn. nov.

Cyclorhipidion japonicum (Nobuchi, 1981a)

Cyclorhipidion miyazakiense (Murayama, 1936)

Xyleborus armipennis Schedl, 1953c

Xyleborus wakayamensis Nobuchi, 1981a

Cyclorhipidion muticum sp. nov.

Cyclorhipidion neocavipenne (Schedl, 1977)

Cyclorhipidion obesulum sp. nov.

Cyclorhipidion ohnoi (Browne, 1980a)

Cyclorhipidion pelliculosum (Eichhoff, 1878a)

Xyleborus seiryorensis Murayama, 1930

Xyleborus quercus Kurentzov, 1948

Xyleborus starki Nunberg, 1956

Cyclorhipidion perpilosellum (Schedl, 1935a)

Xyleborus punctatopilosus Schedl, 1936b

Cyclorhipidion petrosum sp. nov.

Cyclorhipidion pilipenne (Eggers, 1940)

Cyclorhipidion pruinosulum Browne, 1979

Cyclorhipidion pruinosum (Blandford, 1896b)

Xyleborus arcticollis Blandford, 1896b

Xyleborus decipiens Eggers, 1923

Cyclorhipidion sisyrnophorum (Hagedorn, 1910a)

Cyclorhipidion tenuigraphum (Schedl, 1953) stat. res.

Cyclorhipidion trucaudinum sp. nov.

Cyclorhipidion umbratum (Eggers, 1941b)

Cyclorhipidion vigilans (Schedl, 1939b)

Cyclorhipidion xeniolum sp. nov.

Cyclorhipidion xyloteroides (Eggers, 1939b)

Debus Hulcr & Cognato, 2010a

Debus adusticollis (Motschulsky, 1863)

Xyleborus vestitus Schedl, 1931

Debus amphicranoides (Hagedorn, 1908)

Xyleborus amphicranoides latecavatus Eggers, 1927b

Xyleborus amphicranoides parvior Browne, 1981b

Debus birmanus (Eggers, 1930)

Debus detritus (Eggers, 1927a)

Xyleborus maniensis Browne, 1981a

Debus emarginatus (Eichhoff, 1878a)

Xyleborus exesus Blandford, 1894b

Ips cinchonae Veen, 1897

Xyleborus cordatus Hagedorn, 1910a

Xyleborus palmeri Hopkins, 1915a

Xyleborus terminaliae Hopkins, 1915a

Xyleborus emarginatus semicircularis Schedl, 1973

Debus pumilus (Eggers, 1923)

Xyleborus cylindricus Eggers, 1927b

Xyleborus neocylindricus Schedl, 1942a

Ips kelantanensis Browne, 1955

Xyleborus ipidia Schedl, 1972a

Xyleborus planodeclivis Browne, 1974

Debus quadrispinus (Motschulsky, 1863), comb. nov.

Xyleborus fallax Eichhoff, 1878a, syn. nov.

Xyleborus amphicranulus Eggers, 1923

Xyleborus fastigatus Schedl, 1935a

Debus shoreae (Stebbing, 1907)

Tomicus assamensis Stebbing, 1909

Diuncus Hulcr & Cognato, 2009

Diuncus ciliatoformis (Schedl, 1953d) stat. res.

Diuncus corpulentus (Eggers, 1930)

Diuncus dossuarius (Eggers, 1923)

Diuncus haberkorni (Eggers, 1920)

Xyleborus approximatus Schedl, 1951a

Xyleborus taichuensis Schedl, 1952b

Xyleborus potens Schedl, 1964a

Diuncus javanus (Eggers, 1923)

Xyleborus perdix Schedl, 1939a

Diuncus justus (Schedl, 1931)

Xyleborus marginicollis Schedl, 1936c

Xyleborus ciliatus Eggers, 1940

Xyleborus apiculatus Schedl, 1942a

Diuncus mucronatulus (Eggers, 1930)

Diuncus mucronatus (Eggers, 1923)

Diuncus quadrispinulosus (Eggers, 1923)

Xyleborus parvispinosus palembangensis Schedl, 1939b

Xyleborus parvispinosus Schedl, 1951a

Dryoxylon Bright & Rabaglia, 1999

Dryoxylon onoharaense (Murayama, 1934)

Eccoptopterus Motschulsky, 1863

Platydactylus Eichhoff, 1886

Eurydactylus Hagedorn, 1909

Eccoptopterus limbus Sampson, 1911

Xyleborus auratus Eggers, 1923

Xyleborus squamulosus duplicatus Eggers, 1923

Xyleborus squamulosus Eggers, 1923

Eccoptopterus spinosus (Olivier, 1800)

Eccoptopterus sexspinosus Motschulsky, 1863

Xyleborus abnormis Eichhoff, 1869

Platydactylus gracilipes Eichhoff, 1886

Xyleborus sexspinosus multispinosus Hagedorn, 1908

Xyleborus collaris Eggers, 1923

Eccoptopterus sagittarius Schedl, 1939b

Eccoptopterus sexspinosus pluridentatus Schedl, 1942c

Xyleborus eccoptopterus Schedl, 1951b

Euwallacea Hopkins, 1915a

Wallacellus Hulcr & Cognato, 2010a

Euwallacea andamanensis (Blandford, 1896b)

Xyleborus noxius Sampson, 1913

Xyleborus siobanus Eggers, 1923

Xyleborus burmanicus Beeson, 1930

Xyleborus granulipennis Eggers, 1930

Xyleborus intextus Beeson, 1930

Xyleborus senchalensis Beeson, 1930

Xyleborus talumalai Browne, 1966

Euwallacea aplanatus (Wichmann, 1914)

Euwallacea destruens (Blandford, 1896b)

Xyleborus barbatus Hagedorn, 1910a

Xyleborus barbatulus Schedl, 1934b

Xyleborus pseudobarbatus Schedl, 1942a

Xyleborus nandarivatus Schedl, 1950a

Xyleborus procerrimus Schedl, 1969a

Euwallacea fornicatior (Eggers, 1923)

Xyleborus schultzei Schedl, 1951a

Euwallacea fornicatus (Eichhoff, 1868b)

Xyleborus whitfordiodendrus Schedl, 1942a

Xyleborus tapatapaoensis Schedl, 1951b

Euwallacea funereus (Lea, 1910)

Xyleborus nepos Eggers, 1923

Xyleborus nepos robustus Schedl, 1933

Xyleborus signatus Schedl, 1949

Euwallacea geminus sp. nov.

Euwallacea gravelyi (Wichmann, 1914) stat. res.

Xyleborus ovalicollis Eggers, 1930

Xyleborus barbatomorphus Schedl, 1951a, syn. nov.

Euwallacea interjectus (Blandford, 1894c)

Xyleborus pseudovalidus Eggers, 1925

Euwallacea kuroshio Gomez & Hulcr, 2018 (in Gomez et al. 2018b)

Euwallacea luctuosus (Eggers, 1939a)

Euwallacea malloti (Eggers, 1930)

Euwallacea minutus (Blandford, 1894b), comb. nov.

Xyleborus breviusculus Schedl, 1942a

Xyleborus pernitidus Schedl, 1954

Euwallacea neptis sp. nov.

Euwallacea perbrevis (Schedl, 1951a)

Xyleborus molestulus Wood, 1975, syn. nov.

Euwallacea piceus (Motschulsky, 1863)

Xyleborus indicus Eichhoff, 1878a

Xyleborus imitans Eggers, 1927a

Xyleborus indicus subcoriaceus Eggers, 1927b

Xyleborus samoensis Beeson, 1929

Euwallacea semiermis (Schedl, 1934c)

Euwallacea semirudis (Blandford, 1896b) stat. res.

Xyleborus dubius Eggers, 1923

Xyleborus sereinuus Eggers, 1923

Xyleborus hybridus Eggers, 1927b

Xyleborus interruptus Eggers, 1940

Xyleborus neohybridus Schedl, 1942a, syn. nov.

Xyleborus longehirtus Nunberg, 1956

Euwallacea sibsagaricus (Eggers, 1930)

Xyleborus dalbergiae Eggers, 1930

Xyleborus tonkinensis Schedl, 1934a, syn. nov.

Euwallacea similis (Ferrari, 1867)

Bostrichus ferrugineus Bohemann, 1858

Xyleborus parvulus Eichhoff, 1868b

Xyleborus dilatatus Eichhoff, 1878b

Xyleborus submarginatus Blandford, 1896b

Xyleborus bucco Schaufuss, 1897

Xyleborus capito Schaufuss, 1897

Xyleborus novaguineanus Schedl, 1936b

Xyleborus dilatatulus Schedl, 1953a

Euwallacea subalpinus sp. nov.

Euwallacea testudinatus sp. nov.

Euwallacea validus (Eichhoff, 1876a)

Euwallacea velatus (Sampson, 1913)

Xyleborus assamensis Eggers, 1930

Xyleborus rudis Eggers, 1930, syn. nov.

Xyleborus asperipennis Eggers, 1934b

Fortiborus Hulcr & Cognato, 2010a

Fortiborus macropterus (Schedl, 1935b)

Fortiborus major (Stebbing, 1909)

Xyleborus siclus Schedl, 1936d

Fortiborus pseudopilifer (Schedl, 1936a)

Fraudatrix Cognato, Smith & Beaver, 2020

Fraudatrix cuneiformis (Schedl, 1958b)

Fraudatrix melas (Eggers, 1927b)

Fraudatrix simplex (Browne, 1949)

Hadrodemius Wood, 1980

Hadrodemius comans (Sampson, 1919)

Xyleborus amorphus Eggers, 1926

Xyleborus metacomans Eggers, 1930

Hadrodemius globus (Blandford, 1896b)

Xyleborus ursus Eggers, 1923

Xyleborus ursus fuscus Eggers, 1923

Xyleborus tomentosus Eggers, 1939a

Hadrodemius pseudocomans (Eggers, 1930)

Xyleborus artecomans Schedl, 1953c

Heteroborips Reitter, 1913

Heteroborips fastigatus sp. nov.

Heteroborips indicus sp. nov.

Heteroborips seriatus (Blandford, 1894b)

Xyleborus orientalis Eggers, 1933b

Xyleborus todo Kôno, 1938

Xyleborus orientalis aceris Kurentzov, 1941

Xyleborus orientalis kalopanacis Kurentzov, 1941

Xyleborus perorientalis Schedl, 1957

Heteroborips tristis (Eggers, 1930), comb. nov.

Immanus Hulcr & Cognato, 2013

Immanus desectus (Eggers, 1923)

Xyleborus desectus arduus Schedl, 1942a

Immanus sarawakensis (Eggers, 1923)

Leptoxyleborus Wood, 1980

Leptoxyleborus machili (Niisima, 1910), comb. nov.

Xyleborus depressus Eggers, 1923

Xyleborus kojimai Murayama, 1936

Xyleborus sejugatus Schedl, 1942a

Leptoxyleborus sordicauda (Motschulsky, 1863)

Phloeotrogus attenuatus Motschulsky, 1863

Xyleborus concisus Blandford, 1894b

Xyleborus marginatus Eggers, 1927b

Xyleborus sordicaudulus Eggers, 1927b

Xyleborus incurvus Eggers, 1930

Xyleborus sordicaudulus peguensis Eggers, 1930

Microperus Wood, 1980

Microperus alpha (Beeson, 1929)

Microperus chrysophylli (Eggers, 1930)

Microperus corporaali (Eggers, 1923)

Microperus cruralis (Schedl, 1975b), comb. nov.

Microperus diversicolor (Eggers, 1923)

Xyleborus myristicae Schedl, 1939b

Xyleborus brevipilosus Eggers, 1940

Xyleborus theae Eggers, 1940

Xyleborus cylindripennis Schedl, 1954

Xyleborus atavus Schedl, 1979b

Microperus fulvulus (Schedl, 1942c) stat. res.

Xyleborus fulvus Schedl, 1939b

Microperus kadoyamaensis (Murayama, 1934)

Xyleborus denseseriatus Eggers, 1941b, syn. nov.

Xyleborus nameranus Murayama, 1954

Xyleborus pubipennis Schedl, 1974, syn. nov.

Xyleborus huangi Browne, 1983b

Microperus kirishimanus (Murayama, 1955)

Microperus latesalebrinus sp. nov.

Microperus minax sp. nov.

Microperus nudibrevis (Schedl, 1942a)

Microperus nugax (Schedl, 1939a)

Xyleborus pertuberculatus Eggers, 1940

Microperus perparvus (Sampson, 1922b)

Xyleborus tsukubanus Murayama, 1954

Microperus pometianus (Schedl, 1939a)

Microperus quercicola (Eggers, 1926)

Xyleborus izuensis Murayama, 1952

Microperus recidens (Sampson, 1923)

Xyleborus minusculus Eggers, 1923

Xyleborus minutissimus Eggers, 1930

Xyleborus crassitarsus Schedl, 1936d

Xyleborus artegraphus Schedl, 1942c

Xyleborus extensus Schedl, 1955a

Xyleborus tuberculosus Browne, 1981b

Microperus sagmatus sp. nov.

Microperus undulatus (Sampson, 1919)

Xyleborus leprosulus Schedl, 1936d

Planiculus Hulcr & Cognato, 2010a

Planiculus bicolor (Blandford, 1894b)

Xyleborus laevis Eggers, 1923

Xyleborus bicolor unimodus Beeson, 1929

Xyleborus rodgeri Beeson, 1930

Xyleborus rodgeri privatus Beeson, 1930

Xyleborus rameus Schedl, 1940a

Xyleborus artelaevis Schedl, 1942a

Xyleborus ashuensis Murayama, 1954

Xyleborus tumidus Schedl, 1975c

Xyleborus filiformis Schedl, 1975c

Xyleborus glabratulus Browne, 1983a

Planiculus limatus (Schedl, 1942b)

Xyleborus subemarginatus Eggers, 1940

Xyleborus subparallelus Eggers, 1940

Planiculus shiva (Maiti & Saha, 1986), comb. nov.

Pseudowebbia Browne, 1961a

Pseudowebbia trepanicauda (Eggers, 1923)

Schedlia Browne, 1950b

Schedlia allecta (Schedl, 1942c)

Schedlia sumatrana (Hagedorn, 1908)

Stictodex Hulcr & Cognato, 2013

Stictodex dimidiatus (Eggers, 1927a)

Xyleborus dorsosulcatus Beeson, 1930, syn. nov.

Xyleborus tunggali Schedl, 1936d

Xyleborus decumans Schedl, 1953b

Xyleborus cruciatus Schedl, 1973

Streptocranus Schedl, 1939b

Streptocranus bicolor (Browne, 1949)

Streptocranus bicuspis (Eggers, 1940)

Streptocranus recurvus Browne, 1949

Streptocranus fragilis Browne, 1949

Streptocranus mirabilis Schedl, 1939b

Streptocranus petilus sp. nov.

Tricosa Cognato, Smith & Beaver, 2020

Tricosa cattienensis Cognato, Smith & Beaver, 2020 (in Cognato et al. 2020a)

Tricosa indochinensis Cognato, Smith & Beaver, 2020 (in Cognato et al. 2020a)

Tricosa jacula Cognato, Smith & Beaver, 2020 (in Cognato et al. 2020a)

Tricosa metacuneolus (Eggers, 1940)

Xyleborus kaimochii Nobuchi, 1981a

Truncaudum Hulcr & Cognato, 2010a

Truncaudum agnatum (Eggers, 1923)

Xyleborus polyodon Eggers, 1923

Xyleborus gratiosus Schedl 1942a

Xyleborus nutans Schedl, 1942a

Xyleborus delicatus Schedl, 1955a

Xyleborus subagnatus Wood, 1992

Truncaudum bullatum sp. nov.

Webbia Hopkins, 1915b

Xelyborus Schedl, 1939a

Prowebbia Browne, 1962

Webbia biformis Browne, 1958

Webbia cornuta Schedl, 1942a

Webbia dasyura Browne, 1981a

Webbia dipterocarpi Hopkins, 1915b

Webbia diversicauda Browne, 1972

Webbia duodecimspinata Schedl, 1942a

Webbia pabo Sampson, 1922

Webbia quatuordecimspinata Sampson, 1921

Webbia trigintispinata Sampson, 1922

Webbia vigintisexspinata Sampson, 1922

Webbia mucronatus Eggers, 1927, syn. nov.

Webbia turbinata Maiti & Saha, 1986

Xyleborinus Reitter, 1913

Xyleborinus andrewesi (Blandford, 1896b)

Xyleborus persphenos Schedl, 1970a

Xyleborus insolitus Bright, 1972

Cryptoxyleborus gracilior Browne, 1984a

Xyleborinus artestriatus (Eichhoff, 1878b)

Xyleborus laticollis Blandford, 1896b

Xyelborus angustior Eggers, 1925, syn. nov.

Xyleborus rugipennis Schedl, 1953b

Xyleborus undatus Schedl, 1974, syn. nov.

Xyleborus beaveri Browne, 1978

Xyleborinus attenuatus (Blandford, 1894b)

Xyleborus alni Niisima, 1909

Xyleborus canus Niisima, 1909

Xyleborinus cuneatus sp. nov.

Xyleborinus disgregus sp. nov.

Xyleborinus echinopterus sp. nov.

Xyleborinus ephialtodes sp. nov.

Xyleborinus exiguus (Walker, 1859)

Xyleborus muriceus Eichhoff, 1878a

Xyleborus diversus Schedl, 1954b, syn. nov.

Xyleborus perexiguus Schedl, 1971b

Xyleborus ankius Schedl, 1975c

Xyleborinus huifenyinae sp. nov.

Xyleborinus jianghuasuni sp. nov.

Xyleborinus octiesdentatus (Murayama, 1931)

Xyleborinus perpusillus (Eggers, 1927a)

Xyleborus perminutissimus Schedl, 1934b

Xyleborus angustatulus Schedl, 1942c

Xyleborinus saxesenii (Ratzeburg, 1837)

Xyleborus dohrni Wollaston, 1854

Xyleborus decolor Boieldieu, 1859

Xyleborus aesculi Ferrari, 1867

Xyleborus subdepressus Rey, 1883

Xyleborus frigidus Blackburn, 1885

Xyleborus arbuti Hopkins, 1915a

Xyleborus floridensis Hopkins, 1915a

Xyleborus pecanis Hopkins, 1915a

Xyleborus quercus Hopkins, 1915a

Xyleborus sobrinus Eichhoff, 1876a

Xyleborinus librocedri Swaine, 1934

Xyleborinus tsugae Swaine, 1934

Xyleborus pseudogracilis Schedl, 1937c

Xyleborus retrusus Schedl, 1940b

Xyleborus peregrinus Eggers, 1944

Xyleborus pseudoangustatus Schedl, 1948

Xyleborus paraguayensis Schedl, 1949

Xyleborus opimulus Schedl, 1976

Xyleborinus schaufussi (Blandford, 1894b)

Xyleborus kraunhiae Niisima, 1910

Xyleborinus sculptilis (Schedl, 1964b)

Xyleborinus speciosus (Schedl, 1975b)

Xyleborinus spinipennis (Eggers, 1930)

Xyleborinus subgranulatus (Eggers, 1930)

Xyleborinus subspinosus (Eggers, 1930) stat. res.

Xyleborinus thaiphami sp. nov.

Xyleborinus tritus sp. nov.

Xyleborus Eichhoff, 1864

Anaeretus Dugès, 1888

Progenius Blandford, 1896a

Mesoscolytus Broun, 1904

Boroxylon Hopkins, 1915a

Xyleborus affinis Eichhoff, 1868b

Xyleborus affinis fuscobrunneus Eichhoff, 1878b

Xyleborus affinis mascarensis Eichhoff, 1878b

Xyleborus affinis parvus Eichhoff, 1878b

Xyleborus sacchari Hopkins, 1915a

Xyleborus societatis Beeson, 1935a

Xyleborus subaffinis Eggers, 1933a

Xyleborus proximus Eggers, 1943

Xyleborus bidentatus (Motschulsky, 1863)

Xyleborus subcostatus Eichhoff, 1869a

Xyleborus riehlii Eichhoff, 1878b

Progenius fleutiauxi Blandford, 1896a

Xyleborus laeviusculus Blandford, 1896a

Boroxylon stephegynis Hopkins, 1915a

Boroxylon webbi Hopkins, 1915a

Xyleborus subcostatus dearmatus Eggers, 1923

Xyleborus brevidentatus Eggers, 1930

Xyleborus quadridens Eggers, 1930

Xyleborus cognatus Blandford, 1896a

Xyleborus ferrugineus (Fabricius, 1801)

Tomicus trypanaeoides Wollaston, 1867

Xyleborus confusus Eichhoff, 1868a

Xyleborus fuscatus Eichhoff, 1868a

Xyleborus retusicollis Zimmermann, 1868

Xyleborus amplicollis Eichhoff, 1869

Xyleborus insularis Sharp, 1885

Xyleborus tanganus Hagedorn, 1910a

Xyleborus nyssae Hopkins, 1915a

Xyleborus soltaui Hopkins, 1915a

Xyleborus hopkinsi Beeson, 1929

Xyleborus argentinensis Schedl, 1931

Xyleborus rufopiceus Eggers, 1932

Xyleborus schedli Eggers, 1934a

Xyleborus nesianus Beeson, 1940

Xyleborus notatus Eggers, 1941a

Xyleborus subitus Schedl, 1949

Xyleborus festivus Eichhoff, 1876a

Xyleborus pinicola Eggers, 1930

Xyleborus detectus Schedl, 1975a

Xyleborus pinivorus Browne, 1980a

Xyleborus glabratus Eichhoff, 1877

Xyleborus kumamotoensis Murayama, 1934

Xyleborus insidiosus Cognato & Smith, 2019

Xyleborus muticus Blandford, 1894b

Xyleborus lignographus Schedl, 1953c, syn. nov.

Xyleborus conditus Schedl, 1971b, syn. nov.

Xyleborus mysticulus Cognato & Smith, 2019

Xyleborus opacus sp. nov.

Xyleborus perforans (Wollaston, 1857)

Bostrichus testaceus Walker, 1859

Xyleborus duponti Montrouzier, 1861

Anodius denticulus Motschulsky, 1863

Anodius tuberculatus Motschulsky, 1863

Xyleborus kraatzii Eichhoff, 1868b

Xyleborus kraatzii philippinensis Eichhoff, 1878b

Xyleborus immaturus Blackburn, 1885

Xylopertha hirsuta Lea, 1894

Xyleborus whitteni Beeson, 1935b

Xyleborus apertus Schedl, 1939a

Xyleborus criticus Schedl, 1950b

Xyleborus cylindrus Schedl, 1951a

Xyleborus shionomisakiensis Murayama, 1951

Xyleborus minimus Schedl, 1955a

Xyleborus pfeilii (Ratzeburg, 1837)

Bostrichus alni Mulsant & Rey, 1856

Xyleborus vicarius Eichhoff, 1876a

Xyleborus adumbratus Blandford, 1894b

Xyleborus septentrionalis Niisima, 1909

Xyleborus singhi Park & Smith, 2020

Xyleborus sunisae sp. nov.

Xyleborus volvulus (Fabricius, 1775)

Xyleborus torquatus Eichhoff, 1868b

Xyleborus alternans Eichhoff, 1869

Xyleborus badius Eichhoff, 1869

Xyleborus interstitalis Eichhoff, 1878b

Xyleborus guanajuatensis Dugès, 1887

Xyleborus grenadensis Hopkins, 1915a

Xyleborus hubbardi Hopkins, 1915a

Xyleborus rileyi Hopkins, 1915a

Xyleborus schwarzi Hopkins, 1915a

Xyleborus continentalis Eggers, 1920

Xyleborus silvestris Beeson, 1929

Xyleborus vagabundus Schedl, 1949

Xyleborus granularis Schedl, 1950b

Xyleborus yunnanensis sp. nov.

Xylosandrus Reitter, 1913

Apoxyleborus Wood, 1980

Xylosandrus adherescens Schedl, 1971b

Xylosandrus amputatus (Blandford, 1894c)

Xyleborus melli Schedl, 1938

Xylosandrus beesoni Saha, Maiti & Chakraborti, 1992

Xylosandrus bellinsulanus sp. nov.

Xylosandrus borealis Nobuchi, 1981b

Xylosandrus brevis (Eichhoff, 1877)

Xyleborus cucullatus Blandford, 1894b

Xyleborus montanus Niisima, 1910

Xylosandrus compactus (Eichhoff, 1876a)

Xyleborus morstatti Hagedorn, 1912a

Xylosandrus crassiusculus (Motschulsky, 1866)

Xyleborus semiopacus Eichhoff, 1878b

Xyleborus semigranosus Blandford, 1896b

Dryocoetes bengalensis Stebbing, 1908

Xyleborus mascarenus Hagedorn, 1908

Xyleborus ebriosus Niisima, 1909

Xyleborus okoumeensis Schedl, 1935b

Xyleborus declivigranulatus Schedl, 1936d

Xylosandrus dentipennis Park & Smith, 2020

Xylosandrus derupteterminatus (Schedl, 1951a)

Xylosandrus discolor (Blandford, 1898)

Xyleborus posticestriatus Eggers, 1939b

Xylosandrus diversepilosus (Eggers, 1941b)

Xylosandrus eupatorii (Eggers, 1940)

Xylosandrus formosae (Wood), comb. nov.

Xyleborus formosanus Browne, 1981a

Xylosandrus germanus (Blandford, 1894b)

Xyleborus orbatus Blandford, 1894b

Xylosandrus jaintianus (Schedl, 1967)

Xylosandrus mancus (Blandford, 1898)

Xyleborus abruptus Sampson, 1914

Xyleborus mancus formosanus Eggers, 1930

Xylosandrus mesuae (Eggers, 1930)

Xylosandrus metagermanus (Schedl, 1951a)

Xylosandrus morigerus (Blandford, 1894a)

Xyleborus coffeae Wurth, 1908

Xyleborus difficilis Eggers, 1923

Xyleborus luzonicus Eggers, 1923

Xyleborus abruptoides Schedl, 1955a

Xylosandrus spinifer sp. nov.

Xylosandrus subsimiliformis (Eggers, 1939a)

Xylosandrus subsimilis (Eggers, 1930)

Taxonomic treatment

Key to Xyleborini genera of Southeast Asia (females only)

1	Scutellum not easily visible in dorsal view, apparently absent (Fig. 73A), or conical (Fig. 86A), or narrow, minute and convex (Fig. 67E), or visible only on anterior slope of elytral bases (Fig. 62E)	2
–	Scutellum distinctly visible, linguiform, flush with the elytra, or medially depressed below elytra	13
2	Scutellum conical and surrounded by setae (Fig. 86A)	*Xyleborinus*
–	Scutellum apparently absent (Fig. 73A), or narrow, minute and convex (Fig. 67E), or visible only on anterior slope of elytral bases (Fig. 62E)	3
3	Pronotum with a dense basal mycangial tuft (Fig. 62A); antennal scape long and slender, gradually thickening to apex	*Hadrodemius*
–	Pronotum without a mycangial tuft (Fig. 79G); antennal scape short and thick, or of even thickness	4
4	Mesonotal mycangial tuft in two or four pit mycangia located on the elytra either near the scutellum or along the base (Fig. 39A, C, E), or mycangial tufts absent (Fig. 38G); body elongate with elytral apex attenuate or acuminate	*Cryptoxyleborus*
–	Mesonotal mycangial tuft on elytral bases (Fig. 79G); body stouter with rounded or truncate elytral apex	5
5	Anterior margin of pronotum quadrate or subquadrate, and emarginated; posterior face of protibiae inflated, with or without granules	6
–	Anterior margin of pronotum rounded, never emarginated; posterior face of protibiae flat and unarmed by granules	7
6	Pronotum 1.1–2.0× longer than wide; pronotal disc smooth, finely punctate; antennal funicle 2- or 3-segmented; posterior face of protibiae inflated and unarmed by granules	*Webbia*
–	Pronotum wider than long; pronotal disc coarse, finely asperate; antennal funicle 4-segmented; posterior face of protibiae inflated and granulate	*Schedlia*
7	Declivity truncate, circular, completely surrounded by a circle of pointed teeth	*Pseudowebbia*
–	Declivity not as above if truncate, then not surrounded by a circle of pointed teeth	8
8	Antennal club obliquely truncate, type 2 with one or two sutures visible on posterior face (Fig. 2); pronotal disc punctate	Microperus , in part
–	Antennal club flattened, types 3 or 4 with two or three sutures visible on posterior face (Fig. 3); pronotal disc finely asperate or punctate	9
9	Pronotal disc finely asperate (Fig. 35G)	Coptodryas , in part
–	Pronotal disc punctate (Fig. 36E)	10
10	Antennal club circular	Coptodryas amydra sp. nov.
–	Antennal club longer than wide	Microperus , in part
11	Elytral bases straight (Fig. 67C)	*Microperus fulvulus*
–	Elytral bases bisinuate (Fig. 36E)	12
12	Protibiae distinctly triangular, denticles on apical 1/3 of outer margin	Coptodryas inornata sp. nov.
–	Protibiae semi-circular with evenly rounded outer edge, denticles along most of length or obliquely triangular with denticles on apical half	Microperus , in part
13	Elytral with oblong pit mycangia in distinctly impressed area immediately adjacent to the scutellum on each elytron (Fig. 63C)	*Heteroborips*
–	Elytra without pit mycangia (Fig. 45G)	14
14	Mycangial tuft present on basal margin of pronotum (Fig. 94H) (tuft faint in several species, e.g., Fig. 94B)	15
–	Pronotum without mycangial tufts (Fig. 7F)	18
15	Procoxae widely separated	Xylosandrus , in part
–	Procoxae contiguous or narrowly separated	16
16	Metatibiae conspicuously enlarged and flattened; pronotal disc asperate	*Eccoptopterus*
–	Metatibiae similar to pro- and mesotibiae, never enlarged; pronotal disc punctate	17
17	Lateral margins of pronotum carinate (Fig. 33D)	Cnestus , in part
–	Lateral margins of pronotum obliquely costate (Fig. 22D)	Anisandrus , in part
18	Elytral apex divaricate and ornamented with a pair of distal projections; very elongate, 3.85–4.75× as long as wide	*Streptocranus*
–	Elytral apex entire without a pair of distal projections; stout to elongate, 2.1–3.4× as long as wide	19
19	Posterior face of protibiae inflated, granulate	20
–	Posterior face of protibiae flat, without granules	23
20	Declivital face with three striae (Fig. 8L); antennal club flattened, types 4 or 5 with zero or three sutures on posterior face (Fig. 3)	Amasa , in part
–	Declivital face with five or six striae (Fig. 28C); antennal club obliquely truncate, type 1 or 2 with zero or one suture on posterior face (Fig. 2)	21
21	Elytra with distinctive deep strial furrows and interstrial ridges, ridges either granulate or carinate (Fig. 26E)	Arixyleborus , in part
–	Elytra without strial furrows and interstrial ridges (Fig. 28C)	22
22	Declivital posterolateral margin rounded; lateral profile of declivity appearing obliquely truncate; declivity armed with numerous tubercles; declivital striae 1 variably undulating, never parallel to suture (Fig. 74)	*Stictodex*
–	Declivital posterolateral margin carinate forming a circumdeclivital ring; lateral profile of declivity appearing truncate; declivity unarmed; declivital striae 1 parallel to suture (Fig. 28C, D, J)	*Arixyleborus resecans*
23	Scutellum flush with elytra and medially impressed (Fig. 28G), or depressed below elytra (Fig. 30A)	Arixyleborus , in part
–	Scutellum flush with elytra and flat (Fig. 25C)	24
24	Elytra with distinctive deep strial furrows and interstrial ridges, ridges either granulate or carinate	Arixyleborus , in part
–	Elytra without strial furrows and interstrial ridges	25
25	Anterior margin of pronotum feebly emarginate (Fig. 52A); submentum not depressed below ventral surface of head	*Dryoxylon*
–	Anterior margin of pronotum entire (Fig. 12C); submentum depressed below ventral surface of head (except Ancipitis, some Diuncus)	26
26	Pronotal disc asperate (Fig. 12C), coarsely sculptured	27
–	Pronotal disc punctate (Fig. 47C), finely sculptured	29
27	Anterior margin of pronotum with separate asperities of almost equal size, not larger than those on anterior slope (Fig. 12C)	*Ambrosiodmus*
–	Anterior margin of pronotum with two or more distinctly larger asperities, which may be fused to form a recurved carina (Fig. 31E)	28
28	Protibiae with normal socketed denticles, their bases elevated; declivity distinctly flattened and posterolaterally widened, posterolateral margin costate to interstriae 5; declivital interstriae 2 without spines or tubercles (Fig. 31E)	*Beaverium*
–	Protibiae with denticles reduced or absent, only the raised bases present; declivity either convex with posterolateral margin costate to interstriae 7, or truncate, its margin forming a circular rim around the declivity; spines or tubercles present on declivital interstriae 2 (Fig. 64A)	*Immanus*
29	Elytral apex emarginate and/or explanate (Fig. 48A)	30
–	Elytral apex entire (Fig. 33C)	31
30	Elytra never explanate or excavated	Planiculus , in part
–	Elytra explanate and weakly to strongly excavated (not explanate, strongly excavated and apex appearing subquadrate in D. adusticollis)	*Debus*
31	Lateral margin of pronotum carinate (Fig. 33D)	Cnestus , in part
–	Lateral margin of pronotum obliquely costate (Fig. 7D)	32
32	Procoxae narrowly separated	33
–	Procoxae contiguous	35
33	Elytra truncate; antennal club flattened, types 4 or 5 (Fig. 3)	Amasa , in part
–	Elytra rounded; antennal club obliquely truncate, types 1 or 2 (Fig. 2)	34
34	Declivity unarmed, lacking granules or tubercles (some granules on disc)	*Xylosandrus formosae*
–	Declivity bearing granules or tubercles	Anisandrus , in part
35	Antennal club flattened, types 3, 4 or 5 (Fig. 3)	36
–	Antennal club obliquely truncate, types 1 or 2 (Fig. 2)	46
36	Elytral disc with at least interstrial punctures confused (Fig. 42G)	*Cyclorhipidion*
–	Elytral disc with interstrial punctures uniseriate or interstriae impunctate (Fig. 57E)	37
37	Submentum large, distinctly triangular and flat, flush with genae	*Ancipitis*
–	Submentum variable, slightly or deeply depressed below genae	38
38	Protibiae semi-circular with evenly rounded outer edge	Euwallacea , in part
–	Protibiae obliquely or distinctly triangular	39
39	Anterior margin of pronotum conspicuously extended anteriad with prominent serrations (Fig. 60A)	40
–	Anterior margin of pronotum not conspicuously extended anteriad, without serrations (Fig. 14G)	41
40	Elytral apex rounded; eyes very large, deeply emarginate; elytral apex angulate; larger, 4.8–6.6 mm	*Fortiborus*
–	Elytral apex acuminate; eyes small, feebly emarginate, almost entire; smaller, 3.4–3.5 mm	*Xyleborus bidentatus*
41	Anterior margin of pronotum subquadrate or quadrate in dorsal view (Fig. 4)	42
–	Anterior margin of pronotum conical or rounded in dorsal view (Fig. 4)	43
42	Pronotum wider than long; stouter species, 2.3–2.7× as long as wide	Ambrosiophilus osumiensis , in part
–	Pronotum at least 1.15× longer than wide; elongate species, 2.78–2.89× as long as wide	*Euwallacea semiermis*
43	Elytral apex attenuate, sides parallel in basal 30–60%; declivital slope very gradually rounded; scutellum small	*Tricosa*
–	Elytral apex narrowly or broadly rounded, sides parallel in basal 66–80%; declivital slope evenly or steeply rounded; scutellum large	44
44	Protibiae with six or more socketed denticles	Ambrosiophilus , in part
–	Protibiae with five socketed denticles	45
45	Declivital interstriae unarmed by tubercles or granules	Ambrosiophilus lannaensis sp. nov.
–	Declivital interstriae 2 and 3 each bearing three large tubercles	*Xyleborus singhi*
46	Antennal club 2-segmented, elytra attenuate	*Fraudatrix*
–	Antennal club 3- or 4-segmented, elytra variable but never attenuate	47
47	Antennal club type 1, segment 1 encircling anterior face, no sutures on posterior face (Fig. 2); antennal funicle long and slender; anterior margin of pronotum serrate (absent in D. ciliatoformis)	*Diuncus*
–	Antennal club type 2, with at least one suture on posterior face (Fig. 2); antennal funicle regularly thick or short and thick; anterior margin of pronotum without serrations	48
48	Protibiae semi-circular with evenly rounded outer edge	49
–	Protibiae obliquely or distinctly triangular without evenly rounded edge	52
49	Elytral disc with interstrial punctures confused	50
–	Elytral disc with interstrial punctures uniseriate	51
50	Declivity steeply rounded, posterolateral margin costate and tuberculate	Xyleborus , in part
–	Declivity truncate and encircled by a tuberculate circumdeclivital carina	Truncaudum bullatum sp. nov.
51	Pronotal summit prominent	Euwallacea , in part
–	Pronotal summit low, indistinct	*Truncaudum agnatum*
52	Declivity extremely flat, laterally broadened and densely setose, setae star-shaped scales or bristle-like; declivital slope very gradual	*Leptoxyleborus*
–	Declivity variably convex or slightly broadened and slightly to moderately setose, setae hair-like; declivital slope steep or evenly rounded	53
53	Posterolateral margin of declivity acutely carinate; elytral apex laterally broadened	54
–	Posterolateral margin of declivity rounded or costate; elytral apex variably rounded	55
54	Declivital interstriae 2 armed by tubercles and granules; body unicolored	Xyleborus , in part
–	Declivital interstriae 2 unarmed by tubercles, typically unarmed by granules; body typically bicolored	*Planiculus bicolor*
55	Declivital interstriae 1 laterally broadened from base to declivital midpoint and then narrowing towards apex	Xyleborus , in part
–	Declivital interstriae 1 parallel to suture along its length	56
56	Declivity with tubercles on interstriae 1 and 3 equally sized or those of interstriae 3 the largest	Xyleborus , in part (Xyleborus s. s.)
–	Declivity with tubercles on interstriae 1, 2 and 3 equally sized or those of interstriae 1 the largest	Euwallacea , in part

Amasa Lea, 1894

Amasa Lea, 1894: 322.

Pseudoxyleborus Eggers, 1930: 206. Synonymy: Wood 1984: 223.

Anaxyleborus Wood, 1980: 90. Synonymy: Wood 1983: 647.

Type species

Amasa thoracica Lea, 1894 = Tomicus truncatus Erichson, 1842; monotypy.

Diagnosis

2.5–5.0 mm, 2.11–3.4× as long as wide. Amasa is distinguished by the declivity truncate, margined with a circumdeclivital ring; antennal club flattened, types 4 or 5 (typically type 4), club sutures sinuate, two sutures visible on posterior face; protibiae typically slender, inflated and granulate on posterior face (rarely distinctly triangular or unarmed on posterior face); anterior margin of pronotum with a row of serrations; scutellum flat, flush with elytral surface; declivital face with three striae; procoxae contiguous or narrowly separated; and mycangial tufts absent.

Similar genera

Cyclorhipidion, Pseudowebbia, Truncaudum, Webbia, Xylosandrus.

Distribution

Distributed throughout Asia and Australasia, also occurring in Madagascar. One species has been introduced to Brazil, Chile and Uruguay (Flechtmann and Cognato 2011; Gómez et al. 2017; Kirkendall 2018).

Gallery system

This usually comprises a short radial tunnel leading to a single, large, flat brood chamber, extending in the longitudinal plane.

Remarks

Amasa is easily confused with other species possessing truncate declivities in the genera listed above. Most species can be readily distinguished by the type 4 antennal club with sinuate sutures and the presence of only three striae on the declivital face.

Previous morphological studies of Amasa have suggested that species are very morphologically variable (Hulcr and Cognato 2013). As a result, many species were considered conspecific and part of a morphological continuum. Molecular data generated as part of this study has demonstrated that Amasa species are actually morphologically conserved even across broad ranges (Smith et al. 2020). Amasa species outside our coverage area are thus in need of revision. Potentially much of the diversity is awaiting discovery.

Key to Amasa species (females only)

1	Antennal club type 5, with sutures almost or completely reduced, club covered with pubescence (Fig. 3)	2
–	Antennal club type 4, with sutures visible and partly corneous (Fig. 3)	5
2	Eye completely divided (Fig. 7D); declivity with striae impressed, and all interstriae densely punctate; 4.5 mm	*beesoni*
–	Eye moderately to strongly emarginate (Fig. 7F), other characters variable; 2.0–3.9 mm	3
3	Declivital face with strial and interstrial punctures deeply confused, indistinguishable; larger, 3.7–3.9 mm	*aspersa*
–	Declivital face with three striae clearly indicated on each elytron; smaller, 2.0–3.2 mm	4
4	Stout, 2.0× as long as wide; pronotum from dorsal view round and stout, type 1. 2.0–2.4 mm	*cylindrotomica*
–	Slender, 2.8–3.2× as long as wide; pronotum from dorsal view elongate, type 7. 2.0–3.2 mm	*eugeniae*
5	Declivital striae 2 not equidistant between 1 and 3 (Fig. 10I)	6
–	Declivital striae 2 equidistant between 1 and 3 (Fig. 7K)	7
6	Declivital striae 1 clearly laterally displaced, striae 2 nearly touching striae 1, striae 3 displaced near circumdeclivital margin (Fig. 9I)	lini sp. nov.
–	Declivital striae 2 medially displaced toward striae 1; distance between striae 1 and 3 twice the distance between 1 and 2 (Fig. 10I)	youlii sp. nov.
7	Declivity not granulate, or only interstriae 1 granulate, or only interstriae 1 and 2	8
–	Declivity with all interstriae granulate	10
8	All declivital interstriae smooth, never granulate; larger, 4.5–4.8 mm	*opalescens*
–	Interstriae 1, or 1 and 2 granulate; smaller, 2.9–3.6 mm	9
9	Declivital face flat, strongly shagreened to opalescent; interstriae 1 granulate (typically near apex)	*schlichii*
–	Declivital face convex, strongly shiny; interstriae 1 and 2 moderately inflated from apex to near midpoint of declivity	gibbosa sp. nov.
10	Declivital face setose, sparsely to moderately covered with recumbent or semi-recumbent hair-like setae, sometimes difficult to see	11
–	Declivital face without setae	13
11	Declivity strongly shiny; interstriae very finely setose, setae semi-erect; larger, 4.3–4.5 mm	*concitata*
–	Declivity shagreened, dull; interstriae sparsely to moderately covered with semi-recumbent hair-like setae; smaller, 2.5–3.0 mm	12
12	Setae on declivital interstriae short, less than 1/2 width of an interstria; margin of circumdeclivital ring with short, erect, hair-like setae	galeoderma sp. nov.
–	Setae on declivital interstriae approximately as long as the width of an interstria; margin of circumdeclivital ring with long, erect, bristle-like setae	*versicolor*
13	Declivity strongly shiny; declivital interstriae 1 carinate along at least apical 1/2	tropidacron sp. nov.
–	Declivity shagreened, dull; declivital interstriae 1 granulate	14
14	Declivital interstriae convex; larger, 3.4 mm, and more elongate, 3.4× as long as wide	cycloxyster sp. nov.
–	Declivital interstriae 2–4 flat; smaller, 2.8–3.2 mm, and stouter, 2.3–2.4× as long as wide	*resecta*

Amasa aspersa (Sampson, 1921)

Fig. 7A, B, I

Xyleborus aspersus Sampson, 1921: 31.

Amasa aspersus [sic] (Sampson): Wood and Bright 1992: 682.

Type material

Holotype (NHMUK).

Diagnosis

3.7–3.9 mm long (mean = 3.82 mm; n = 2); 2.11–2.17× as long as wide. This species is distinguished by the dense and strongly confused declivital strial and interstrial punctures with striae and interstriae indistinguishable.

Similar species

None.

Distribution

Brunei, East & West Malaysia, Thailand.

Host plants

All host records are from the genus Eugenia (Myrtaceae), and the species appears to have a fixed host association with this family (Browne 1961b).

Figure 7.

Dorsal, lateral and declivital view of Amasa aspersa, 3.7–3.9 mm (A, B, I), A. beesoni paratype, 5.0 mm (C, D, J), A. concitata, 4.3–4.5 mm (E, F, K), and A. cycloxyster holotype, 3.4 mm (G, H, L).

Amasa beesoni (Eggers, 1930)

Fig. 7C, D, J

Pseudoxyleborus beesoni Eggers, 1930: 207.

Amasa beesoni (Eggers): Wood 1984: 223.

Type material

Holotype (FRI), paratype (NHMW, 1).

Diagnosis

5.0 mm long (n = 1); 2.17× as long as wide. This species is distinguished from all other species in Southeast Asia, except the Malaysian species, A. glauca (Sampson, 1921), by the completely divided eye. It is easily distinguished from A. glauca by the presence of a small tooth on the first interstriae at the top of the declivity, the impressed declivital striae, and densely punctured declivital interstriae.

Similar species

Amasa glauca (from Indomalayan region), A. opalescens.

Distribution

‘Borneo’, West Malaysia, Myanmar, Thailand.

Host plants

The only host records are from the family Sapindaceae (Nephelium, Xerospermum), and the species may have a fixed host association with this family (Browne 1961b).

Amasa concitata (Schedl, 1969)

Fig. 7E, F, K

Xyleborus concitatus Schedl, 1969a: 214.

Amasa concitatus [sic] (Schedl): Wood and Bright 1992: 682.

Type material

Holotype (PPST). Not examined.

New records

China: Jiangxi, Longnan County, Jiulianshan, 24.58; 114.44, 382 m, 1.vii.2018, Lv-Jia, S.C. Lai, ex unknown [host tree] (LYLC, 1).

Diagnosis

4.3–4.5 mm long (n = 2); 2.32–2.5× as long as wide. This species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface smooth, strongly shiny; large size; declivital interstriae very finely setose, setae semi-erect; declivital face convex towards suture; declivital interstriae 1 inflated from apex to near midpoint of declivity; declivital striae 1–3 approximately equidistant.

Similar species

Amasa gibbosa, A. lini, A. tropidacron, A. youlii.

Distribution

China* (Jiangxi), Taiwan.

Host plants

Recorded only from ‘Formosan hardwood’ and ‘angiosperm wood’ (Beaver and Liu 2010).

Amasa cycloxyster sp. nov.

http://zoobank.org/AAE768AB-65F5-4427-91F5-1FA7CEFFC93A

Fig. 7G, H, L

Type material

Holotype , female, Thailand: Surat Thani, Khao Sok National Park, 22.iii.2006, Hulcr et al., ex “Mai Naun Pang” tree (MSUC).

Diagnosis

3.4 mm long (n = 1); 3.4× as long as wide. The species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface shagreened, dull, opaque; declivity glabrous; declivital interstriae 1–3 multiseriate granulate, granules strongly confused; and declivital interstriae convex.

Similar species

Amasa galeoderma, A. resecta, A. schlichii, A. versicolor.

Description

(female). 3.4 mm long (n = 1); 3.4× as long as wide. Body bicolored: pronotum, head, legs, antennae and abdomen orange, elytra dark brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; median impression between eyes; surface shagreened, impunctate, alutaceous, asperate; asperities longitudinal, smaller, rounder, denser above epistoma, increasing in size and length and decreasing in density dorsally and laterally. Eyes very deeply emarginate just above antennal insertion, upper part smaller than lower part. Submentum triangular, deeply impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club approximately circular and flat, type 4; segment 1 corneous, transverse on anterior face, occupying basal 1/5; segment 2 narrow, larger than segment 1, corneous; segments 1–3 present on posterior face. Pronotum: 1.08× as long as wide. In dorsal view basic and parallel-sided, type 2, sides parallel in basal 1/2, rounded anteriorly; anterior margin with a row of five serrations. In lateral view basic, type 0, disc flat, summit at midpoint. Anterior slope shagreened, with densely spaced, fine asperities, becoming lower and more strongly transverse towards summit, bearing long, fine, semi-recumbent, hair-like setae. Disc shiny, alutaceous, impunctate, glabrous. Lateral margins obliquely costate. Base transverse, posterior angles narrowly rounded. Elytra: 1.4× as long as wide, 1.3× as long as pronotum. Scutellum moderately sized, broad, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then sharply angulate to apex. Disc ascending posteriorly, shiny, glabrous; striae and interstriae laterally diverging from base to declivital summit; striae not impressed, punctures separated by 1–4 diameters of a puncture; interstriae flat, finely punctate, punctures 1/2 the size of strial punctures, strongly confused. Declivity truncate, face convex, strongly shagreened, dull, glabrous; three striae present, striae moderately impressed, equidistant, strial punctures shiny, very large, shallow, much larger than on disc, punctures subcontiguous to spaced by two diameters of a puncture; interstriae impunctate, convex, interstriae 1 more strongly convex, interstriae 1–3 multiseriate granulate, granules strongly confused. Posterolateral margin forming a circumdeclivital carina, carina glabrous. Legs: procoxae contiguous; prosternal coxal piece bulging. Protibiae slender, broadest at apical 1/3; posterior face inflated, coarsely granulate; apical 1/2 of outer margin with six small socketed denticles, their length as long as basal width. Meso- and metatibiae broad, flattened, outer margins evenly rounded with 11 small and nine small to minute socketed denticles, respectively; posterior faces unarmed; anterior faces finely granulate.

Etymology

G. kyklos = circle; xyster = rasp. In reference to acute granules on the round declivital face. A noun in apposition.

Distribution

Thailand.

Host plants

Unknown.

Remarks

The holotype specimen is a DNA voucher, SAX40. The head and pronotum were separated from the specimen prior to DNA extraction and point mounted with the elytra.

Amasa cylindrotomica (Schedl, 1939)

Fig. 8A, B, I

Pseudoxyleborus cylindrotomicus Schedl, 1939b: 40.

Xyleborus cylindrotomicus (Schedl): Schedl 1942c: 6.

Xylosandrus cylindrotomicus (Schedl): Wood 1989: 177.

Amasa cylindrotomica (Schedl): Dole and Cognato 2010: 525.

Xyleborus semitruncatus Schedl, 1942c: 35. Synonymy: Schedl 1951a: 79; Wood 1989: 177.

Xyleborus truncatellus Schedl, 1951a: 79. Synonymy: Kalshoven 1959a: 95.

Xyleborus jucundus Schedl, 1954a: 138 (new name for Xyleborus truncatellus Schedl, 1951 nec Schedl 1949). Synonymy: Kalshoven 1959a: 95.

Type material

Lectotype (NHMW). Not examined.

Diagnosis

2.1–2.4 mm long (mean = 2.25 mm; n = 2); 2.0× as long as wide (Sittichaya et al. 2019). This species is distinguished by its minute size, stout form with the pronotum approximately as long as the elytra; declivital surface shagreened, dull, glabrous; and antennal club type 5.

Similar species

Amasa opalescens.

Distribution

Indonesia (Java, Sumatra), Thailand.

Host plants

Recorded from Syzygium aromaticum (Myrtaceae) (Sittichaya et al. 2019).

Figure 8.

Dorsal, lateral and declivital view of Amasa cylindrotomica, 2.1–2.4 mm (A, B, I), A. eugeniae paratype, 2.8–3.2 mm (C, D, J), A. galeoderma holotype, 3.0 mm (E, F, K), and A. gibbosa holotype, 3.5–3.6 mm (G, H, L).

Amasa eugeniae (Eggers, 1930)

Fig. 8C, D, J

Xyleborus eugeniae Eggers, 1930: 183.

Amasa eugeniae (Eggers): Wood and Bright 1992: 683.

Type material

Holotype (FRI), paratypes (FRI, 1; NHMW, 1; NMNH, 1).

Diagnosis

2.8–3.2 mm long (mean = 2.65 mm; n = 5); 2.8–3.2× as long as wide. This species is distinguished by its very elongate body and pronotum (type 7) when viewed dorsally; antennal club type 5; and declivital surface shagreened, dull.

Similar species

Cyclorhipidion amasoides.

Distribution

India (Uttarakhand, West Bengal), Sri Lanka.

Host plants

Recorded from two species of Eugenia (Myrtaceae), and Elaeocarpus (Elaeocarpaceae) (Maiti and Saha 2004).

Amasa galeoderma sp. nov.

http://zoobank.org/77A6AA8D-E16C-4FBD-ADA5-A77AE0CACD50

Fig. 8E, F, K

Type material

Holotype , female, Vietnam: Dong Nai, Cat Tien N.P., 11.44221, 107.43114, 379 m, 20.ii.2017, VN79, A.I. Cognato, T.A. Hoang, ex 4 cm diameter branch (MSUC). Paratypes, female, as holotype (NHMW, 1; NHMUK, 1; NMNH, 1; VMNH, 1).

Diagnosis

3.0 mm long (mean = 3.0 mm; n = 5); 2.5× as long as wide. This species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface shagreened, dull, opaque; declivital interstriae granulate, granules multiseriate, confused; declivital interstriae 1 moderately covered with semi-recumbent fine hair-like setae, less than 1/2 width of an interstria; and circumdeclivital carina margin setose, setae short, erect, hair-like.

Similar species

Amasa cycloxyster, A. resecta, A. schlichii, A. versicolor.

Description

(female). 3.0 mm long (mean = 3.0 mm; n = 5); 2.5× as long as wide. Body bicolored: pronotum, head, legs and antennae orange, elytra and abdomen dark brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; median impression between eyes; surface shagreened, impunctate, alutaceous, asperate; asperities longitudinal, smaller, rounder, denser above epistoma, increasing in size and length and decreasing in density dorsally and laterally. Eyes very deeply emarginate just above antennal insertion, upper part smaller than lower part. Submentum triangular, deeply impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club approximately circular and flat, type 4; segment corneous, 1 convex on anterior face, occupying approximately basal 1/4; segment 2 narrow, larger than segment 1, corneous; segments 1–3 present on posterior face. Pronotum: 1.0× as long as wide. In dorsal view basic and parallel-sided, type 2, sides parallel in basal 1/2, rounded anteriorly; anterior margin with a row of 5–7 serrations. In lateral view basic, type 0, disc flat, summit at midpoint. Anterior slope strongly shiny, with widely spaced, moderate asperities, becoming lower and more strongly transverse towards summit, bearing long, fine, semi-recumbent, hair-like setae. Disc shiny, alutaceous, sparsely finely punctate, glabrous. Lateral margins obliquely costate. Base transverse, posterior angles narrowly rounded. Elytra: 1.35× as long as wide, 1.25× as long as pronotum. Scutellum moderately sized, broad, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then sharply angulate to apex. Disc ascending posteriorly, shiny, glabrous; striae and interstriae laterally diverging from base to declivital summit; striae not impressed, punctures separated by 2–3 diameters of a puncture; interstriae flat, finely uniseriate punctate, punctures 1/3 size of strial punctures. Declivity truncate, face flattened, strongly shagreened, dull, glabrous; three striae present, striae moderately impressed, striae 2 equidistant between striae 1 and 3, strial punctures shiny, very large, shallow, much larger than on disc, punctures subcontiguous; interstriae impunctate, convex, interstriae 1 more strongly convex, interstriae 1–3 multiseriate granulate, granules multiseriate, confused, interstriae 1 moderately covered with fine, semi-recumbent, hair-like setae, less than 1/2 width of an interstria. Posterolateral margin forming a circumdeclivital carina; carina setose, setae short, erect, hair-like. Legs: procoxae contiguous, prosternal coxal piece flat, inconspicuous. Protibiae slender, broadest at apical 1/3; posterior face inflated, finely granulate; apical 1/2 of outer margin with six small socketed denticles, their length as long as basal width. Meso- and metatibiae broad, flattened, outer margins evenly rounded with nine and 11 small socketed denticles, respectively, posterior faces unarmed; anterior faces finely granulate.

Etymology

G. galeos = shark; derma = skin. In reference to the shagreened face of the declivity. Noun in apposition.

Distribution

Vietnam.

Host plants

Unknown.

Amasa gibbosa sp. nov.

http://zoobank.org/359F611F-95A0-4631-94B6-67F1A8DC5BCD

Fig. 8G, H, L

Type material

Holotype , female, Thailand: Kanchanaburi, Thong Pha Phoom Dist., Phu Yae subdist[rict], 400 m, 14.944N, 98.674E, 16.vii.2002, Cognato, Gillogly, Harlin (MSUC). Paratypes, female, as holotype (MSUC, 1; NHMUK, 1; RABC, 1); Suratthani, Khao Sok N.P., 1.ii.2015, 19°21'41.8"N, 98°55'03.4"E, W. Sittichaya, ex ethanol baited trap, tropical rain forest (MSUC, 1).

Diagnosis

3.5–3.6 mm long (mean = 3.53 mm; n = 3); 2.33–2.41× as long as wide. This species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface glabrous, smooth, strongly shiny; moderate size; declivital face convex, interstriae 1 and 2 moderately inflated from apex to near midpoint of declivity; declivital striae 1–3 approximately equidistant.

Similar species

Amasa concitata, A. lini, A. tropidacron, A. youlii.

Description

(female). 3.5–3.6 mm long (mean = 3.53 mm; n = 3); 2.33–2.41× as long as wide. Body dark red-brown. Legs and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; median impression between eyes; surface shagreened, impunctate, alutaceous, asperate; asperities longitudinal, smaller, rounder, denser above epistoma, increasing in length and decreasing in width and density dorsally. Eyes very deeply emarginate just above antennal insertion, upper part smaller than lower part. Submentum triangular, deeply impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club approximately circular and flat, type 4; segment 1 corneous, convex on anterior face, occupying approximately basal 1/4; segment 2 broad, larger than segment 1, corneous; segments 1–3 present on posterior face. Pronotum: 1.02× as long as wide. In dorsal view basic and parallel-sided, type 2, sides parallel in basal 1/2, rounded anteriorly; anterior margin with a row of six serrations. In lateral view basic, type 0, disc flat, summit at midpoint. Anterior slope strongly shiny with densely spaced, fine asperities, becoming lower and more strongly transverse towards summit, bearing long, fine, semi-recumbent, hair-like setae. Disc shiny, alutaceous, densely finely punctate behind summit, punctures decreasing in density toward base, glabrous. Lateral margins obliquely costate. Base transverse, posterior angles narrowly rounded. Elytra: 1.48× as long as wide, 1.45× as long as pronotum. Scutellum moderately sized, broad, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then sharply angulate to apex. Disc ascending posteriorly, shiny, glabrous; striae and interstriae laterally diverging from base to declivital summit; striae not impressed, punctures separated by five diameters of a puncture; interstriae flat, finely punctate, punctures 1/2 size of strial punctures, strongly confused. Declivity truncate, face convex, strongly shiny, smooth, glabrous; three striae present, striae weakly impressed, striae 2 equidistant between striae 1 and 3, strial punctures subshiny, very large and deep, much larger and deeper than on disc, punctures subcontiguous to spaced by one diameter of a puncture; interstriae impunctate, convex, interstriae 1 and 2 moderately inflated from apex to near midpoint of declivity; apical 1/4 of interstriae 1 and 2 with a row of uniseriate rugae. Posterolateral margin forming a circumdeclivital carina; carina setose, setae short, erect hair-like. Legs: procoxae contiguous; prosternal coxal piece flat, inconspicuous. Protibiae distinctly triangular, broadest at apical 1/3; posterior face inflated, coarsely granulate; apical 1/2 of outer margin with six or seven small socketed denticles, their length as long as basal width. Meso- and metatibiae broad, flattened; outer margins evenly rounded with 11 and nine small to minute socketed denticles, respectively; posterior faces unarmed; anterior faces finely granulate.

Etymology

L. gibbosa = humped. In reference to the rather bulging declivity. A variable adjective.

Distribution

Thailand.

Host plants

Unknown.

Amasa lini sp. nov.

http://zoobank.org/AE746EB3-4A92-4977-BD5B-DF3BC45FE625

Fig. 9A, B, I

Type material

Holotype , female, Taiwan: Nantou Dist., Sun Moon Lake, 23.vi.2016, C.-S. Lin (TARI).

Diagnosis

3.5 mm long (n = 1); 2.33× as long as wide. This species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface smooth, shiny; large size; declivity glabrous; declivital interstriae 1 strongly tumescent and granulate; declivital striae 1 strongly laterally displaced, nearly touching striae 2, striae 3 displaced to near circumdeclivital carina margin; and declivital striae 2 not appearing equidistant between striae 1 and 3.

Similar species

Amasa concitata, A. gibbosa, A. tropidacron, A. youlii.

Description

(female). 3.5 mm long (n = 1); 2.33× as long as wide. Body bicolored: pronotum reddish, elytra and abdomen dark brown, head, legs, and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; median impression between eyes; surface shagreened, impunctate, alutaceous, asperate; asperities longitudinal, larger, rounder, denser above epistoma, increasing in length and decreasing in width and density dorsally. Eyes very deeply emarginate just above antennal insertion, upper part smaller than lower part. Submentum triangular, deeply impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club approximately circular and flat, type 4; segment 1 corneous, sinuate on anterior face, occupying approximately 1/5 of club; segment 2 narrow, larger than segment 1, corneous; segments 1–3 present on posterior face. Pronotum: 1.4 × as long as wide. In dorsal view basic and parallel-sided, type 2, sides parallel in basal 1/2, rounded anteriorly; anterior margin with a row of eight serrations. In lateral view basic, type 0, disc flat, summit at midpoint. Anterior slope shagreened, with densely spaced, fine asperities, becoming lower and more strongly transverse towards summit, bearing long, fine, semi-recumbent hair-like setae. Disc subshiny, alutaceous, densely, finely punctate, glabrous. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Elytra: 1.4× as long as wide, 1.43× as long as pronotum. Scutellum moderately sized, broad, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then sharply angulate to apex. Disc flat, shiny, glabrous; striae and interstriae laterally diverging from base to declivital summit; striae not impressed, punctures separated by 3–5 diameters of a puncture; interstriae flat, finely punctate, punctures 1/3 size of strial punctures, strongly confused. Declivity truncate, face convex, smooth, shiny, glabrous; three striae present, striae weakly impressed, striae 1 strongly laterally displaced, striae 2 nearly touching striae 1, striae 3 displaced to near circumdeclivital carina, strial punctures dull, small, shallow, larger than on disc, punctures spaced by a diameter of a puncture; interstriae impunctate, convex, interstriae 1 strongly tumescent and granulate, granules strongly confused, apical 1/6 of interstriae 1 carinate. Posterolateral margin forming a circumdeclivital carina; carina glabrous. Legs: procoxae contiguous, prosternal coxal piece flat, inconspicuous. Protibiae slender, broadest at apical 1/3; posterior face inflated, finely granulate; apical 1/2 of outer margin with five small socketed denticles, their length as long as basal width. Meso- and metatibiae broad, flattened, outer margins evenly rounded with 11 and nine small socketed denticles, respectively, posterior faces unarmed; anterior faces finely granulate.

Etymology

The species is named for Mr. Ching-Shan Lin, the collector, for his contributions to our knowledge of bark and ambrosia beetles. Noun in genitive.

Distribution

Taiwan.

Host plants

Unknown.

Figure 9.

Dorsal, lateral and declivital view of Amasa lini holotype, 3.5 mm (A, B, I), A. opalescens lectotype, 4.5–4.8 mm (C, D, J), A. resecta, 2.85–3.2 mm (E, F, K), and A. schlichii, 2.9–3.5 mm (G, H, L).

Amasa opalescens (Schedl, 1937)

Fig. 9C, D, J

Xyleborus opalescens Schedl, 1937a: 550.

Amasa opalescens (Schedl): Wood and Bright 1992: 684.

Type material

Lectotype (NHMW).

Diagnosis

4.5–4.8 mm long (4.7 mm long; n = 3); 2.4–2.5× as long as wide. This species is distinguished by its large size; pronotum rounded, robust from lateral view (type 5); declivital interstriae 1 unarmed (lacking granules) and flat; declivital strial punctures very large, irregularly spaced; and declivital surface appearing smooth and opalescent.

Similar species

Amasa beesoni, A. cylindrotomica, A. schlichii.

Distribution

East & West Malaysia, Thailand, Vietnam.

Host plants

Recorded only from species of Eugenia and Tristania (Myrtaceae), and possibly with a fixed association with this family (Browne 1961b).

Amasa resecta (Eggers, 1923)

Fig. 9E, F, K

Xyleborus abruptus Eggers, 1923: 169.

Xyleborus resectus Eggers, 1927a: 391 (new name for X. abruptus Eggers, 1923 nec Sampson 1914).

Amasa resectus [sic] (Eggers): Wood and Bright 1992: 684.

Xyleborus opacicauda Eggers, 1940: 136. Synonymy: Kalshoven 1959b: 159.

Type material

Syntype Xyleborus resectus (MIZ, 1).

New records

China: Hainan, Wu-zhi-shan Town, 18.902N, 109.663E, 703 m, 2.xii.2016, Tian-Shang, Lv-Jia (RABC, 2).

Diagnosis

2.85–3.2 mm long (mean = 2.94 mm; n = 4); 2.29–2.38× as long as wide. This species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface shagreened, dull, opaque; declivity glabrous; declivital interstriae 1–3 multiseriate granulate, granules strongly confused; and declivital interstriae 2–4 flat.

Similar species

Amasa cycloxyster, A. galeoderma, A. schlichii, A. versicolor, A. youlii.

Distribution

China (Hainan), Indonesia (Java, Sumatra), East Malaysia, New Guinea, Sri Lanka, Thailand.

Host plants

Recorded by Kalshoven (1959b) from five genera in five different families. Evidently polyphagous.

Remarks

Hulcr and Cognato (2013) synonymized Xyleborus fulgens Schedl, 1975c with this species, but we believe it to be distinct. Hence it is not included in the list of synonyms.

Amasa schlichii (Stebbing, 1907)

Fig. 9G, H, L

Acanthotomicus truncatus Stebbing, 1907: 40.

Xyleborus schlichii Stebbing, 1914: 592 (new name for Xyleborus (Acanthotomicus) truncatus (Stebbing, 1907) nec Erichson 1842).

Amasa schlichi [sic] (Stebbing): Wood 1989: 169.

Xyleborus glaber Eggers, 1930: 185. Synonymy: Wood 1989: 169.

Xyleborus uniseriatus Eggers, 1936b: 89. Synonymy: Schedl 1963b: 268.

Xyleborus verax Schedl, 1939b: 43. Synonymy: Kalshoven 1959a: 95.

Type material

Holotype , Xyleborus glaber (FRI), paratype (NHMW, 1). Syntype Xyleborus schlichii (FRI, 1).

New records

China: Hong Kong, Tai Po Kau, vi.2017, J. Skelton (MSUC, 1). S-Yunnan, Xishuangbanna, Sanchahe Nat. Res., 22°09.784'N, 100°52.256'E, 2186 m, 29–30.v.2008, A.I. Cognato (MSUC, 2); as previous except: 23 km NW Jinghong, vic. Na Ban village (NNNR), 22°10'N, 100°39'E, 700–1000 m, v–vii. 2009, L. Meng (RABC, 2). Japan: Okinawa Pref., Iriomote-jima Island, 26.vi.2016, H. Kajimura, ex Machilus thunbergii (MSUC, 1). Vietnam: Cao Bang, 22°33.9981'N, 105°52.591'E, 1051 m, 12–17.iv.2014, VN11, Cognato, Smith, Pham, ex FIT (MSUC, 3). N. Ninh Binh, 90 km SW Hanoi, Cuc Phuong N.P., primate rescue centre, 20°14'24"N, 105°42'53"E, 190 m, 25.iv.2012, A. Weigel, ex light trap (NKME, 1). Thua Thien-Hue, Bach Ma N.P., 16.22897, 107.85349, 415 m, 15.ii.2017, VN61, A.I. Cognato, T.A. Hoang, ex 5 cm diameter branch (MSUC, 1).

Diagnosis

2.9–3.5 mm long (mean = 3.21 mm; n = 10); 2.23–2.54× as long as wide. This species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface shagreened to opalescent, dull, opaque; declivity glabrous; and declivital interstriae 1 granulate (typically near apex), interstriae 2 and 3 unarmed.

Similar species

Amasa cycloxyster, A. galeoderma, A. resecta, A. versicolor, A. youlii.

Distribution

China* (Hong Kong, Yunnan), India (Assam, West Bengal), Indonesia (Java), Japan*, East & West Malaysia, Thailand, Vietnam*.

Host plants

Apparently polyphagous (Beeson 1961; Beaver and Browne 1979; Maiti and Saha 2004).

Remarks

This species had previously been considered to be extremely morphologically variable (Hulcr and Cognato 2013) but Cognato et al. (2020b) and Smith et al. (2020) demonstrated that very little intraspecific morphological variation is present and removed the Papua New Guinean species A. striatotruncata (Schedl, 1936) and A. umbratula (Schedl, 1975) from synonymy.

Wood (1989: 169) considered Xyleborus glaber to be a synonym of X. schlichii. Beaver et al. (2014: 20) later considered it to be a distinct species. Upon our examination of the photos of the holotype and a paratype specimen we found this species to be conspecific with Amasa schlichii and it is here returned to synonymy.

Amasa tropidacron sp. nov.

http://zoobank.org/D6F42632-2E07-47D3-BBDA-8C97934C7E12

Fig. 10A, B, G

Type material

Holotype , female, Japan: Okinawa, Iriomote-jima, Isd. Code. 1, 9.xi.2012, Kajimura (MSUC). Paratypes, female, as holotype (MSUC, 1); as previous except: Yona, 1.xi.2010, J. Hulcr, ex Castanopsis, uffeID 7348 (UFFE, 2), uffeID 7389 (UFFE, 4); Vietnam: Ninh Binh, Cuc Phuong N.P., Mac Lake, 20°15'29.0"N, 105°42'27.5"E, 155 m, 4–7.v.2009, J.B. Heppner, ex blacklight trap (FSCA, 1).

Diagnosis

2.5–2.8 mm long (mean = 2.65 mm; n = 2); 2.5–2.54× as long as wide. This species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface glabrous, smooth, strongly shiny; small size; declivital face flattened; and interstriae 1 carinate, weakly inflated from apex to near midpoint of declivity; declivital striae 1–3 approximately equidistant.

Similar species

Amasa concitata, A. gibbosa, A. lini, A. youlii.

Description

(female). 2.5–2.8 mm long (mean = 2.65 mm; n = 2); 2.5–2.54× as long as wide. Body light red-brown. Head, legs, and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; median impression between eyes; surface shagreened, impunctate, alutaceous, asperate; asperities longitudinal, smaller, rounder, denser above epistoma, increasing in length and decreasing in width and density dorsally and laterally. Eyes deeply emarginate just above antennal insertion, upper part smaller than lower part. Submentum triangular, deeply impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club approximately circular and flat, type 4; segment 1 corneous, convex on anterior face, occupying approximately basal 1/5; segment 2 broad, larger than segment 1, corneous; segments 1–3 present on posterior face. Pronotum: 1.13× as long as wide. In dorsal view basic, type 2, sides parallel in basal 1/2, rounded anteriorly; anterior margin with a row of 6–8 serrations. In lateral view basic, type 0, disc flat, summit at midpoint. Anterior slope shagreened with densely spaced, fine asperities, becoming lower and more strongly transverse towards summit; bearing long, fine, semi-recumbent hair-like setae. Disc shiny, alutaceous, impunctate, glabrous. Lateral margins obliquely costate. Base transverse, posterior angles narrowly rounded. Elytra: 1.4× as long as wide, 1.23× as long as pronotum. Scutellum moderately sized, broad, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then sharply angulate to apex. Disc ascending posteriorly, shiny, glabrous; striae and interstriae laterally diverging from base to declivital summit; striae not impressed, punctures separated by 1–4 diameters of a puncture; interstriae flat, finely punctate, punctures 1/5 size of strial punctures, strongly confused. Declivity truncate, face flattened, strongly shiny, smooth, glabrous; three striae present, striae weakly impressed, equidistant, strial punctures strongly shiny, very large, deep, much larger and deeper than on disc, punctures subcontiguous to spaced by one diameter of a puncture; interstriae impunctate, convex, interstriae 1 weakly inflated from apex to below declivital midpoint, interstriae 1 uniseriate granulate, 2–4 multiseriate granulate, granules strongly confused; apical 1/4 of interstriae 1 and 2 costate with a row of rugae. Posterolateral margin forming a circumdeclivital carina; carina setose, setae short, erect, hair-like. Legs: procoxae contiguous; prosternal coxal piece flat, inconspicuous. Protibiae slender, broadest at apical 1/3; posterior face inflated, finely granulate; apical 1/2 of outer margin with five small socketed denticles, their length as long as basal width. Meso- and metatibiae broad, flattened, outer margins evenly rounded with 11 and ten small socketed denticles, respectively; posterior faces unarmed; anterior faces finely granulate.

Etymology

G. tropis = keel, ridge; akron = end. In reference to the inflated costate apex of the declivity. Noun in apposition.

Distribution

Japan, Vietnam.

Host plants

This species has been recorded from Castanopsis (Fagaceae).

Figure 10.

Dorsal, lateral and declivital view of Amasa tropidacron holotype, 2.5–2.8 mm (A, B, G), A. versicolor, 2.5–2.6 mm (C, D, H), and A. youlii holotype, 2.9–3.0 mm (E, F, I).

Amasa versicolor (Sampson, 1921)

Fig. 10C, D, H

Xyleborus versicolor Sampson, 1921: 29.

Amasa versicolor (Sampson): Wood and Bright 1992: 685.

Type material

Holotype (NHMUK), allotype (NHMUK).

New records

Ceylon [Sri Lanka]: Kalutara Dist., Kanneliya, 250 m, 23.v.1973, S.L. Wood, ex limbs (NMNH, 1); Morapitiya, 250 m, 27.v.1973, S.L. Wood (NMNH, 2).

Diagnosis

2.5–2.6 mm long (mean = 2.57 mm; n = 5); 2.27–2.43× as long as wide. The species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface shagreened, dull, opaque; declivital interstriae granulate, granules multiseriate, confused; declivity setose, interstriae moderately covered with semi-recumbent hair-like setae, approximately as long as the width of an interstria; and circumdeclivital carina margin setose, setae long, erect, bristle-like.

Similar species

Amasa cycloxyster, A. galeoderma, A. resecta, A. schlichii, A. youlii.

Distribution

Federated States of Micronesia, India (‘Bengal’), Indonesia (Java), East & West Malaysia, Myanmar, Sri Lanka*, Thailand.

Host plants

Polyphagous (Browne 1961b; Beaver and Browne 1979).

Amasa youlii sp. nov.

http://zoobank.org/5AAB68A6-B0EC-46B6-A0E8-D3A649A4B11C

Fig. 10E, F, I

Type material

Holotype , female, China: Fujian, Fuzhou, Qishan, 31.iii.2018, Y. Li, ex 5 cm diameter twig, possibly Fagaceae (IZAS). Paratypes, female, as holotype (MSUC, 1; NMNH, 1)

Diagnosis

2.9–3.0 mm long (mean = 2.93 mm; n = 3); 2.42–2.5× as long as wide. This species is distinguished by the pronotum appearing basic (type 2) when viewed dorsally, anterior margin serrate; declivital surface smooth, moderately shiny; small size; declivital interstriae setose, setae recumbent; declivital face flattened; and interstriae 1 weakly inflated from apex to near midpoint of declivity; and declivital striae 2 medially displaced, not appearing equidistant between striae 1 and 3.

Similar species

Amasa concitata, A. gibbosa, A. lini, A. tropidacron.

Description

(female). 2.9–3.0 mm long (mean = 2.93 mm; n = 3); 2.42–2.5 × as long as wide. Body bicolored: pronotal disc, head, legs, and antennae reddish, anterior slope of pronotum, elytra, and abdomen dark brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; median impression between eyes; surface shagreened, impunctate, alutaceous, asperate; asperities longitudinal, smaller, rounder, denser above epistoma, increasing in size and length and decreasing in density dorsally and laterally. Eyes deeply emarginate just above antennal insertion, upper part smaller than lower part. Submentum triangular, deeply impressed. Antennal scape regularly thick, longer than club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club approximately circular and flat, type 4; segment 1 corneous, sinuate on anterior face, occupying approximately basal 1/4; segment 2 narrow, larger than segment 1, corneous; segments 1–3 present on posterior face. Pronotum: 0.88× as long as wide. In dorsal view basic and parallel-sided, type 2, sides parallel in basal 1/2, rounded anteriorly; anterior margin with a row of 4–6 serrations. In lateral view basic, type 0, disc flat, summit at midpoint. Anterior slope strongly shagreened with densely spaced, short fine asperities, becoming lower and more strongly transverse towards summit, bearing long, fine, semi-recumbent hair-like setae. Disc shiny, alutaceous, densely minutely punctate, glabrous. Lateral margins obliquely costate. Base transverse, posterior angles narrowly rounded. Elytra: 1.45× as long as wide, 1.65× as long as pronotum. Scutellum moderately sized, broad, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then sharply angulate to apex. Disc flat, shiny, glabrous; striae and interstriae laterally diverging from base to declivital summit; striae not impressed, punctures separated by 1–4 diameters of a puncture; interstriae flat, finely punctate, punctures 1/2 size of strial punctures, strongly confused. Declivity truncate, face flattened, moderately shiny, smooth, setose; three striae present, striae weakly impressed, striae 2 medially displaced near striae 1, strial punctures shiny, moderately large, moderately deep, much larger than on disc, punctures subcontiguous to spaced by three diameters of a puncture; interstriae impunctate, convex, interstriae 1 moderately inflated from apex to above declivital midpoint, interstriae 1 uniseriate granulate, 2–4 multiseriate granulate, granules strongly confused; apical 1/2 of interstriae 1 carinate to just before apex, becoming flattened, apical 1/4 of interstriae 2 costate, nearly carinate, with a row of rugae. Posterolateral margin forming a circumdeclivital carina; carina setose, setae short, erect hair-like. Legs: procoxae contiguous, prosternal coxal piece flat, inconspicuous. Protibiae slender, broadest at apical 1/3; posterior face inflated, coarsely granulate; apical 1/2 of outer margin with six small socketed denticles, their length as long as basal width. Meso- and metatibiae broad, flattened, outer margins evenly rounded with 11 and nine small socketed denticles, respectively, posterior faces unarmed; anterior faces finely granulate.

Etymology

Named after the collector Dr. You Li for his generous contributions to this project. Noun in genitive, invariable.

Distribution

China (Fujian).

Host plants

Unknown but potentially collected from Fagaceae.

Ambrosiodmus Hopkins, 1915

Ambrosiodmus Hopkins, 1915a: 55.

Phloeotrogus Motschulsky, 1863: 512. Wood 1969: 113.

Brownia Nunberg, 1963: 37. Synonymy: Wood 1980: 96.

Type species

Xyleborus tachygraphus Zimmerman, 1868; original designation.

Diagnosis

2.5–4.8 mm, 1.7–2.8× as long as wide, body usually stout and darkly colored. Ambrosiodmus is distinguished by the pronotum short and rounded, types 1 or 2 in dorsal view; pronotal disc entirely asperate; pronotum anterior margin without a carina or serrations; elytral disc convex; declivity rounded and steep at apex; antennal club flattened, type 4; scutellum flat, flush with elytra; mycangial tufts absent; and procoxae contiguous.

Similar genera

Ambrosiophilus, Beaverium, Immanus.

Distribution

Temperate and tropical regions of the world.

Gallery system

This consists of a radial entrance tunnel leading to branched tunnels. These usually lie predominantly in one horizontal plane but may extend into three dimensions. They lack enlarged brood chambers. Many gallery systems are often started in a small area of the tree. Unlike many xyleborines, the galleries of different individuals often interconnect so that beetles can move between galleries (Beeson 1961; Kasson et al. 2016).

Remarks

Recent studies suggest that all Ambrosiodmus and Ambrosiophilus species (see below) are associated with a single species of polypore basidiomycete ambrosia fungus (Flavodon ambrosius) (Kasson et al. 2016; Li et al. 2017). This fungus has greater ability to break down lignocellulose than most ambrosia fungi. This enables the beetles to colonize wood at a more advanced state of decay than most ambrosia beetles, and to persist in the same tree over several generations (Kasson et al. 2016; Li et al. 2017).

Key to Ambrosiodmus species (females only)

1	Declivity granulate (Fig. 12F)	2
–	Declivity tuberculate or denticulate, never granulate (Fig. 12E)	4
2	Declivital interstriae with uniformly sized and spaced granules from base to apex; declivital interstriae bearing erect hair-like setae	*rubricollis*
–	Declivity with uniformly sized and spaced granules on declivital interstriae from base to declivity midpoint, apical 1/2 of interstriae with granules irregularly spaced; declivital interstriae slightly elevated and bearing erect thick setae	3
3	Larger, 3.2–3.4 mm; apical 1/2 of declivital interstriae 1 with five or six granules	*brunneipes*
–	Smaller, 2.9–3.1 mm; apical 1/2 of declivital interstriae 1 with three or four granules	*conspectus*
4	Declivital interstriae tuberculate, except interstriae 1 unarmed (rarely a few granules in some individuals); smaller, 2.5–2.8 mm	*asperatus*
–	All declivital interstriae tuberculate; larger, 3.4–4.8 mm	5
5	Tubercles of declivital interstriae 2 distinctly larger than those of other interstriae (Fig. 11L); usually larger, 3.4–4.8 mm	*lewisi*
–	Tubercles of declivital interstriae 2 similarly sized to those of other interstriae (Fig. 12E); usually smaller, 3.5–4.0 mm	*minor*

Ambrosiodmus asperatus (Blandford, 1895)

Fig. 11A, B, I

Xyleborus asperatus Blandford, 1895: 321.

Ambrosiodmus asperatus (Blandford): Wood 1989: 169.

Xyleborus nepotulus Eggers, 1923: 179. Synonymy: Schedl 1958c: 151.

Xyleborus citri Beeson, 1930: 215. Synonymy: Wood 1989: 169.

Xyleborus nepotulomorphus Eggers, 1936b: 88. Synonymy: Schedl 1958c: 151.

Type material

Holotype Xyleborus asperatus (NHMUK). Paratype Xyleborus nepotulomorphus (MFNB).

New records

China: Guangxi, Shiwandashan, 25.iii.2018, Y. Li, ex Quercus griffithii (UFFE, 1). Hong Kong, Tai Po Kau, vi.2017, J. Skelton (MSUC, 1). Japan: South-western Japan, Okinawa, Iriomote-jima Island, H. Kajimura, ex Machilus thunbergii tree (MSUC, 1). Vietnam: Thua Thien-Hue, Bach Ma N.P., 16.22897, 107.85349, 415 m, 15.ii.2017, VN57, A.I. Cognato, T.A. Hoang, ex 5 cm diameter branch; twig (MSUC, 1).

Diagnosis

2.5–2.8 mm long (mean = 2.64 mm; n = 5); 2.4–2.8× as long as wide. This species is distinguished by declivital interstriae 2 bearing a row of 3–5 denticles that are larger than those on other interstriae, and declivital interstriae 1 distinctly impressed.

Similar species

Ambrosiophilus cristatulus, A. osumiensis, A. subnepotulus.

Distribution

Australia, Brunei, China (Guizhou, Guangxi*, Hainan, Hong Kong*, Xizang), India (Tamil Nadu, West Bengal), Indonesia (Java, Sulawesi, Sumatra), Japan (Ryukyu Is), West Malaysia, Nepal, New Guinea, Sri Lanka, Taiwan, Thailand, Vietnam*.

Host plants

Polyphagous (Beaver and Liu 2010).

Remarks

This species has a very similar appearance and size to several Ambrosiophilus species which also have three or four denticles on declivital interstriae 2. The two genera are easily separated by the pronotal disc sculpturing: punctate in Ambrosiophilus and asperate in Ambrosiodmus.

Figure 11.

Dorsal, lateral and declivital view of Ambrosiodmus asperatus, 2.5–2.8 mm (A, B, I), A. brunneipes, 3.2–3.4 mm (C, D, J), A. conspectus paratype, 2.9–3.1 mm (E, F, K), and A. lewisi, 3.4–4.8 mm (G, H, L).

Ambrosiodmus brunneipes (Eggers, 1940)

Fig. 11C, D, J

Xyleborus brunneipes Eggers, 1940: 138.

Ambrosiodmus brunneipes (Eggers): Wood and Bright 1992: 671.

Type material

Allotype (NHMW).

Diagnosis

3.2–3.4 mm long (mean = 3.38 mm; n = 5); 2.43–2.5× as long as wide. This species is distinguished by the declivital interstriae with uniformly sized and spaced granules from base to declivital midpoint, apical 1/2 of interstriae with granules irregularly spaced; declivital interstriae slightly elevated and bearing thick, erect setae, setae located ventrad of granules; declivital surface strongly shagreened; and dark brown color.

This species is very closely related to A. conspectus and is distinguished by the larger size and five or six granules on the apical 1/2 of declivital interstriae 1.

Similar species

Ambrosiodmus conspectus, A. rubricollis.

Distribution

Indonesia (Java), East & West Malaysia, Thailand.

Host plants

Recorded from Parartocarpus (Moraceae), Octomeles (Tetramelaceae), and rattans (Arecacae). Probably polyphagous (Beaver et al. 2014).

Ambrosiodmus conspectus (Schedl, 1964)

Fig. 11E, F, K

Xyleborus conspectus Schedl, 1964b: 247.

Ambrosiodmus conspectus (Schedl): Wood and Bright 1992: 672.

Type material

Paratypes (NHMW, 2).

Diagnosis

2.9–3.1 mm long (mean = 3.01 mm; n = 5); 2.48–2.73× as long as wide. This species is distinguished by declivity with uniformly sized and spaced granules on declivital interstriae from base to declivity midpoint, apical 1/2 of interstriae with granules irregularly spaced; declivital interstriae slightly elevated and bearing thick, erect setae, setae located ventrad of granules; declivital surface strongly shagreened; and dark brown color.

This species is very closely related to A. brunneipes and is distinguished by the smaller size and the and three or four granules on the apical 1/2 of declivital interstriae 1.

Similar species

Ambrosiodmus brunneipes, A. rubricollis.

Distribution

East Malaysia, Thailand.

Host plants

Recorded only from rattan (Arecacae) (Schedl 1964b).

Ambrosiodmus lewisi (Blandford, 1894)

Fig. 11G, H, L

Xyleborus lewisi Blandford, 1894b: 104.

Ambrosiodmus lewisi (Blandford): Wood 1989: 170.

Ozopemon tuberculatus Strohmeyer, 1912: 38. Synonymy: Beaver and Liu 2010: 20.

Xyleborus lewekianus Eggers, 1923: 181. Synonymy: Wood 1989: 170.

Xyleborus tegalensis Eggers, 1923: 181. Synonymy: Schedl 1962a: 208.

Type material

Syntypes Xyleborus lewisi (NHMUK). Syntypes Ozopemon tuberculatus (SDEI).

New records

China: Hong Kong, Sheung Shui, 22.vi.1964, ex soaked in oil (BPBM, 1); Tai Po Kau, 23.ix.1965, Lee Kit Ming, Hui Wai Ming, ex light trap (BPBM, 1), as previous except: 30.vi.1964, (BPBM, 1), as previous except: 2–6.vii.1964 (BPBM, 1), as previous except: 3–4.vii.1965 (BPBM, 1). India: Arunachal Pradesh, Hunli vicinity, 28°19'32"N, 95°57'31"E, 1300±100 m, 26.v.2012, L. Dembický (ZFMK, 1). Vietnam: Cao Bang, Phia Oac Hotel, 22°37.702'N, 105°54.5467'E, 847 m, 10–17.iv.2014, VN1, Cognato, Smith, Pham, ex in flight (MSUC, 1). Lao Cai, pass 8 km NW Sapa, 22°21'13"N, 103°46'01"E, 2030 m, 10.viii.2013, forested margin, V. Assing (MFNB, 1); Hoang Lien N.P., 22.35, 103.77, 1500–2000 m, 19.v.2019, VN168, S.M. Smith, A.I. Cognato, ex 10 cm branch (MSUC, 7). Ninh Binh, Cuc Phuong N.P., Mac Lake, 20°15'29.0"N, 105°42'27.5"E, 155 m, 4–7.v.2009, J.B. Heppner, ex blacklight trap (FSCA, 1). Thua Thien-Hue, Bach Ma N.P., 16.18902, 107.8498, 1193 m, 15.ii.2017, VN54, A.I. Cognato, T.A. Hoang, ex 1–4 cm diameter branch (MSUC, 1).

Diagnosis

3.4–4.8 mm long (mean = 4.26 mm; n = 5); 1.7–2.53× as long as wide. This species is distinguished by each declivital interstriae variously tuberculate, never granulate; and red-brown color.

This species strongly resembles A. minor from which it can usually be distinguished by the larger size and the tubercles on declivital interstriae 2 distinctly larger than those of other interstriae.

Similar species

Ambrosiodmus minor.

Distribution

China (Guangdong, Guizhou, Guangxi, Hainan, Hong Kong*, Sichuan, Xizang, Yunnan), India (Arunachal Pradesh*, Assam, Tamil Nadu, West Bengal), Indonesia (Java, Kalimantan, Sumatra), Japan, East & West Malaysia, Myanmar, Philippines, South Korea, Sri Lanka, Taiwan, Thailand, Vietnam. Established in USA (Hoebeke 1991; Gomez et al. 2018a).

Host plants

The species is polyphagous but may show some preference for Dipterocarpaceae in the southern part of its range, and for Fagaceae in the northern part (Browne 1961b).

Ambrosiodmus minor (Stebbing, 1907)

Fig. 12A, B, E

Phloeosinus minor Stebbing, 1907: 37.

Dryocoetes minor (Stebbing): Stebbing 1914: 549.

Xyleborus minor (Stebbing): Beeson 1930: 70.

Ambrosiodmus minor (Stebbing): Wood and Bright 1992: 676.

Xyleborus crassus Hagedorn, 1910a: 8. Synonymy: Schedl 1962a: 208.

Type material

Holotype Phloeosinus minor (FRI).

New records

China: Chongqing, NanShan, 15.viii.2015, J-G Wang, Lv-Jia, Tian-Shang (RABC, 3). Jiangsu, Nanjing, Zijinshan, 10.viii.2017, Y. Li, ex unknown log (MSUC, 1). Jiangxi, Jinggang Shan Mts, Jingzhushan Zhufeng, forested slopes of river valley, 26°32.0'N, 114°08.6'E, 805 m, 29.iv.2011, M. Ficáček, J. Hájek (MNHP, 1). Zhejiang, Tianmu Shan, pass 25 km NW Linan, 620–820 m, 30°25'40"N, 119°35'30"E, creek valley with bamboo and mixed forest, litter, sifted, 16.vi.2007, M. Schülke (MFNB, 1). Laos: Louangnantha, Nantha to Muang Sing, 21°09'N, 101°19'E, 900–1200 m, 5–31.v.1997, V. Kubáň (NHMB, 3). NE, Hua Phan, Ban Saluei, Phou Pan (Mt.), 20°12'N, 104°01'E, 1300–1900 m, 7.iv–25.v.2010, C. Holzschuh (NHMUK, 2); NW, 5 km SW Muang Sing, Chiang Tung (Stupa) GH, 750 m, 26.iii–5.iv.2010, S. Murzin (IRSNB, 4); N, 10 km N Luang Prabang, Mekon [sic] riv., 240 km N. Vientiane, hill county [sic], sparse, settled primary vegetation, ix.1992, I. Somay (RABC, 1). Taiwan: [Pingtung Co.], Henchun, Kuraru [Kenting Forestry Park], 250 m, 3.iv.1965, C.M. Yoshimoto (BPBM, 1). Vietnam: Hoa Binh, 1929, A. DeCooman (MNHN, 1); as previous except: 1934 (MNHN, 1). Lao Cai, Nam Tha, 22.01218, 104.37685, 9.v.2015, Pham Thu, ex funnel trap (RJRC, 1).

Diagnosis

3.5–4.0 mm long (mean = 3.74 mm; n = 5); 2.19–2.53× as long as wide. This species is distinguished by each declivital interstriae variously tuberculate, never granulate; and red-brown color.

This species strongly resembles A. lewisi from which it can usually be distinguished by the smaller size and tubercles on interstriae 2 not distinctly larger than those of other interstriae.

Similar species

Ambrosiodmus lewisi.

Distribution

Bangladesh, Bhutan, China (Chongqing, Guangxi, Jiangsu, Jiangxi*, Sichuan, Yunnan, Zhejiang), India (Assam, Meghalaya, Madhya Pradesh, Maharashtra, Uttarakhand, Uttar Pradesh, West Bengal), Laos*, East Malaysia, Myanmar, Nepal, Taiwan, Thailand, Vietnam. Established in the USA (Rabaglia and Okins 2011; Gomez et al. 2018a).

Host plants

Polyphagous (Beeson 1930, 1961; Ohno 1990; Maiti and Saha 2004; Lin et al. 2019).

Remarks

Wood and Bright (1992) considered the species as being described by Stebbing in 1909 (Stebbing 1909: 20) rather than in 1907 despite listing this publication under the taxonomy section for the species. This error undoubtedly occurred because Stebbing classified Phloeosinus minor as a new species in both publications. However, in 1909 he states “this amplifies the description given of this insect in [1907]” at the end of the species description. This error has been unknowingly perpetuated throughout the literature published since 1992.

Figure 12.

Dorsal, lateral and declivital view of Ambrosiodmus minor, 3.5–4.0 mm (A, B, E), and A. rubricollis, 2.5–2.8 mm (C, D, F).

Ambrosiodmus rubricollis (Eichhoff, 1876)

Fig. 12C, D, F

Xyleborus rubricollis Eichhoff, 1876a: 202.

Ambrosiodmus rubricollis (Eichhoff): Wood 1989: 170.

Xyleborus taboensis Schedl, 1952b: 65. Synonymy: Wood 1989: 170.

Xyleborus strohmeyeri Schedl, 1975b: 457. Synonymy: Wood 1989: 170.

Type material

Holotype (IRSNB) . Not examined.

New records

China: Chongqing, Simian mtn, 7.v.2015, Tian-Shang, Lv-Jia (RABC, 1); as previous except: Jinfo mtn, 10.v.2015 (RABC, 2). Guangdong, Lantau Is., Shi Bi pool, hardwood plantation, 4.vi.2004, Li, Z-R. (RABC, 1). Guangxi, Jiangidi, 25°55.6'N, 110°14.8'E, 365 m, terraced fields surrounded with shrubs and bamboo forest, 12.iv.2013, M. Ficáček, J. Hájek, J. Růžička (MNHP, 1). Hong Kong Is., Shek O, secondary broadleaf trees & bamboo forest, 24.viii.2004 (RABC, 1); as previous except: Tai Po Kau, vi.2017, J. Skelton (MSUC, 1). Jiangxi, Gan Zhou, 7.vii.2016, Lv-Jia & Lai-S-C, ex Senna surattensis (RABC, 1); as previous except: Jinggang Shan, Jingzhushan Zhufeng, 26°31.0'N, 114°05.9'E, 640 m, stream valley, 25.iv.2011, M. Ficáček, J. Hájek (MNHP, 1); as previous except: 26°32.0'N, 114°08.6'E, 805 m, forested slopes of river valley, 29.iv.2011 (RABC, 1). Laos: Vientiane, Ban Van Eue, 15.xii.1965, native collector (BPBM, 1). Vietnam: Cao Bang, 22°33.118'N, 105°52.537'E, 1048 m, 12–17.vi.2014, VN9, Cognato, Smith, Pham, FIT (MSUC, 5). Thua Thien-Hue, Bach Ma N.P., 16.18902, 107.8498, 1193 m, 15.ii.2017, VN54, A.I. Cognato, T.A. Hoang, ex 1–4 cm diameter branch (MSUC, 1). Tuyen Quang, Doi Can Tuyen Quang, 21.72740, 105.22742, 15.iv.2015, R.J. Rabaglia, funnel trap (RJRC, 1). Yen Bai, Tan Huong, 21.82410, 104.89651, 15.iv.2015, funnel trap (RJRC, 1).

Diagnosis

Host plants

Polyphagous (Faccoli 2008; Beaver and Liu 2010).

Remarks

Kasson et al. (2016) have shown that the symbiotic association of the species with the fungus, Flavodon ambrosius, has allowed niche expansion with large, long-lived, interconnecting colonies, overlapping generations, and pre-dispersal oviposition by young females.

Figure 13.

Dorsal, lateral and declivital view of Ambrosiophilus atratus, 3.3–3.5 mm (A, B, I), A. caliginestris holotype, 2.9 mm (C, D, J), A. consimilis holotype, 2.6–3.5 mm (E, F, K), and A. cristatulus, 2.1–2.3 mm (G, H, L).

Ambrosiophilus caliginestris sp. nov.

http://zoobank.org/03B24F5A-E7B4-4AE8-AA98-FC688F1B0828

Fig. 13C, D, J

Type material

Holotype , female, Vietnam: Cao Bang, 22°36.454'N, 105°52.083'E, 1661 m, 15.iv.2014, VN35, Cognato, Smith, Pham, ex phloem (MSUC).

Diagnosis

2.9–3.1 mm long (mean = 3.0 mm; n = 2); 2.42–2.58× as long as wide. This species is distinguished by all declivital interstriae granulate along the entire length; pronotum from lateral view tall (type 2); declivital striae 1 weakly impressed; declivital interstriae moderately and uniformly granulate, granules spaced by a distance of four diameters of a granule; interstrial setae long, hair-like; and moderate size.

Similar species

Ambrosiophilus atratus, A. satoi, A. sulcatus.

Description

(female). 2.9 mm long (n = 1); 2.42× as long as wide. Body ferruginous. Legs and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; surface shagreened, punctate; punctures dense, becoming shallower and sparser on reticulate upper part of frons. Eyes feebly emarginate just above antennal insertion, upper part smaller than lower part. Submentum narrow, triangular, slightly impressed. Antennal scape regularly thick, longer than club. Pedicel as wide as scape, much shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club approximately circular and flat, type 3; segment 1 corneous, transverse on anterior face, occupying approximately basal 1/2; segment 2 narrow, corneous; segments 1 and 2 present on posterior face. Pronotum: 0.9× as long as wide. In dorsal view basic and parallel-sided, type 2, sides parallel in basal 1/2, rounded anteriorly; anterior margin without serrations. In lateral view tall, type 2, disc flat, summit pronounced. Anterior slope with densely spaced small asperities, becoming lower and more strongly transverse towards summit. Disc shagreened with sparse, fine punctures bearing long, fine, erect hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Elytra: 1.54× as long as wide, 1.6× as long as pronotum. Scutellum moderately sized, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then broadly rounded to apex. Disc shiny, striae not impressed, punctures moderately coarse, shallow, separated by less than one diameter of a puncture, glabrous; interstriae flat, finely punctate, punctures more widely separated than those of striae, with long, fine, erect hair-like setae. Declivity steep, strongly convex, shiny; strial punctures larger than on disc, striae 1 weakly impressed; interstriae moderately and uniformly granulate, granules spaced by a distance of four diameters of a granule, each granule with a moderately long, erect hair-like seta. Posterolateral margin carinate, granulate to interstriae 7. Legs: procoxae contiguous; prosternal coxal piece tall, pointed. Protibiae slender, obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with eight large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with eight large socketed denticles.

Etymology

L. caligo = fog; -estris = belonging to. In reference to the climate of the type localities. An adjective.

Distribution

Vietnam.

Host plants

Unknown.

Ambrosiophilus consimilis (Eggers, 1923), comb. nov.

Fig. 13E, F, K

Xyleborus consimilis Eggers, 1923: 180.

Ambrosiodmus consimilis (Eggers): Wood and Bright 1992: 672.

Type material

Holotype (MCG).

New records

India: Bengal [West Bengal], Samsing, xi.1933, B. Singh, ex Litsea sp. (NMNH, 3).

Diagnosis

Host plants

Unknown.

Figure 14.

Dorsal, lateral and declivital view of Ambrosiophilus indicus holotype, 2.4 mm (A, B, I), A. lannaensis holotype holotype, 1.95–2.05 mm (C, D, J), A. latisulcatus, 3.9–4.2 mm (E, F, K), and A. osumiensis, 2.3–3.2 mm (G, H, L).

Ambrosiophilus lannaensis sp. nov.

http://zoobank.org/A9A545CB-1BEC-4B49-93AA-9A7208B0A24A

Fig. 14C, D, J

Type material

Holotype , female, Thailand: Chiang Mai, Doi Pui, 18°50'23"N, 98°53'53"E, 1200–1300 m, 2-BM-Jun-B-23 (2016), [vi.2016], S. Sanguansub et al., ex Butea monosperma (MSUC). Paratypes, female, as holotype except: 3-BM-Jun-B-26 (NHMUK, 1); as previous except: 2-CAJun-B-14, ex Castanopsis armata (QSBG, 1); as previous except: 4-CA-Jun-B-114 (RABC, 1); as previous except: 4-CA-Jun-B-8 (SSC, 1); as previous except: 4-CA-Jun-B-87 (SSC, 1); as previous except: 2-LT-Jun-B-91, ex Lithocarpus tenuinervis (SSC, 1); as previous except: Doi Pui, Chiang Khian Highl. Res. Stn, 27.vii.2013, S. Buranapanichpan, ex persimmon, Diospyros kaki (RABC, 1); as previous except: ex Mangifera indica (MSUC, 1); as previous except: Doi Pui, 1400 m, 25.v–2.vi.2006, W. Puranasakul, ex EtOH trap (RABC, 1).

Diagnosis

1.95–2.05 mm long (mean = 2.01 mm; n = 5); 2.67–2.73× as long as wide. This species is distinguished by all declivital interstriae granulate on upper 1/2 of declivity; pronotum from dorsal view conical and elongate (type 5), from lateral view type 7; pronotal disc shiny, punctures moderately fine and separated by several times their diameter; posterolateral margins of elytra rounded; lower part of declivity flattened; and declivital striae not impressed, interstriae finely, sparsely granulate on upper part of declivity only.

Similar species

Ambrosiophilus atratus, A. caliginestris, A. satoi, A. wantaneeae.

Description

(female). 1.95–2.05 mm long (mean = 2.01 mm; n = 5); 2.67–2.73 × as long as wide. Body dark brown, antennae and legs light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; surface subshiny, punctate; punctures moderately dense, becoming shallower and sparser on reticulate upper part. Eyes deeply emarginate just above antennal insertion, upper part smaller than lower part. Submentum distinctly triangular, slightly impressed. Antennal scape regularly thick, approximately as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club approximately circular and flat, type 3; segment 1 corneous, transverse on anterior face, occupying approximately basal 1/3; segment 2 narrow, corneous; segments 1 and 2 present on posterior face. Pronotum: 1.13× as long as wide. In dorsal view conical and elongate, type 5, sides almost parallel in basal 1/2, conical anteriorly; anterior margin without serrations. In lateral view elongate, disc longer than anterior slope, type 7, summit not pronounced, on anterior 1/3. Anterior slope with widely spaced, small coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny with moderately dense small, deep punctures bearing short, fine, erect hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Elytra: 1.63× as long as wide, 1.67× as long as pronotum. Scutellum small, triangular, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then broadly rounded to apex. Disc shiny, striae not impressed, with moderately coarse, shallow punctures separated by one width of their diameter, each bearing a short, semi-erect hair-like seta; interstriae flat, finely punctate, punctures more widely separated than those of striae, with fine, semi-erect setae. Declivity steep, strongly convex, shagreened; strial punctures larger than on disc, striae 1 and 2 very weakly impressed; interstriae unarmed by granules, each puncture bearing a moderately long, erect hair-like seta. Posterolateral margin rounded, unarmed. Legs: procoxae contiguous; prosternal coxal piece short, pointed. Protibiae slender, obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with five large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with seven large socketed denticles.

Etymology

The specific name refers to the old Northern Thai kingdom ‘Lan Na’. Latinized adjective.

Distribution

Thailand.

Host plants

This species is evidently polyphagous and is here reported from Mangifera indica (Anacardiaceae), Diospyros kaki (Ebenaceae), Butea monosperma (Fabaceae), Castanopsis armata, and Lithocarpus tenuinervis (Fagaceae).

Ambrosiophilus latisulcatus (Eggers, 1940)

Fig. 14E, F, K

Xyleborus latisulcatus Eggers, 1940: 142.

Ambrosiodmus latisulcatus (Eggers): Wood and Bright 1992: 675.

Ambrosiophilus latisulcatus (Eggers): Beaver et al. 2014: 25.

Type material

Holotype Xyleborus latisulcatus (NMNH).

Diagnosis

3.9–4.2 mm long (mean = 4.05 mm; n = 2); 2.52–2.8× as long as wide. This species is distinguished by declivital interstriae 1–3 each armed by one major tubercle surrounding declivital sulcus; pronotum from dorsal view conical frontally (type 6); pronotal anterior slope steep, flat; anterior margin with a row of six serrations; pronotum from lateral view tall (type 2); pronotal surface reticulate, discal punctures coarse, dense, spaced less than the diameter of a puncture; declivity moderately sulcate to interstriae 3, margins of sulcus armed with three equally sized tubercles: one at the base of interstriae 1, one on interstriae 2 just ventrad to the first, and one at the midpoint of interstriae 3.

Similar species

Ambrosiophilus sexdentatus, A. sulcatus.

Distribution

Indonesia (Java), Thailand.

Host plants

Unknown.

Ambrosiophilus osumiensis (Murayama, 1934)

Fig. 14G, H, L

Xyleborus osumiensis Murayama, 1934: 292.

Ambrosiophilus osumiensis (Murayama): Smith et al. 2018b: 393.

Xyleborus metanepotulus Eggers, 1939b: 119. Synonymy: Smith et al. 2018b: 393.

Xyleborus nodulosus Eggers, 1941b: 233. syn. nov.

Xyleborus pernodulus Schedl, 1957: 85. Unnecessary replacement name. Synonymy: Browne 1961c: 50.

Xyleborus hunanensis Browne, 1983b: 33. Synonymy: Beaver 2011: 283.

Ambrosiophilus peregrinus Smith & Cognato, 2015: 216. Synonymy: Smith et al. 2017: 552.

Type material

Holotype Xyleborus hunanensis (IZAS). Holotype Xyleborus metanepotulus (TARI). Holotype Xyleborus nodulosus (ZMFK). Holotype Xyleborus osumiensis (NMNH). Holotype Ambrosiophilus peregrinus (NMNH), paratypes (MSUC, 5).

New records

China: Anhui, Chuxian, 32.25N, 118.28E, 1.v.1965, Pistacia chinensis (NMNH, 4). Chongqing, Nan Shan, 20.viii.2015, Wang, J-G., Lv-Jia, Tian-Shang (RABC, 4); as previous except: Simian mtn, 7.v.2016, Tian-Shang, Lv-Jia (RABC, 2); as previous except: Youyang, 5.vi.2016, Tian-Shang (RABC, 1). Guangxi, Malu, 27.iii.2018, Y. Li, ex Cinnamomum cassia (UFFE, 1); as previous except: Shangsi, 25.iii.2018, ex Broussonetia papyrifera (UFFE, 1); as previous except: unknown host (UFFE, 1); as previous except: Shiwandashan, 25.iii.2018, Y. Li, ex Quercus griffithii (UFFE, 1); as previous except: Shangsi, 26.iii.2018, Y. Li, ex Quercus griffithii (UFFE, 1). Jiangxi, Ganzhou, Lv-Jia, ex Ligustrum lucidum (RABC, 1). Sichuan, Emei mtn, 18.viii.2016, Tian-Shang (RABC, 1). Yunnan, Xishuangbanna, Sanchahe Nat. Res., 22°09.784'N, 100°52.256'E, 2186 m, 29–30.v.2008, A.I. Cognato (MSUC, 1). Vietnam: Cao Bang, 22°34.118'N, 105°52.537'E, 1048 m, 12–17.iv.2014, VN9, Cognato, Smith, Pham, FIT (MSUC, 1). Ninh Binh, Cuc Phuong N.P., 7.iii.2018, 20.34932, 105.59669, 431 m, A.I. Cognato, S.M. Smith, VN 130, ex standing dead laurel (MSUC, 2). Thua Thien-Hue, Bach Ma N.P., 16.18902, 107.8498, 1193 m, 15.ii.2017, VN54, A.I. Cognato, T.A. Hoang, ex 1–4 cm diameter branch (MSUC, 1).

Diagnosis

2.3–3.2 mm long (mean = 2.6 mm; n = 7); 2.3–2.67× as long as wide. This species is distinguished by declivital interstriae 1 unarmed, 2 armed by 3–5 pointed tubercles along its length, major declivital tubercles on interstriae 2; weakly to moderately sulcate to striae 1, interstriae 2 convex, bearing 3–5 pointed tubercles and several small granules (near apical and basal margins) along its length; pronotum from dorsal view basic or subquadrate (type 2 or 3); and pronotum from lateral view basic (type 0).

Similar species

Ambrosiophilus papilliferus, A. subnepotulus, A. wantaneeae.

Distribution

China (Anhui, Chongqing*, Fujian, Guangxi*, Guizhou, Hunan, Jiangxi*, Sichuan*, Yunnan), Japan, Taiwan, Vietnam. Imported and established in USA (Smith and Cognato 2015; Schiefer 2018).

Host plants

This species is likely polyphagous and has been recorded from numerous host families including Pistacia (Anacardiaceae), Ilex (Aquifoliaceae), Quercus (Fagaceae), Cinnamomum (Lauraceae), Broussonetia (Moraceace), and Ligustrum (Oleaceae).

Remarks

The morphology of A. osumiensis is highly variable in regard to numerous characteristics that are routinely used to diagnose other xyleborine species. Such variation includes: the antennal club type either 3 or 4; pronotum basic (type 2) or subquadrate (type 3) from dorsal view; declivity shiny or shagreened; pronotal disc shiny or shagreened; number and size of tubercles on declivital interstriae 2; and a large size range with individuals differing by up to 0.9 mm in length. This variation led to A. osumiensis being described several times. Types of each species are distinct and diagnosable. Examination of the specimens listed above in ‘new records’ as well as the holotypes showed that these species formed a continuous spectrum of variation. During our fieldwork we were able to collect and sequence specimens that fell within the concept of X. metanepotulus (Vietnam), X. hunanensis (China), X. nodulosus (China) and A. peregrinus (Georgia, USA) and an additional larger morphospecies from multiple localities in Vietnam. COI sequences showed that all populations differed by no more than 7.4% supporting the hypothesis of one morphologically variable species. Typical intraspecific variation in xyleborines is below 10% (Cognato et al. 2020b). Xyleborus hunanensis, X. metanepotulus, X. nodulosus and A. peregrinus are thus all conspecific and considered synonyms of the oldest name, A. osumiensis.

The identification of A. nodulosus from East Malaysia by Browne (1980b) and Ohno (1990) appears to be incorrect. We have therefore omitted East Malaysia from the distribution, and also omitted the associated host records. The host records reported in Smith et al. (2017) are therefore incorrect.

Ambrosiophilus papilliferus sp. nov.

http://zoobank.org/B552010D-0595-470E-8A24-01C0C81F7964

Fig. 15A, B, I

Type material

Holotype , female, 贵州平塘核桃 1981.VI.6 采集者：罗禄怡 [China: Guizhou, Pingtang, 6.vi.1981, Luyi Luo, ex Carya sp.] (NMNH). Paratype, female, Vietnam: Thua Thien-Hue, Bach Ma N.P., 16.18902, 107.8498, 1193 m, 15.ii.2017, VN54, A.I. Cognato, T.A. Hoang, ex 1–4 cm diameter branch (MSUC).

Diagnosis

2.5 mm long (n = 1); 2.5× as long as wide. This species is distinguished by declivital interstriae 1 unarmed, interstriae 2 armed by four or five moderately sized and variably spaced denticles along its length, interstriae 3 armed by five larger denticles; declivital striae 1 and 2 moderately impressed; and pronotum from dorsal view basic (type 2), lateral view basic (type 0).

Similar species

Ambrosiophilus osumiensis, A. wantaneeae.

Description

(female). 2.5 mm long (n = 1); 2.5× as long as wide. Body color red-brown, antennae and legs light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; surface subshiny, punctate; punctures moderately dense, becoming shallower and sparser on reticulate upper part. Eyes deeply emarginate just above antennal insertion, upper part smaller than lower part. Submentum narrow, triangular, slightly impressed. Antennal scape regularly thick, shorter than club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club tall and oval, flat, type 3; segment 1 convex and small on anterior face, occupying approximately basal 1/6; segment 2 corneous, narrow; segments 1–3 present on posterior face. Pronotum: 0.79× as long as wide. In dorsal view basic, type 2, sides parallel in basal 1/2, rounded anteriorly; anterior margin without serrations. In lateral view basic, type 0, disc flat, summit pronounced. Anterior slope with widely spaced, small coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny with moderately dense small, shallow punctures bearing short, fine, erect hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Elytra: 1.64× as long as wide, 2.1× as long as pronotum. Scutellum moderately sized, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then broadly rounded to apex. parallel-sided in basal 3/4, then broadly rounded to apex. Disc opalescent, striae weakly impressed, with moderately coarse, shallow, and irregular punctures separated by 0.5–1 diameter of a puncture, glabrous; interstriae flat, finely punctate, punctures more widely separated than those of striae, with fine, erect hair-like setae. Declivity steep, strongly convex, shagreened; strial punctures larger than on disc, striae 1 and 2 moderately impressed, strial punctures bearing short, recumbent setae 1× width of a puncture; interstriae 1 unarmed by granules, interstriae 2 with four or five coarse granules, interstriae 3 and 4 with four or five slightly smaller granules, each granule with a moderately long, erect hair. Posterolateral margin carinate to interstriae 7. Legs: procoxae contiguous, prosternal coxal piece short, pointed. Protibiae slender, obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with six large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened, outer margins evenly rounded with eight large socketed denticles.

Etymology

L. papilla = nipple; adjectival suffix ferus = bearer. In reference to the denticles on the declivity. An adjective.

Distribution

China (Guizhou), Vietnam.

Host plants

Recorded only from Carya (Juglandaceae).

Remarks

Locality labels on the holotype are in Chinese and were translated by You Li. An English locality label has been placed on the specimen below the original locality labels.

Figure 15.

Dorsal, lateral and declivital view of Ambrosiophilus papilliferus holotype, 2.5 mm (A, B, I), A. satoi paratype, 2.5–2.75 mm (C, D, J), A. sexdentatus, 2.7–3.0 mm (E, F, K), and A. subnepotulus, 2.5–2.8 mm (G, H, L).

Ambrosiophilus satoi (Schedl, 1966)

Fig. 15C, D, J

Xyleborus satoi Schedl, 1966b: 39.

Ambrosiophilus satoi (Schedl): Beaver and Liu 2010: 22.

Type material

Paratype (NHMW).

New records

Thailand: Chiang Mai, Doi Suthep, 1400 m, EtOH trap, 16–20.v.2005, W. Puranasakul (RABC, 1); as previous except: 20.vii.2016, S. Sanguansub et al. (RABC, 1); as previous except: 18°50'23"N, 98°53'53"E, 1200–1300 m, vi.2016, S. Sanguansub et al., ex Castanopsis armata (RABC, 1); as previous except: ex Lithocarpus tenuinervis (RABC, 1).

Diagnosis

2.5–2.75 mm long (mean = 2.67 mm; n = 5); 2.5–2.57× as long as wide. This species is distinguished by all declivital interstriae granulate on upper 1/2 of declivity; pronotum from lateral view basic (type 0); declivity rounded, face flattened; declivital interstriae sparsely and uniformly granulate, granules spaced by a distance of at least four diameters of a granule; interstrial setae short, bristle-like; and small size.

Similar species

Ambrosiophilus atratus, A. caliginestris, A. latisulcatus, A. sulcatus.

Distribution

Bhutan, Taiwan, Thailand*.

Host plants

Recorded from a ‘camphor log’ (probably Cinnamomum camphora (Lauraceae)) (Schedl 1966b), and from Castanopsis armata and Lithocarpus tenuinervis (Fagaceae).

Ambrosiophilus sexdentatus (Eggers, 1940)

Fig. 15E, F, K

Xyleborus sexdentatus Eggers, 1940: 148.

Ambrosiodmus sexdentatus (Eggers): Wood and Bright 1992: 680.

Ambrosiophilus sexdentatus (Eggers): Hulcr and Cognato 2009: 24.

Type material

Holotype (NMNH).

Diagnosis

2.7–3.0 mm long (mean = 2.84 mm; n = 5); 2.7–2.9× as long as wide. This species is distinguished by declivital interstriae 2 (1 spine), interstriae 3 (2 spines) surrounding declivital sulcus; pronotum from dorsal view basic (type 2); pronotal anterior slope rounded, convex; pronotum anterior margin lacking serrations; pronotum from lateral view tall (type 2); pronotal discal punctures small, fine spaced by at least two diameters of a puncture, surface shiny; declivity strongly sulcate to interstriae 3, lateral margins of sulcus rounded, margin armed with three large spines, one at the base of interstriae 2, one at the declivital midpoint of interstriae 3 and one on the apical 1/3 of interstriae 3.

Similar species

Ambrosiophilus latisulcatus, A. sulcatus.

Distribution

Indonesia (Java), New Guinea, Thailand.

Host plants

Recorded from Quercus (Fagaceae) and Tectona (Lamiaceae) in Java (Kalshoven 1959b).

Remarks

A mycocleptic associate of Beaverium species (Hulcr and Cognato 2010b, 2013).

Ambrosiophilus subnepotulus (Eggers, 1930)

Fig. 15G, H, L

Xyleborus subnepotulus Eggers, 1930: 178.

Ambrosiodmus subnepotulus (Eggers): Wood and Bright 1992: 680.

Ambrosiophilus subnepotulus (Eggers): Beaver and Liu 2010: 22.

Xyleborus cristatuloides Schedl, 1971a: 284. syn. nov.

Type material

Holotype Xyleborus subnepotulus (FRI). Lectotype Xyleborus cristatuloides (NHMW).

New records

China: Guizhou, Guiyang, Huaxi, 31.iv.2015, Y. Li, ex in flight (UFFE, 9). Hong Kong, vi.2017, J. Skelton (MSUC, 15). Laos: Vientiane, Ban Van Eue, 31.xii.1965, native collector (BPBM, 2).

Diagnosis

2.5–2.8 mm long (mean = 2.64 mm; n = 7); 2.27–2.6× as long as wide. This species is distinguished by declivital interstriae 1 unarmed, interstriae 2 armed by one tubercle at declivital summit, remainder of interstriae 2 unarmed, interstriae 3 with two large tubercles; declivity weakly bisulcate from sutural margin to striae 2; interstriae 3 weakly convex; pronotal surface shiny, discal punctures minute, very fine, widely spaced by 2–6 diameters of a puncture; and declivital surface smooth, shiny; and moderate size.

Similar species

Ambrosiophilus consimilis, A. cristatulus, A. indicus, A. osumiensis.

Distribution

China* (Guizhou, Hong Kong*), Indonesia (Java), Laos*, Myanmar, Sri Lanka, Taiwan.

Host plants

The only recorded host is Albizia lebbeck (Fabaceae) (Beeson 1930).

Remarks

Wood (1989) considered X. cristatuloides Schedl as a synonym of Ambrosiodmus asperatus. However, the lectotype has a punctate pronotal disc and declivital sculpturing that is almost identical with Ambrosiophilus subnepotulus and it is here placed in synonymy.

Ambrosiophilus sulcatus (Eggers, 1930)

Fig. 16A, B, E

Xyleborus sulcatus Eggers, 1930: 180.

Ambrosiodmus sulcatus (Eggers): Wood and Bright 1992: 680.

Cyclorhipidion sulcatum (Eggers): Maiti and Saha 2004: 118.

Ambrosiophilus sulcatus (Eggers): Beaver and Liu 2018: 537.

Xyleborus sulcatulus Eggers, 1939a: 13. syn. nov.

Xyleborus sinensis Eggers, 1941b: 224. syn. nov.

Type material

Holotype Xyleborus sinensis (ZMFK). Holotype Xyleborus sulcatus (FRI). Holotype Xyleborus sulcatulus (NHRS).

New records

China: Jiangxi, Wu-Yi Mt., 19.vii.2017, Lai, S-C, Tian Shang et al. (RABC, 1). India: Bengal [West Bengal], Darjeeling, Debrepani, 6000 ft, 15.ix.1929, J.C.M. Gardner, unknown wood (NMNH, 1). Taiwan: [Formosa], Taiheizan, 9.v.[19]32, L. Gressitt (NMNH, 1). Chiayi Co., Fenkihu, 1370 m, 10–12.iv.1965, C.M. Yoshimoto, B.D. Perkins (BPBM, 1). Vietnam: Hoa Binh, 1940, A. DeCooman (MNHN, 1), Lao Cai, 16 km W of Sa Pa, 1800 m, at light, 17.iii.1998, L. Peregovits, T. Vásárhelyi (RABC, 1).

Diagnosis

3.4–4.5 mm long (mean = 3.94 mm; n = 5); 2.5–2.87× as long as wide. This species is distinguished by all declivital interstriae granulate along the entire length; pronotum from dorsal view basic (type 2); pronotal anterior slope rounded; pronotal anterior margin without a row of serrations; pronotum from lateral view tall (type 2); declivity weakly to strongly bisulcate between striae 1 and interstriae 3; interstriae densely and uniformly granulate, granules on interstriae 3 spaced by a distance of less than the diameter of a granule; interstrial setae long, hair-like, and of large size.

Ambrosiophilus sulcatus is variable in body length, the degree of bisulcation of the declivity and in the size of the declivital granules, but all specimens form a continuous spectrum of variation. Specimens from India and China (Fujian) are larger, more strongly bisulcate and have slightly larger granules than specimens occurring further south (Myanmar and Vietnam).

Similar species

Ambrosiophilus atratus, A. caliginestris, A. latisulcatus, A. satoi.

Distribution

China (Fujian, Jiangxi*), India (Assam, West Bengal*), Myanmar, Nepal, Taiwan*, Vietnam*.

Host plants

Recorded only from Artocarpus (Moraceae) (Beeson 1930).

Remarks

The type specimens of Xyleborus sinensis, X. sulcatulus and type images of X. sulcatus, were directly examined. The specimens differ in size (2.8 mm X. sulcatulus, 3.0 mm, X. sulcatus, 4.2 mm X. sinensis), the depth of the declivital sulci and in the degree development of interstrial granules. Additional non-type specimens were also examined. We found that size, depth of the declivital sulci and development of interstrial granules formed a continuum of variation and should be considered a single morphologically variable species.

Figure 16.

Dorsal, lateral and declivital view of Ambrosiophilus sulcatus holotype, 3.4–4.5 mm (A, B, E), and A. wantaneeae holotype, 2.0–2.1 mm (C, D, F).

Ambrosiophilus wantaneeae sp. nov.

http://zoobank.org/1BF7E0E1-F0EC-4121-8CA5-02D5122BD6C1

Fig. 16C, D, F

Type material

Holotype , female, Thailand: Chiang Mai, Doi Pui, 1400 m, 17.iv.–8.v.2006, W. Puranasakul, ex EtOH trap (NHMUK). Paratypes, female, as holotype except: 25.iv.–16.v.2005 (QSBG, 1; RABC, 1); as previous except: flight intercept trap (MSUC, 1); as previous except: 18°50'23"N, 98°53'53"E, 1200–1300 m, 30.iv.2014, S. Sanguansub et al., ex EtOH trap (RABC, 1).

Diagnosis

2.0–2.1 mm long (mean = 2.03 mm; n = 4); 2.63–2.77× as long as wide. This species is distinguished by declivital interstriae 1 unarmed, 2 armed by four or five coarse granules along its length, interstriae 3 with four or five slightly smaller granules; declivital striae 1 and 2 very weakly impressed; and pronotum from dorsal view conical (type 0) to subelongate (type 7), lateral view long (type 7).

Similar species

Ambrosiophilus osumiensis, A. papilliferus.

Description

(female). 2.0–2.1 mm (mean = 2.02 mm; n = 4); 2.63–2.67× as long as wide. Body dark brown to pitchy black, antennae and legs light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes; surface subshiny, punctate; punctures moderately dense, becoming shallower and sparser on reticulate upper part. Eyes deeply emarginate just above antennal insertion, upper part smaller than lower part. Submentum narrow, triangular, slightly impressed. Antennal scape regularly thick, approximately as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club approximately circular, type 3; segment 1 corneous, transverse on anterior face, occupying approximately basal 1/3; segment 2 narrow, corneous; segments 1 and 2 present on posterior face. Pronotum: 1.0–1.1× as long as wide. In dorsal view conical and elongate, type 5, sides almost parallel in basal 1/2, conical anteriorly; anterior margin without serrations. In lateral view elongate, disc as long as anterior slope, type 7, summit not pronounced, at midpoint. Anterior slope with widely spaced, small asperities, becoming lower and more strongly transverse towards summit. Disc strongly shiny with sparse, small, deep punctures bearing short, fine, erect hair-like setae. Some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Elytra: 1.6–1.7× as long as wide, 1.6–1.7× as long as pronotum. Scutellum small, triangular, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then broadly rounded to apex. Disc shiny, striae not impressed, parallel, with moderately coarse, shallow punctures separated by 1–2× their diameter, without hair-like setae; interstriae flat, finely punctate, punctures more widely separated than those of striae, with fine, erect hair-like setae. Declivity shiny, steep, strongly convex; strial punctures larger than on disc, striae 1 and 2 very weakly impressed; interstriae 1 without granules, interstriae 2 with four or five coarse granules, interstriae 3 and 4 with four or five slightly smaller granules, each granule with a moderately long, erect hair-like seta. Posterolateral margin rounded, unarmed. Legs: procoxae contiguous; prosternal coxal piece short, pointed. Protibiae slender, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with six moderate socketed denticles, their length slightly longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with eight small socketed denticles.

Etymology

The species is named for Ms. Wantanee Puranasakul (then at Chiang Mai University, Thailand) who collected several new species of Scolytinae during her MSc studies. Noun in genitive.

Distribution

Thailand.

Host plants

Unknown.

Ancipitis Hulcr & Cognato, 2013

Ancipitis Hulcr & Cognato, 2013: 41.

Type species

Xyleborus puer Eggers, 1923; original designation.

Diagnosis

1.9–5.4 mm long, 2.08–2.73× as long as wide. Ancipitis is distinguished by the flat submentum that is flush with genae and shaped as a distinct large triangle; elytra extremely long, flattened, very gradually descending, broadened laterally and elongated apically; declivital face appearing somewhat depressed below posterolateral costa and covered with hair-like setae; pronotum extended anteriad, appearing conical, type 0 in dorsal view, without serrations on anterior margin; antennal club flattened, type 3 with three sutures visible on the posterior face; scape long and slender; protibiae slender, all tibia bearing large denticles; procoxae appearing tall, longer than basal width; scutellum flat, flush with elytra; procoxae narrowly separated; mycangial tufts absent; elytra unarmed.

Similar genera

Diuncus, Leptoxyleborus.

Distribution

Distributed in temperate and tropical Asia and Melanesia.

Gallery system

This consists of branched tunnels without brood chambers (Browne 1961b). There may also be surface galleries between the bark and the sapwood (Kalshoven 1959b; Browne 1961b).

Key to Ancipitis species (females only)

1	Declivity weakly sulcate between interstriae 3 in middle of declivity; sutural interstriae weakly raised and striae 1 impressed in apical third; larger, 4.9–5.4 mm	*punctatissimus*
–	Declivity not sulcate between interstriae 3 in middle of declivity; sutural interstriae not raised and striae 1 not impressed in apical third; smaller, 3.0–3.6 mm	*puer*

Ancipitis puer (Eggers, 1923)

Fig. 17A, B, E

Xyleborus puer Eggers, 1923: 191.

Ancipitis puer (Eggers): Hulcr and Cognato 2013: 42.

Xyleborus ceramensis Schedl, 1937a: 549. Synonymy: Hulcr and Cognato 2013: 42.

Type material

Holotype (MCG), syntypes (MCG, 2; NMNH, 2).

Diagnosis

Moderately sized, 3.0–3.6 mm long (mean = 3.2 mm; n = 5); 2.14–2.73× as long as wide. This species is distinguished by its moderate size, declivity not sulcate between interstriae 3 in middle of declivity; sutural interstriae not raised and striae 1 not impressed in apical third; and declivital striae and interstriae both bearing long hair-like setae that are erect on the interstriae and semi-recumbent on the striae.

Similar species

Leptoxyleborus machili, L. sordicauda.

Distribution

Indonesia (Ceram, Sumatra), East & West Malaysia, New Guinea, Thailand.

Host plants

Recorded only from Shorea (Dipterocarpaceae) and Intsia (Fabaceae) (Browne 1961b), but probably polyphagous.

Figure 17.

Dorsal, lateral and declivital view of Ancipitis puer, 3.0–3.6 mm (A, B, E), and A. punctatissimus, 4.9–5.4 mm (C, D, F).

Ancipitis punctatissimus (Eichhoff, 1880)

Fig. 17C, D, F

Xyleborus punctatissimus Eichhoff, 1880: 189.

Leptoxyleborus punctatissimus (Eichhoff): Wood and Bright 1992: 660.

Ancipitis punctatissimus (Eichhoff): Beaver et al. 2014: 26.

Xyleborus spatulatus Blandford, 1896b: 218. Synonymy: Kalshoven 1959a: 95.

Type material

Syntypes Xyleborus spatulatus (NHMUK).

Diagnosis

The largest Ancipitis species, 4.9–5.4 mm long (mean = 5.16 mm; n = 5); 2.08–2.41× as long as wide. This species is distinguished by its large size; declivity weakly sulcate between interstriae 3 in middle of declivity; sutural interstriae weakly raised and striae 1 impressed in apical third; and declivital interstriae with three rows of mixed short erect and recumbent hair-like setae.

Similar species

Ancipitis puer, Leptoxyleborus sordicauda.

Distribution

Indonesia (Java, Sumatra), East & West Malaysia, Thailand.

Host plants

Recorded from four different families of angiosperm trees, and from Pinus merkusii (Pinaceae) (Browne 1961b; Ohno 1990; Schedl 1969a).

Anisandrus Ferrari, 1867

Anisandrus Ferrari, 1867: 24.

Type species

Apate dispar Fabricius, 1793; monotypy.

Diagnosis

2.1–5.9 mm, 1.88–2.78× as long as wide, body usually stout and dark. Anisandrus is distinguished most easily by the antennal club obliquely truncate type 1 (A. achaete type 2), club taller than wide (A. achaete wider than tall), procoxae narrowly separated, protibiae slender, obliquely or distinctly triangular, outer margin with 5–8 large socketed denticles on distal 1/2, posterior face unarmed, mesonotal mycangial tufts typically present along the pronotal base (missing in three species), either as a small tuft the length of the scutellum and directly opposite it or extending laterally from the scutellum to striae 3 and with elytral base broadly, shallowly emarginated from the scutellum to striae 3. Additional diagnostic characters include: pronotum from dorsal view typically types 0 and 1 (A. cryphaloides, type 6), pronotum from lateral view tall (type 3), or rounded and robust (type 5), pronotum anterior margin with a row of serrations, pronotum lateral margins obliquely costate, scutellum flat, flush with elytra, and the elytral disc either convex or variously transversely impressed with a saddle-like depression. Species range from nearly glabrous to densely setose and are typically black or dark brown.

Similar genera

Cnestus, Cyclorhipidion, Hadrodemius, Xylosandrus. Anisandrus is closely related to Cnestus, Hadrodemius and Xylosandrus, all of which possess a mesonotal mycangium and the associated dense tuft of hair-like setae at the scutellar area and pronotal base (Gohli et al. 2017; Johnson et al. 2018).

Distribution

Uncommon genus with species occurring in forests of the Holarctic and Paleotropical regions.

Gallery system

The species usually attack stems of small diameter, and the gallery system consists of a radial or circumferential gallery with several longitudinal branches without brood chambers. SMS collected several species (A. cristatus, A. lineatus, A. longidens) in northern Vietnam that had a preference for attacking small saplings just above the soil line.

Remarks

This genus is remarkably diverse in montane habitats across Asia but most species are poorly known. It is very likely that many additional new species await description.

Key to Anisandrus species (females only)

1	Pronotal mycangial tuft moderate to densely setose, very broad, extending laterally from the scutellum to striae 3 (Fig. 23E)	2
–	Pronotal mycangial tuft absent (Fig. 23C) or just anteriad and roughly equal in width to scutellum, lightly to moderately setose (Fig. 22C)	7
2	Posterolateral margin of elytra rounded (Fig. 18L); declivital face convex or flattened; smaller, 2.8–3.1 mm	3
–	Posterolateral margin of elytra costate or carinate to interstriae 5 (Fig. 23K); declivital face variably sulcate; larger, 3.9–5.6 mm	4
3	Elytral disc flat; declivital face moderately steep and convex; declivital summit with interstriae 1 unarmed, a small denticle on interstriae 2 and a minute denticle on interstriae 3; declivity shiny	auratipilus sp. nov.
–	Elytral disc with a broad and weak transverse saddle-like depression; declivital face steep, flattened; declivital summit with a minute denticle on interstriae 1, a small denticle on interstriae 2, and interstriae 3 unarmed; declivity opalescent	venustus sp. nov.
4	At least punctures of declivital striae 2 strongly confused, minute; pronotal asperities large, widely spaced; elytral disc with a profound transverse saddle-like depression; declivity broadly sulcate to interstriae 5	*percristatus*
–	Declivital strial punctures all uniseriate, large; pronotal asperities small, densely spaced; elytral disc with a weak to deep transverse saddle-like depression; declivity sulcate to interstriae 3	5
5	Elytral disc with a weak transverse saddle-like depression (Fig. 21B); declivital interstriae uniseriately punctate, and setose, setae erect, very long, very fine and hair-like	hera sp. nov.
–	Elytral disc with a deep transverse saddle-like depression (Fig. 21H); declivital interstriae impunctate or with biseriate punctures, and setae semi-erect, short, thick, or scale-like	6
6	Declivital interstriae impunctate, setose, setae semi-erect, short and thick; declivital summit with large incurved spine on interstriae 2; declivital interstriae 3 with six additional unequally sized incurved spines on basal 1/2 of declivity; larger, 5.4–5.6 mm	*klapperichi*
–	Declivital interstriae minutely biseriately punctate, setose, setae bristle-like, erect; declivital summit with a large incurved spine on interstriae 2, interstriae 3 unarmed; smaller, 4.0–4.15 mm	xuannu sp. nov.
7	Mesonotal mycangial tuft absent (Fig. 23C)	8
–	Mesonotal mycangial tuft just anteriad and roughly equal in width to scutellum, lightly to moderately setose (Fig. 22C)	10
8	Antennal club wider than longer, type 2, one suture visible on posterior face (Fig. 2); protibiae distinctly triangular; anterior margin of the pronotum without serrations	*achaete*
–	Antennal club longer than wide, type 1, no sutures visible on posterior face (Fig. 2); elytral disc convex; protibiae obliquely triangular; anterior margin of the pronotum with a row of serrations	9
9	Declivital interstriae 1 and 3 armed by 4–5 unequally sized tubercles; declivital striae strongly impressed; elytral disc with a weak transverse saddle-like depression; pronotal disc coarsely punctate; larger, 4.5 mm	*carinensis*
–	Declivital interstriae uniseriate granulate on basal 1/2, granules equally sized; striae clearly impressed; elytral disc convex; pronotal disc finely punctate; smaller, 2.8 mm	paragogus sp. nov.
10	Interstriae 2 and 3 of equal width at midpoint of declivity (Fig. 18K)	17
–	Interstriae 2 and 3 not equal in width at midpoint of declivity (Fig. 22J)	11
11	Interstriae 2 narrower than interstriae 3 at midpoint of declivity (Fig. 22J)	12
–	Interstriae 3 narrower than interstriae 2 at midpoint of declivity (Fig. 20J)	15
12	Declivity rounded, posterolateral margin rounded	13
–	Declivity obliquely truncate, posterolateral margin costate	14
13	Elytral disc with a weak transverse saddle-like depression; declivital interstriae 2 armed with a blunt tubercle at summit, interstriae 3 armed by one or two denticles near declivital summit ventrad to tubercle on interstriae 2	sinivali sp. nov.
–	Elytral disc convex; declivity unarmed	*hirtus*
14	Declivity weakly bisulcate, margins ornamented by large sharp spines on interstriae 2–7, spine on interstriae 3 the largest; declivital interstriae impunctate; posterolateral margin costate to interstriae 5.	*longidens*
–	Declivity steeply rounded and flat, declivital summit armed by a minute denticle on interstriae 2 and 3; granules present on basal 1/2 of interstriae 2–4; declivital interstriae clearly punctate, posterolateral margin costate to interstriae 7	*improbus*
15	Declivity steeply rounded and flat; elytral apex sharply angulate, nearly subquadrate; posterolateral margin costate to interstriae 5; pronotum rounded, type 1, in dorsal view	*eggersi*
–	Declivity gradual and convex, elytral apex broadly rounded; posterolateral margin rounded; pronotum conical, type 0, in dorsal view	16
16	Declivity strongly shagreened or opalescent; striae weakly impressed; smaller, 2.1–2.4 mm	cryphaloides sp. nov.
–	Declivity strongly shiny, striae deeply impressed; larger, 2.6–3.3 mm	*lineatus*
17	Declivital interstriae 2 punctate, punctures either uniseriate or confused	18
–	Declivital interstriae 2 impunctate, punctures may be replaced by granules	22
18	Declivital interstriae 2 punctures multiseriate and confused; body densely covered by erect dark brown pubescence	*ursulus*
–	Declivital interstriae 2 punctures uniseriate; body nearly glabrous or at most moderately setose	19
19	Declivity rounded and convex; posterolateral margin rounded	auco sp. nov.
–	Declivity steep and face variably impressed; posterolateral margin costate or carinate	20
20	Declivital summit unarmed; declivital face flat and weakly depressed below lateral margins	*mussooriensis*
–	Declivital summit ornamented by two small sharp incurved spines at the base of interstriae 2 and 3; declivital face flat and moderately bisulcate or concave	21
21	Declivity moderately bisulcate; declivital interstriae bearing erect fine hair-like setae	feronia sp. nov.
–	Declivital face concave; declivital interstriae bearing erect pointed bristle-like setae	*geminatus*
22	Posterolateral margins of elytra rounded; larger, 5.8–5.9 mm	*niger*
–	Posterolateral margins of elytra costate or carinate; smaller, 2.2–3.7 mm	23
23	Declivital summit without a sharp hooked spine on interstriae 2; declivital interstriae 2 face densely granulate or denticulate; elytral disc typically without a weak transverse saddle-like depression	24
–	Declivital summit with a sharp hooked spine on interstriae 2; declivital interstriae 2 face sparsely granulate; elytral disc flat, with a weak transverse saddle-like depression (rarely flat in some apicalis and cristatus)	25
24	Declivital interstriae denticulate; elytral discal interstriae punctures uniseriate; declivity appearing bisulcate with declivity impressed from striae 1 to interstriae 2, interstriae 3 distinctly raised; smaller, 2.2–2.5 mm	*maiche*
–	Declivital interstriae granulate; elytral discal interstriae with 2–3 confused rows of punctures; declivital interstriae 1 slightly raised, interstriae 2 and 3 flush; larger, 3.1–3.5 mm	*dispar*
25	Spine at declivital summit of interstriae 2 backwardly pointed; smaller, 2.6–2.8 mm	congruens sp. nov.
–	Spine at declivital summit of interstriae 2 incurved; larger, 3.05–3.7 mm	26
26	Spines interstriae 3 not backwardly hooked, much smaller than spine at the summit of interstriae 2; smaller, 3.05–3.4 mm; declivity weakly sulcate	*apicalis*
–	Spines interstriae 3 backwardly hooked, subequal to the spine at the summit of interstriae 2; larger, 3.35–3.7 mm; declivity moderately sulcate	*cristatus*

Anisandrus achaete sp. nov.

http://zoobank.org/53ED4F36-7BC7-4354-945D-2EE41160D8D9

Fig. 18A, B, I

Type material

Holotype , female, 云南勐养 700m 寄主:栎 1984.VII.19 [China: Yunnan, Mengyang, 700 m, 19.vii.1984, ex Fagaceae] (NMNH). Paratype, female, as holotype (IZAS).

Diagnosis

3.5 mm long (mean = 3.5 mm; n = 2); 2.33× as long as wide. This species is distinguished by the mesonotal mycangial tuft absent; antennal club type 2, one suture on posterior face; elytral disc with a weak transverse saddle-like depression near declivital summit; declivity unarmed by spines; declivital striae strongly impressed, interstriae granulate; and anterior margin of pronotum without serrations.

Similar species

Anisandrus apicalis.

Description

(female). 3.5 mm long (mean = 3.5 mm; n = 2); 2.33× as long as wide. Body bicolored with pronotal and elytral bases light brown, remainder of elytra red-brown. Head, legs, and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, strongly shiny, finely punctate; lateral areas weakly rugose, setose; each shallow ruga or puncture bearing a very long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club wider than long, obliquely truncate, type 2; segment 1 corneous, transverse on anterior face, occupying basal 2/5, nearly covering posterior face; segment 2 narrow, corneous; segment 1 present on posterior face. Pronotum: 0.89× as long as wide. In dorsal view basic, type 2, sides parallel in basal 1/2, rounded anteriorly; anterior margin without serrations. In lateral view basic, type 0, disc as long as anterior slope, summit at apical 2/5. Anterior slope with densely spaced, large fine asperities, becoming lower and more strongly transverse towards summit. Disc impressed behind summit, shiny, impunctate, glabrous, some long hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles acutely rounded. Mycangial tuft absent. Elytra: 1.55 × as long as wide, 1.75× as long as pronotum. Scutellum narrow, moderately sized, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 2/3, then broadly rounded to apex; surface shiny. Disc with a weak medial transverse saddle-like depression, striae 1–3 distinctly impressed, other striae not impressed, punctures small, deep, separated by 2–4 diameters of a puncture, glabrous; interstriae glabrous, unarmed, interstriae 1–4 feebly convex, punctate, punctures minute, confused. Declivity occupying approximately 1/3 of elytra, steeply rounded, declivital face flattened; striae deeply impressed, strial punctures much larger and deeper than those of disc; interstriae impunctate, uniseriate granulate, granules bearing setae 1.5× width of interstriae 2, erect, hair-like, interstriae 3 narrower than interstriae 2 at midpoint of declivity. Posterolateral margin rounded, unarmed by granules. Legs: procoxae contiguous, prosternal coxal piece tall and pointed. Protibiae distinctly triangular, broadest at apical 4/5, posterior face smooth; apical 1/2 of outer margin with eight moderate socketed denticles, their length slightly longer than basal width. Mesotibiae flattened, distinctly triangular, apical 1/2 with nine moderate socketed denticles on outer margin; metatibiae flattened, obliquely triangular, apical 1/2 with nine moderate socketed denticles on outer margin.

Etymology

G. a = without; chaite = long hair. In reference to the uncharacteristically reduced number of elytral setae. Noun in apposition.

Distribution

China (Yunnan).

Host plants

Recorded from Fagaceae.

Remarks

Locality labels on the holotype and paratype are in Chinese and were translated by You Li. An English locality label has been placed on each specimen below the original locality labels.

Anisandrus apicalis (Blandford, 1894)

Fig. 18C, D, J

Xyleborus apicalis Blandford, 1894b: 105.

Ambrosiodmus apicalis (Blandford): Wood 1989: 169.

Anisandrus apicalis (Blandford): Hulcr et al. 2007: 578.

Type material

Holotype (NHMUK).

New records

China: Jiangxi, Wu-Yi Mt., 17.vii.2017, Lai, S-C, Tian, S et al. (RABC, 1). Sichuan, Jiuzhago Nature Reserve, 33°08.865'N, 103°55.134'E, 2483 m, 5.vii.2005, A.I. Cognato, ex Pinus armandii (MSUC).

Diagnosis

3.05–3.4 mm long (mean = 3.17 mm; n = 5); 2.33–2.43× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc with or without a weak transverse saddle-like depression; declivital posterolateral margin costate to interstriae 5; declivity appearing bisulcate, weakly impressed from striae 1 and 2, interstriae 3 feebly inflated and tuberculate from base to apical 1/2 then becoming flattened and unarmed to apex; and moderately sized sharp incurved spine at base of declivity on interstriae 2.

This species strongly resembles A. cristatus and A. congruens and is most easily distinguished by the moderate size, the less strongly impressed declivital sulci and smaller spines on interstriae 3 that are not backwardly hooked and much smaller than the spine at the summit of interstriae 2.

Similar species

Anisandrus congruens, A. cristatus, A. geminatus, A. niger, A. sinivali, A. venustus.

Distribution

China (Anhui, Guangxi, Guizhou, Hainan, Jiangxi*, Shanxi, Sichuan, Xizang, Yunnan), India (Meghalaya, Sikkim, West Bengal), Japan, South & North Korea, Kuril Islands, Nepal, Thailand.

Host plants

A polyphagous species usually attacking angiosperms, but also recorded from Pinus (Pinaceae) (Murayama 1936; Nobuchi 1966).

Remarks

Published records from India, Nepal, Thailand, and some Chinese provinces may refer to Anisandrus cristatus or A. congruens, with which A. apicalis has been confused previously.

Anisandrus auco sp. nov.

http://zoobank.org/97A9EB18-B9CC-4BDF-91FC-E636111196F5

Fig. 18E, F, K

Type material

Holotype , female, Vietnam: Cao Bang, 22°36.3'N, 105°52.6'E, 1435–1601 m, 13–17.iv.2014, VN16, Cognato, Smith, Pham, ex FIT (MSUC).

Diagnosis

2.9 mm long (n = 1); 2.23× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc flat; declivital interstriae clearly punctate; declivity gradual and convex, posterolateral margins rounded; pronotum rounded when viewed dorsally (type 1); and pronotum armed by four uniformly sized coarse serrations on anterior margin.

Similar species

Anisandrus cryphaloides.

Description

(female). 2.9 mm long (n = 1); 2.23× as long as wide. Body bicolored with pronotal and elytral bases lighter than rest of body. Pronotal and elytral bases, head, legs, and antennae light brown, remainder of elytra red-brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, subshiny, punctate; punctures large, shallow, dense; punctures bearing a long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 as long as pedicel. Club longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 0.85× as long as wide. In dorsal view rounded, type 1, sides convex, rounded anteriorly; anterior margin with a row of four very large, coarse serrations. In lateral view short and tall, type 3, disc as long as anterior slope, summit at midpoint. Anterior slope with densely spaced, very large coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny with moderately dense, large, shallow punctures bearing moderate, semi-recumbent, hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles acutely rounded. Mycangial tuft present along basal margin, tuft moderately setose, approximately the width of scutellum. Elytra: 1.49× as long as wide, 1.75× as long as pronotum. Scutellum broad, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 1/2, then broadly rounded to apex; surface shiny. Disc flat, striae not impressed, with moderately-sized, deep punctures separated by less than one diameter of a puncture, setose, setae as long as two punctures, recumbent, hair-like; interstriae flat, punctate, punctures strongly confused, setose, setae 1× width of interstriae 2, erect, hair-like, unarmed by granules. Declivity occupying approximately 2/5 of elytra, gradually rounded, declivital face convex; striae weakly impressed, strial punctures larger and deeper than those of disc, punctures setose, setae slightly longer than the diameter of a puncture, semi-erect, hair-like; interstriae uniseriate punctate, setae 2× width of interstriae 2, erect, hair-like, interstriae 2 as wide as interstriae 3 at midpoint of declivity. Posterolateral margin rounded, unarmed by granules. Legs: procoxae contiguous. Protibiae obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with six large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with nine and ten small socketed denticles, respectively.

Etymology

Vietnamese mythology, Âu Cơ – mountain fairy that gave birth to the ancestors of the Vietnamese people. Pronunciation – ò-ghá. Noun in apposition.

Distribution

Vietnam.

Host plants

Unknown.

Figure 18.

Dorsal, lateral and declivital view of Anisandrus achaete holotype, 3.5 mm (A, B, I), A. apicalis, 3.05–3.4 mm (C, D, J), A. auco holotype, 2.9 mm (E, F, K), and A. auratipilus holotype, 2.8 mm (G, H, L).

Anisandrus auratipilus sp. nov.

http://zoobank.org/07D53FDF-F903-4459-8B69-EE3C1D1DFEFE

Fig. 18G, H, L

Type material

Holotype , female, China: Fujian, Fuzhou, 18.iii.2018, Y. Li, ex unknown twig (IZAS). Paratypes, female, as holotype (MSUC, 2).

Diagnosis

2.8 mm long (n = 1); 2.15× as long as wide. This species is distinguished by the moderately dense mesonotal mycangial tuft that extends laterally from the scutellum to striae 3; declivital posterolateral margin rounded; elytral disc flat; declivital face moderately steep, convex; declivital interstriae 1 unarmed; declivital summit with a small denticle on interstriae 2 and a minute denticle on interstriae 3; interstriae 3 with three denticles on basal 1/2; declivital striae weakly impressed, punctures small, shallow and seriate; interstriae convex, minutely punctate, punctures strongly confused, setose, setae erect hair-like; body shiny, abundantly covered with long erect hair-like setae; elytral disc finely punctate; and pronotal asperities large, coarse, moderately spaced.

Similar species

Anisandrus apicalis, A. hera, A. klapperichi, A. percristatus, A. venustus, A. xuannu.

Description

(female). 2.8 mm long (n = 1); 2.15× as long as wide. Body bicolored with pronotal and elytral bases lighter than rest of body. Pronotal and elytral bases brown, remainder of elytra and head dark brown. Legs and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, impunctate, median area of with a small ovate smooth, glabrous, strongly shiny area; lateral areas shagreened, weakly rugose, setose, each shallow ruga bearing a long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 0.7× as long as wide. In dorsal view conical, type 0, sides convex, conical anteriorly; anterior margin with a row of four moderate serrations. In lateral view type 3, short and tall, disc as long as anterior slope, summit at midpoint. Anterior slope with moderately spaced, large, coarse, asperities, becoming lower and more strongly transverse towards summit. Disc subshiny with dense, large, shallow punctures bearing short to moderate, erect hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Mycangial tuft present along basal margin tuft broad, moderately setose, laterally extending to elytral striae 3. Elytra: 1.6× as long as wide, 2.26× as long as pronotum. Scutellum broad, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 2/3, then narrowly rounded to apex; surface shiny. Disc flat, striae not impressed, with small, shallow punctures separated by one diameter of a puncture, setose, setae as long as a puncture, semi-recumbent, hair-like; interstriae flat, minutely punctate, punctures strongly confused, setose, setae 1× width of interstriae 2, erect hair-like, unarmed by granules. Declivity occupying approximately 2/5 of elytra, steeply rounded, declivital face convex; striae weakly impressed, strial punctures somewhat larger and deeper than those of disc, and bearing setae as described for disc; interstriae sparsely minutely uniseriate punctate, setae 1–1.5× width of interstriae 2, erect, hair-like, interstriae 2 as wide as interstriae 3 at midpoint of declivity, declivital summit with a small denticle on interstriae 2 and a minute denticle on interstriae 3; interstriae 3 with three denticles on basal 1/2. Posterolateral margin rounded, unarmed by granules. Legs: procoxae contiguous; prosternal coxal piece short, inconspicuous. Protibiae obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with five large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with seven and eight large socketed denticles, respectively.

Etymology

L. auratus = golden; pilus = hair. In reference to the golden setae covering the elytra. Noun in apposition.

Distribution

China (Fujian).

Host plants

Unknown.

Anisandrus carinensis (Eggers, 1923), comb. nov.

Fig. 19A, B, I

Xyleborus carinensis Eggers, 1923: 180.

Type material

Holotype (MCG).

Diagnosis

4.5 mm long (n = 1); 2.25× as long as wide. This species is distinguished by the mesonotal mycangial tuft absent; antennal club type 1 with segment 1 encircling anterior face; elytral disc with a weak transverse saddle-like depression; declivital interstriae 1 and 3 armed by four or five unequally sized tubercles; and a row of serrations on anterior margin of pronotum.

Similar species

Anisandrus achaete.

Distribution

Myanmar.

Host plants

Unknown.

Remarks

The species has the generic characters of Anisandrus and is here transferred to that genus.

Figure 19.

Dorsal, lateral and declivital view of Anisandrus carinensis holotype, 4.5 mm (A, B, I), A. congruens holotype, 2.6–2.8 mm (C, D, J), A. cristatus, 3.35–3.7 mm (E, F, K), and A. cryphaloides holotype, 2.1–2.4 mm (G, H, L).

Anisandrus congruens sp. nov.

http://zoobank.org/BD7E0ACE-07E2-47D2-A3CE-A160BFA5ED5C

Fig. 19C, D, J

Type material

Holotype , female, Vietnam: Cao Bang, 22°36.3'N, 105°52.6'E, 1435–1601 m, 13–17.iv.2014, VN16, Cognato, Smith, Pham, ex FIT (MSUC). Paratypes, female, Thailand: Chiang Mai, Doi Pui, 1400 m, 20–24.xii.2004, W. Puranasakul, ex EtOH trap (RABC, 1); Vietnam: Lao Cai, Hoang Lien N.P., 22.35, 103.77, 1500–2000 m, 19.v.2019, VN169, S.M. Smith, A.I. Cognato (MSUC, 1); as previous except: 18–19.v.2019, ex FIT (MSUC, 1).

Diagnosis

2.6–2.8 mm long (mean = 2.7 mm; n = 2); 2.16–2.36× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc with a weak to moderate transverse saddle-like depression; posterolateral margin costate to interstriae 5; declivity appearing bisulcate, moderately impressed from striae 1 and 2, interstriae 3 strongly inflated, tuberculate from summit to apical 1/4 then becoming flattened and unarmed to apex; and moderate sharp backwardly pointed spine at base of declivital interstriae 2.

This species strongly resembles A. apicalis and A. cristatus and is most easily distinguished by the smaller size, more strongly impressed declivital sulci than A. apicalis and larger spines on interstriae 3 that are sharply pointed but not strongly backwardly hooked.

Similar species

Anisandrus apicalis, A. cristatus, A. geminatus, A. niger, A. sinivali.

Description

(female). 2.6–2.8 mm long (mean = 2.7 mm; n = 2); 2.16–2.36× as long as wide. Body uniformly dark brown, except dark red-brown declivity. Legs and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, alutaceous, subshiny, punctate; punctures large, shallow, setose; punctures bearing a long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 1.0× as long as wide. In dorsal view rounded, type 1, sides convex, rounded anteriorly; anterior margin with a row of four serrations. In lateral view robust and rounded, type 5, disc as long as anterior slope, summit at midpoint. Anterior slope with densely spaced, large coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny, alutaceous with sparse fine punctures bearing short, recumbent, hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles acutely rounded. Mycangial tuft present along basal margin, tuft moderately setose, approximately the width of scutellum. Elytra: 1.5× as long as wide, 1.5× as long as pronotum. Scutellum broad, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 1/2, then broadly rounded to apex; surface shiny. Disc with a weak to moderate medial transverse saddle-like depression, striae not impressed, with small, deep punctures separated by two diameters of a puncture, setose, setae as long as a puncture, recumbent, hair-like; interstriae flat, punctate, punctures uniseriate subequal to those of striae, setose, setae 1× width of interstriae 2, erect, hair-like, unarmed by granules. Declivity occupying approximately 1/2 elytra, evenly rounded, declivital face weakly bisulcate, moderately impressed from striae 1 and 2, interstriae 3 strongly inflated, tuberculate from summit to apical 1/4 then becoming flattened and unarmed to apex; striae not impressed, strial punctures much larger and deeper than those of disc, and bearing setae as described for disc; interstriae impunctate, sparsely minutely granulate, setae 1–2× width of interstriae 2, erect, hair-like, interstriae 2 as wide as interstriae 3 at midpoint of declivity, declivital summit with a moderate sharp backwardly pointed spine at base of declivital interstriae 2. Posterolateral margin costate to interstriae 5. Legs: procoxae contiguous; prosternal coxal piece short, inconspicuous. Protibiae obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with seven large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with eight and ten large socketed denticles, respectively.

Etymology

L. congruens = agreeing with. In reference to its similarity to apicalis and cristatus. A participle.

Distribution

Thailand, Vietnam.

Host plants

Unknown.

Anisandrus cristatus (Hagedorn, 1908) comb. nov., stat. res.

Fig. 19E, F, K

Xyleborus cristatus Hagedorn, 1908: 377.

Xyleborus fabricii Schedl, 1964c: 217. Unnecessary replacement name.

Type material

Syntypes (IRSNB). Not examined.

New records

China: Yunnan, Gaoligong Mts, 24.57; 98.45, 2200–2500 m, 8–16.v.1995, V. Kuban (NHMB, 3; RABC, 1). India: [West Bengal], Darjeeling D[istrict], Rally, 850 m, 3.iv.1979, Bhakta B. (NHMB, 1); as previous except: Lepchajagat 7000 ft, 11.ix.1929, J.C.M. Gardner, ex Symplocos theaefolia (NMNH, 1); as previous except: Rangirum, 6000 ft (NMNH, 1). Laos: NE, Hua Phan, Ban Saluei, Phou Pan (Mt.), 20°12'N, 104°01'E, 1300–1900 m, 7.iv–25.v.2010, C. Holzschuh (NHMUK, 4; RABC, 1). Myanmar: Kambaiti, 7000 ft, 22.iv.1934, R. Malaise (NMNH, 1). Nepal: Arun Valley, Deurali, 27°30'N, 87°16'E, ~ 2100 m NN, 10.v.2014, J. Schmidt (NKME, 3); Koshi, Gorza, 2100 m, 5–6.vi.1985, M. Brancucci (NHMB, 9; RABC, 2); Kathmandu V[alley], Gufa–Gorza, 2800–2100 m, M. Brancucci (NHMB, 4); Koli Gandaki Khola, Chitra, Ghar Khola, 2400 m, Bhakta B. (NHMB, 1); Manaslu Mts, E slope of Ngadi Khola valley, 28°22'N, 84°29'E, 2000–2300 m, 14–16.v.2005, J. Schmidt (RABC, 2). Thailand: Chiang Mai, Doi Inthanon, 5.viii.[20]02, R. A. Beaver (RABC, 1). Vietnam: Cao Bang, 22°36.402'N, 105°52.397'E, 1601 m, 13.iv.2014, VN17, Cognato, Smith, Pham, ex standing stump (MSUC, 1). Lao Cai, Hoang Lien N.P., 22.35, 103.77, 1500–2000 m, 19.v.2019, VN171, S.M. Smith, A.I. Cognato, ex 1 cm DBH dead sapling (MSUC, 1).

Diagnosis

3.35–3.7 mm long (mean = 3.55 mm; n = 5); 2.2–2.47× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc with or without a weak transverse saddle-like depression; declivital posterolateral margin costate to interstriae 5; declivity appearing bisulcate, moderately impressed from striae 1 and 2, interstriae 3 moderately inflated, tuberculate from base to apical 1/4 then becoming flattened and unarmed to apex; and large sized sharp incurved spine on interstriae 2 at base of declivity.

This species strongly resembles A. apicalis and A. congruens and is most easily distinguished by the larger size, more strongly impressed declivital sulci than A. apicalis and larger spines on interstriae 3 that are sharply pointed and backwardly hooked and subequal in size to the spine at the summit of interstriae 2.

Similar species

Anisandrus apicalis, A. congruens, A. geminatus, A. niger, A. sinivali.

Distribution

Bhutan*, China* (Yunnan), India (Meghalaya, ‘Naga Hills’, Sikkim, West Bengal), Laos*, Myanmar*, Nepal*, Thailand*, Vietnam*.

Host plants

This species has been recorded from Alnus (Betulaceae), Quercus (Fagaceae), Symplocos (Symplocaceae) (Beeson 1930).

Remarks

Xyleborus cristatus has the generic characters of Anisandrus and is here transferred to that genus. This species was synonymized with Ambrosiodmus apicalis (Blandford) [sic] by Wood (1989). It is here removed from synonymy and reinstated as a distinct species, based on the characters given above, and differences in DNA (Cognato et al. 2020b).

Anisandrus cryphaloides sp. nov.

http://zoobank.org/7C75FBB1-6168-4E35-BC38-6C62FE3882DD

Fig. 19G, H, L

Type material

Holotype , female, Vietnam: Cao Bang, 22°36.804'N, 105°51.982'E, 1831 m, 17.iv.2014, VN42, Cognato, Smith, Pham, ex 0.3–3 cm twigs/branches (NMNH). Paratypes, female, as holotype (MSUC, 4; NHMUK, 2; NMNH, 2; VMNH 1); Lao Cai, Hoang Lien N.P., 22.35, 103.77, 1500 m, 17.v.2019, VN152, S.M. Smith, A.I. Cognato, ex 1–3 cm branch (MSUC, 3); as previous except: VN153, ex branch; 1–2 cm (MSUC, 1); as previous except: 1500–2000 m, 20.v.2019, VN185, ex branch; 1–2 cm (MSUC, 1); 1500–2000 m, 20.v.2019, VN186, ex branch; 1–2 cm (MSUC, 1).

Diagnosis

2.1–2.4 mm long (mean = 2.26 mm; n = 5); 2.2–2.4× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc convex; declivity gradual and convex, with rounded posterolateral margins; pronotum conical frontally when viewed dorsally (type 0); pronotum armed by four coarse serrations on anterior margin (median pair larger than lateral pair); elytra strongly shagreened or opalescent; and declivital striae weakly impressed.

Similar species

Anisandrus auco.

Description

(female). 2.1–2.4 mm long (mean = 2.26 mm; n = 5); 2.2–2.4× as long as wide. Body dark brown. Antennae and legs light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, alutaceous, subshiny, punctate, punctures large, shallow, setose; punctures bearing a long, erect hair-like seta. Eyes feebly emarginate, almost entire, just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 as long as pedicel. Club longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 0.89× as long as wide. In dorsal view conical, type 0, sides convex, conical anteriorly; anterior margin with a row of four coarse serrations, median pair larger than lateral pair. In lateral view type 3, short and tall, disc as long as anterior slope, summit at midpoint. Anterior slope with moderately spaced, large coarse asperities, becoming lower and more strongly transverse towards summit. Disc strongly shiny with moderately dense, large, shallow punctures bearing moderate, erect, hair-like setae or short, recumbent, hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles acutely rounded. Mycangial tuft present along basal margin, tuft moderately setose, approximately the width of scutellum. Elytra: 1.26× as long as wide, 1.4× as long as pronotum. Scutellum broad, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 1/2, then broadly rounded to apex; surface opalescent to shagreened. Disc convex, striae not impressed, with small, shallow punctures separated by less than one diameter of a puncture, setose, setae as long as two punctures, recumbent, hair-like; interstriae flat, punctate, punctures strongly confused, setose, setae longer than the width of interstriae 2, erect hair-like, unarmed by granules. Declivity occupying approximately 1/2 elytra, gradually rounded, declivital face convex; striae weakly impressed, strial punctures somewhat larger and deeper than those of disc; interstriae sparsely uniseriate punctate, setae 2–3× width of an interstria, erect, hair-like, interstriae 3 narrower than interstriae 2 at midpoint of declivity, interstriae 2 with a small incurved spine at declivital summit. Posterolateral margin rounded, unarmed by granules. Legs: procoxae contiguous; prosternal coxal piece short, inconspicuous. Protibiae obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with six very large socketed denticles, their length much longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with eight very large socketed denticles.

Etymology

Resembling Cryphalus Erichson, 1836, in reference to the coarse asperities in concentric rows on the anterior half of the pronotum. Noun in apposition.

Distribution

Vietnam.

Host plants

Unknown.

Anisandrus dispar (Fabricius, 1792)

Fig. 20A, B, I

Apate dispar Fabricius, 1792: 363.

Anisandrus dispar (Fabricius): Ferrari 1867: 24.

Xyleborus dispar (Fabricius): Hagedorn 1910b: 98.

Anisandrus dispar (Fabricius): Hulcr et al. 2007: 578.

Bostrichus thoracicus Panzer, 1793: 34. Synonymy: Hagedorn 1910b: 102.

Scolytus pyri Peck, 1817: 207. Synonymy: Hubbard 1897: 22; Swaine 1918: 124.

Bostrichus tachygraphus Sahlberg, 1836: 152. Synonymy: Eichhoff 1876b: 378.

Bostrichus ratzeburgi Kolenati, 1846: 39. Synonymy: Ferrari 1867: 27.

Xyleborus ishidai Niisima, 1909: 156. Synonymy: Smith et al. 2018b: 393.

Anisandrus aequalis Reitter, 1913: 81. Synonymy: Knížek 2011: 242.

Anisandrus swainei Drake, 1921: 203. Synonymy: Wood 1957: 403.

Xyleborus dispar rugulosus Eggers, 1922: 17. Synonymy: Schedl 1964d: 314.

Xyleborus cerasi Eggers, 1937: 335. Synonymy: Schedl 1964c: 220.

Xyleborus khinganensis Murayama, 1943: 100. Synonymy: Knížek 2011: 242.

Type material

Holotype Anisandrus swainei (NMNH). Lectotype Xyleborus dispar rugulosus (NMNH). Lectotype Xyleborus ishidai (NIAES). Holotype Xyleborus khinganensis (NMNH).

Diagnosis

3.1–3.5 mm long (mean = 3.4 mm; n = 5); 2.27–2.5× as long as wide. This species is distinguished by the mesonotal mycangial tuft sparse, the length of the scutellum; declivital interstriae uniseriate granulate; discal interstriae with two or three confused rows of punctures; declivital interstriae 1 slightly raised, interstriae 2 and 3 even; declivital face smooth, shiny; and declivital interstrial setae erect, 1.5× the width of an interstria.

Similar species

Anisandrus maiche, A. paragogus, Xylosandrus germanus.

Distribution

Europe and North Africa, through Russia and Central Asia to China (Heilongjiang, Shaanxi), North Korea, and Japan. Introduced to Canada and USA (Wood 1977; Gomez et al. 2018a).

Host plants

Polyphagous attacking both angiosperms and conifers (Wood and Bright 1992; Beaver et al. 2014).

Remarks

The biology of the species is described by Palm (1959), Chararas (1962), Egger (1973), and French and Roeper (1975). Speranza et al. (2009) examine the effects of temperature and rainfall on flight activity. Like many xyleborines, the species is attracted to ethanol (Saruhan and Akyol 2012; Galko et al. 2014). It is an important pest of hazel (Corylus avellana) (Betulaceae) in the Mediterranean area (e.g., Bucini et al. 2005; Saruhan and Akyol 2012), and an occasional pest of fruit trees in the USA (Wood 1982).

Figure 20.

Dorsal, lateral and declivital view of Anisandrus dispar, 3.1–3.5 mm (A, B, I), A. eggersi, 3.1–3.2 mm (C, D, J), A. feronia holotype, 2.9 mm (E, F, K), and A. geminatus, 2.9–3.2 mm (G, H, L).

Anisandrus eggersi (Beeson, 1930)

Fig. 20C, D, J

Xyleborus eggersi Beeson, 1930: 215.

Cyclorhipidion eggersi (Beeson): Maiti and Saha 2004: 105.

Anisandrus eggersi (Beeson): Hulcr et al. 2007: 578.

Type material

Paratypes (FRI, 1; NMNH, 1).

New records

Bhutan: km 87 von Phuntsholing, 22.v.1972, Nat.-Hist. Museum Basel – Bhutan Expedition (NHMB, 1) [Misdetermined by K. E. Schedl as Xyleborus fabricii Schedl]. Thailand: Chiang Mai, Doi Inthanon, 5.viii.[20]02, R.A. Beaver, K. Koivisto (RABC, 1); as previous except: 13.xi.[20]11, W. Sittichaya (RABC, 2). Loei, Phu Hin Rongkla N. Park, Huai Man Daeng Naoi @ trail, 16°57'N, 101°03'E, 17.iii-10.iv.2003, G.W. Courtney, ex malaise trap (MSUC, 2). Vietnam: Cao Bang, 22°36.804'N, 105°51.982'E, 1831 m, 17.iv.2014, Cognato, Smith, Pham, 0.3–3.0 cm twigs/branches (MSUC, 10). Lao Cai, Hoang Lien N.P., 22.35, 103.77, 1500–2000 m, 20.v.2019, VN185, S.M. Smith, A.I. Cognato, ex branch; 1–2 cm (MSUC, 1); as previous except: 20.v.2019, VN194, ex dead sapling; 1 cm at base (MSUC, 1).

Diagnosis

3.1–3.2 mm long (mean = 3.12 mm; n = 5); 2.21–2.29× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc convex; declivity appearing flat when viewed laterally; two or three small tubercles present on basal 1/2 of interstriae 2; declivital posterolateral margin costate to interstriae 5; declivital face strongly shagreened; and declivital interstriae clearly punctate.

Similar species

Anisandrus feronia, A. improbus, A. mussooriensis.

Distribution

Bhutan*, India (West Bengal), Myanmar, Nepal, Thailand*, Vietnam*.

Host plants

Polyphagous, recorded from five genera in five different families (Euphorbiaceae, Lauraceae, Rosaceae, Staphyleaceae, Symplocaceae) (Maiti and Saha 2004).

Remarks

Maiti and Saha (2004) suggest that it is a high-altitude species.

Anisandrus feronia sp. nov.

http://zoobank.org/E32E26AC-AA3F-40BC-B0C7-A72C1F3CEB2D

Fig. 20E, F, K

Type material

Holotype , female, 福建崇安 1500m 芥桔子 1978.V.7 采集者:黄復生 [China: Fujian, Chong’an, 1500 m, 7.v.1978, Shuyong Wang, ex Fortunella margarita] (NMNH). Paratypes, female, as holotype (IZAS, 1; NMNH, 1).

Diagnosis

2.9 mm long (mean = 2.9 mm; n = 3); 2.23× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc flat; declivital interstriae punctate; declivital posterolateral margin carinate to interstriae 5; declivity moderately bisulcate; declivital margins ornamented by only two small sharp incurved spines at the base of interstriae 2 and 3; and declivital interstriae bearing fine erect hair-like setae.

Similar species

Anisandrus eggersi, A. longidens, A. mussooriensis.

Description

(female). 2.9 mm long (mean = 2.9 mm; n = 3); 2.23× as long as wide. Body dark red-brown. Legs and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, alutaceous, subshiny, punctate; punctures large, shallow, setose; punctures bearing a long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 as long as pedicel. Club longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 0.77× as long as wide. In dorsal view rounded, type 1, sides convex, rounded anteriorly; anterior margin with a row of 6–8 serrations. In lateral view short and tall, type 3, disc shorter than anterior slope, summit at basal 2/5. Anterior slope with densely spaced, large coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny with dense, fine punctures bearing moderate, semi-erect hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles acutely rounded. Mycangial tuft present along basal margin, tuft moderately setose, approximately the width of scutellum. Elytra: 1.52× as long as wide, 1.97× as long as pronotum. Scutellum broad, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/5, then narrowly rounded to apex; surface opalescent. Disc weakly convex, striae not impressed, with small, deep punctures separated by approximately one diameter of a puncture, setose, setae as long as two punctures, recumbent, hair-like; interstriae flat, punctate, punctures strongly confused, setose, setae 1–1.5× width of interstriae 2, erect, hair-like, unarmed by granules. Declivity occupying approximately 1/2 elytra, steeply rounded, declivital face moderately bisulcate to interstriae 4; striae not impressed, strial punctures much larger and deeper than those of disc, and bearing setae as described for disc; interstriae minutely uniseriate punctate, setae 1–1.5× width of interstriae 2, erect, hair-like, interstriae 2 as wide as interstriae 3 at midpoint of declivity, declivital margins ornamented by only two small sharp incurved spines at base of interstriae 2 and 3. Posterolateral margin carinate to interstriae 5. Legs: procoxae contiguous; prosternal coxal piece short, inconspicuous. Protibiae obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with seven very large socketed denticles, their length much longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with nine and ten large socketed denticles, respectively.

Etymology

Roman mythology, Feronia – goddess of wildlife, fertility, abundance. Noun in apposition.

Distribution

China (Fujian).

Host plants

Recorded from Fortunella margarita (Rutaceae).

Remarks

Locality labels on the holotype and paratypes are in Chinese and were translated by You Li. An English locality label has been placed on the specimen below the original locality labels.

Anisandrus geminatus (Hagedorn, 1904)

Fig. 20G, H, L

Xyleborus geminatus Hagedorn, 1904: 126.

Amasa geminata (Hagedorn): Wood and Bright 1992: 683.

Anisandrus geminatus (Hagedorn): Beaver and Liu 2018: 537.

Type material

The holotype was destroyed in the bombing of UHZM in World War II (Wood and Bright 1992).

New records

India: Darjeeling, Rangirum, 6000 ft, J.C.M. Gardner, 3.ix.1929, ex misc. timber (NMNH, 1).

Diagnosis

2.9–3.2 mm long (mean = 3.03 mm; n = 3); 2.31–2.37× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc flat; declivital interstriae punctate; and posterolateral margin costate to interstriae 7; declivital face concave; declivital interstriae 2 and 3 each armed with a small sharp incurved spine at the summit; and declivital interstriae bearing erect pointed bristle-like setae.

Similar species

Anisandrus apicalis, A. congruens, A. cristatus, A. niger, A. sinivali.

Distribution

India (West Bengal), Nepal.

Host plants

Unknown.

Anisandrus hera sp. nov.

http://zoobank.org/1155CC1E-4DAF-40B9-8C45-DE700FBA0AF4

Fig. 21A, B, I

Type material

Holotype , female, 四川峨边 1900公尺木合川 1960-VI-29 采集者：殷惠芬 [China: Sichuan, E’bian; 1900 m, 29.vi.1960, Huifen Yin, ex Schima superba] (NMNH).

Diagnosis

3.9 mm long (n = 1); 2.05× as long as wide. This species is distinguished by the dense mesonotal mycangial tuft that extends laterally from the scutellum to striae 3; declivital posterolateral margin obliquely costate to interstriae 5; elytral disc with a weak transverse saddle-like depression; declivital summit with large incurved spine on interstriae 2, interstriae 3 with two additional unequally sized denticles ventrad to large spine; declivity weakly sulcate to interstriae 3; declivital strial punctures large each bearing a recumbent seta, interstriae minutely punctate, punctures uniseriate, setose, setae erect, hair-like; body moderately sized and abundantly covered with long erect hair-like setae; declivity shiny; and pronotal asperities small, coarse, densely spaced.

Similar species

Anisandrus auratipilus, A. klapperichi, A. percristatus, A. venustus, A. xuannu.

Description

(female). 3.9 mm long (n = 1); 2.05× as long as wide. Body dark brown. Legs and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons moderately impressed above epistoma then weakly convex to upper level of eyes, impunctate, median area of with a oval-shaped smooth, glabrous, strongly shiny area; lateral areas shagreened, coarsely rugose, setose; each ruga bearing a long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 as long as pedicel. Club longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 0.89× as long as wide. In dorsal view rounded, type 1, sides convex, rounded anteriorly; anterior margin with a row of six large serrations. In lateral view type 3, short and tall, disc as long as anterior slope, summit at midpoint. Anterior slope with widely spaced, large coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny, median area weakly rugose, lateral areas with dense, large, shallow punctures bearing moderate, erect hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Mycangial tuft present along basal margin tuft broad, densely setose, laterally extending to elytral striae 3. Elytra: 1.0× as long as wide, 1.13× as long as pronotum. Scutellum narrow, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 2/3, then narrowly rounded to apex; surface shiny. Disc with a weak medial transverse saddle-like depression, striae not impressed, with small, shallow punctures separated by 2–4 diameters of a puncture, setose, setae as long as a puncture, recumbent, hair-like; interstriae flat, punctate, punctures strongly confused, setose, setae 1.5× width of interstriae 2, erect, hair-like, unarmed by granules. Declivity occupying approximately 1/2 elytra, evenly rounded, declivital face weakly sulcate to interstriae 3; striae not impressed, strial punctures somewhat larger and deeper than those of disc, and bearing setae as described for disc; interstriae sparsely minutely uniseriate punctate, setae 1–2× width of interstriae 2, erect, hair-like, interstriae 2 narrower than interstriae 3 at midpoint of declivity, declivital summit with a large incurved spine on interstriae 2, interstriae 3 costate with two additional unequally sized denticles ventrad to large spine. Posterolateral margin costate to interstriae 5. Legs: procoxae contiguous; prosternal coxal piece short, inconspicuous. Protibiae distinctly triangular, broadest at apical 9/10; posterior face smooth; apical 1/2 of outer margin with seven large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened; outer margins obliquely triangular with 11 and 14 small socketed denticles, respectively.

Etymology

Greek mythology, Hera – goddess of women, marriage, family, and childbirth. Noun in apposition.

Distribution

China (Sichuan).

Host plants

Recorded from Schima (Theaceae).

Remarks

Locality labels on the holotype are in Chinese and were translated by You Li. An English locality label has been placed on the specimen below the original locality labels.

Figure 21.

Dorsal, lateral and declivital view of Anisandrus hera holotype, 3.9 mm (A, B, I), A. hirtus, 3.4–4.5 mm (C, D, J), A. improbus holotype, 3.3–3.4 mm (E, F, K), and A. klapperichi 5.4–5.6 mm (G, H, L).

Anisandrus hirtus (Hagedorn, 1904)

Fig. 21C, D, J

Xyleborus hirtus Hagedorn, 1904: 126.

Cyclorhipidion hirtum (Hagedorn): Wood and Bright 1992: 700.

Anisandrus hirtus (Hagedorn): Hulcr et al. 2007: 578.

Xyleborus hagedorni Stebbing, 1914: 596 nec Iglesias 1914.

Xyleborus hirtuosus Beeson, 1930: 217. Synonymy: Wood 1989: 175.

Xyleborus hagedornianus Schedl, 1952d: 164. Unnecessary replacement name for hagedorni.

Xyleborus tectonae Nunberg, 1956: 209. Unnecessary replacement name for hagedorni.

Xyleborus hirtipes Schedl, 1969b: 53. syn. nov.

Xyleborus taiwanensis Browne, 1980b: 386. Synonymy: Beaver and Liu 2010: 22.

Type material

China* (Sichuan), India (Uttarakhand, West Bengal), Nepal, Vietnam*.

Host plants

Recorded from Machilus (Lauraceae), Symplocos (Symplocaceae) (Beeson 1930) and Alnus (Betulaceae).

Figure 22.

Dorsal, lateral and declivital view of Anisandrus lineatus, 2.6–3.3 mm (A, B, I), A. longidens, 3.0 mm (C, D, J), A. maiche, 2.2–2.5 mm (E, F, K), and A. mussooriensis paratype, 3.0–3.25 mm (G, H, L).

Anisandrus longidens (Eggers, 1930)

Fig. 22C, D, J

Xyleborus longidens Eggers, 1930: 181.

Anisandrus longidens (Eggers): Hulcr et al. 2007: 578.

Type material

Holotype (FRI), paratype (NHMW, 1).

New records

Vietnam: Lao Cai, Hoang Lien N.P., 22.35, 103.77, 1500–2000 m, 20.v.2019, VN185, S.M. Smith, A.I. Cognato, ex 1–2 cm branch (MSUC, 1); as previous except: 19–20.v.2019, ex FIT (MSUC, 2; NMNH, 1).

Diagnosis

3.0–3.2 mm long (mean = 3.1 mm; n = 2); 2.5–2.83× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; convex elytral disc; declivity weakly bisulcate, margins ornamented by large sharp spines on interstriae 2–7, spine on interstriae 3 the largest; posterolateral margin costate to interstriae 5; and declivital interstriae impunctate.

Similar species

Anisandrus feronia.

Distribution

India (Meghalaya), Vietnam*.

Host plants

Unknown.

Anisandrus maiche (Kurentzov, 1941)

Fig. 22E, F, K

Xyleborus maiche Kurentzov, 1941: 192.

Anisandrus maiche (Kurentzov): Nikulina et al. 2015: 43.

Anisandrus maiche Stark, 1936: 142 [sic]. Hulcr et al. 2007: 578.

Xyleborus maiche Eggers, 1942: 36. Homonym. Synonymy: Pfeffer 1944: 131.

Type material

Syntypes (ZIN). Not examined.

New records

China: Shanghai, Dongchuan, vii–viii.2017, Lei Gao, ex trap w/ querciverol (MSUC, 4). Japan: Honshu, Saitama, Chichibu, Takikawa Catchm., 35°55'N, 138°49'E, 850–1060 m, 6.viii.2013 (RABC, 1).

Diagnosis

2.2–2.5 mm long (mean = 2.3 mm; n = 5); 2.3–2.78× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; declivital interstriae 1–4 uniseriate denticulate; discal interstriae punctures uniseriate; declivity appearing bisulcate with impressed from striae 1 to interstriae 2, interstriae 3 distinctly raised; declivital punctures small, uniseriate; shiny appearance; and small body size.

Similar species

Anisandrus dispar, A. paragogus, Xylosandrus germanus.

Distribution

China (Heilongjiang, Shanghai*), Japan*, South & North Korea, Russia (European (introduced), Far East), Ukraine. Introduced to USA (Rabaglia et al. 2009; Gomez et al. 2018a).

Host plants

Polyphagous, recorded from eight families of trees (Rabaglia et al. 2009).

Remarks

Kurentzov (1941) and Terekhova and Skrylnik (2012) provide information on the biology and gallery system, which are similar to A. dispar (see above).

Preliminary phylogenies suggest that Anisandrus maiche is sister to Xylosandrus (Cognato et al. 2020b). Kurenzov (1941) provided the first valid description of Xyleborus maiche rather than Eggers (1942), which has been widely and incorrectly cited in the literature (Nikulina et al. 2015).

Anisandrus mussooriensis (Eggers, 1930)

Fig. 22G, H, L

Xyleborus mussooriensis Eggers, 1930: 179.

Cyclorhipidion mussooriense (Eggers): Maiti and Saha 2004: 116.

Anisandrus mussooriensis (Eggers): Beaver and Liu 2018: 538.

Type material

Holotype (FRI), cotype (NMNH, 1).

Diagnosis

3.0–3.25 mm long (mean = 3.1 mm; n = 5); 2.3–2.33× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc flat; declivital interstriae clearly punctate; declivital posterolateral margin carinate to interstriae 5; declivity appearing flat when viewed laterally, weakly depressed below lateral margins; and basal 1/2 of declivital interstriae 2 with two or three small tubercles.

Similar species

Anisandrus eggersi, A. feronia, A. improbus.

Distribution

India (Uttarakhand), Nepal.

Host plants

Recorded only from Berberis (Berberidaceae) (Beeson 1930).

Anisandrus niger (Sampson)

Fig. 23A, B, I

Xyleborus niger Sampson, 1912: 247.

Anisandrus niger (Sampson): Beaver and Liu 2018: 538.

Type material

Holotype (NHMUK).

New records

Laos: NE, Houa Phan, Ban Saluei, Phou Pan Mt, 20°12–13.5'N, 103°59.5–104°01'E, 1340–1780 m, 15.iv.–15.v.2008, Lao collectors (RABC, 1); as previous except: 20°12'N, 104°01'E, 1300–1900 m, 7.iv.–25.v.2010, C. Holzschuh (RABC, 1). Vietnam: Cao Bang, 22°36.3'N, 105°52.6'E, 1435–1601 m, 13–17.iv.2014, VN16, Cognato, Smith, Pham, FIT (MSUC, 1).

Diagnosis

5.8–5.9 mm long (mean = 5.87 mm; n = 3); 2.0–2.19× as long as wide. This species is distinguished by its large size, mesonotal mycangial tuft the length of the scutellum; elytral disc convex; declivital interstriae impunctate; elytral surface smooth, shiny to weakly shagreened; declivital face flattened when viewed laterally; declivity appearing weakly bisulcate; declivital interstriae 2 weakly impressed, declivital interstriae 1 and 3 tuberculate to apex, interstriae 2 with a tubercle at summit and three or four irregularly spaced granules along its length; and declivital posterolateral margin rounded.

Similar species

Anisandrus apicalis, A. congruens, A. cristatus, A. geminatus, A. sinivali.

Distribution

Laos*, Myanmar, Nepal, Vietnam*.

Host plants

Unknown.

Figure 23.

Dorsal, lateral and declivital view of Anisandrus niger, 5.8–5.9 mm (A, B, I), A. paragogus holotype, 2.8 mm (C, D, J), A. percristatus, 5.5 mm (E, F, K), and A. sinivali holotype, 3.9 mm (G, H, L).

Anisandrus paragogus sp. nov.

http://zoobank.org/CB28B458-4610-45AB-9C89-1EFA27E88EBF

Fig. 23C, D, J

Type material

Holotype , female, 西藏 73084 察隅洞穷1973.7.15 桢楠采集者 : 黄复生 [China: Tibet [Xizang], Dongqiong, Chayu, 15.vii.1973, Fusheng Huang, ex Machilus sp.] (NMNH).

Diagnosis

2.8 mm long (n = 1); 2.55× as long as wide. This species is distinguished by the mesonotal mycangial tuft absent; declivital interstriae uniseriate granulate on basal 1/2; declivital face opalescent; declivital interstrial setae erect, 3× width of an interstria; and a row of serrations on anterior margin of pronotum.

Similar species

Anisandrus dispar, A. maiche, Xylosandrus germanus.

Description

(female). 2.8 mm long (n = 1); 2.55× as long as wide. Body brown. Legs and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, subshiny, punctate; punctures large, shallow, moderately dense; punctures bearing a long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, shorter than club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club much longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 0.73× as long as wide. In dorsal view rounded, type 1, sides convex, rounded anteriorly; anterior margin with a row of seven very large serrations. In lateral view robust and rounded, type 5, disc longer than anterior slope, summit at apical 2/5. Anterior slope with densely spaced, large coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny with dense, small, fine punctures bearing short erect hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Mycangial tuft absent. Elytra: 1.6× as long as wide, 2.2× as long as pronotum. Scutellum broad, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then broadly rounded to apex. Disc flat, opalescent, striae not impressed, with small, shallow punctures separated by 1–2 diameters of a puncture, setose, setae short, in-curved, hair-like; interstriae flat, punctate, punctures strongly confused, setose, setae long, erect hair-like, unarmed by granules. Declivity occupying approximately 1/3 of elytra, steeply rounded, declivital face convex, opalescent; striae distinctly impressed, strial punctures much larger and deeper than those of disc; interstriae impunctate, granulate, granules widely and regularly spaced from base to apex, granules setose, setae 3× width of interstriae 2, erect, hair-like, interstriae weakly laterally broadened from declivital summit to midpoint then narrowed to apex. Posterolateral margin costate, granulate to interstriae 7. Legs: procoxae narrowly separated. Protibiae obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with five large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with nine and ten moderate socketed denticles, respectively.

Etymology

G. paragogos = misleading. In reference to its resemblance to Ambrosiophilus.

Distribution

China (Xizang).

Host plants

Recorded only from Machilus (Lauraceae).

Remarks

Locality labels on the holotype are in Chinese and were translated by You Li. An English locality label has been placed on the specimen below the original locality labels.

Anisandrus percristatus (Eggers, 1939), comb. nov.

Fig. 23E, F, K

Xyleborus percristatus Eggers, 1939a: 12.

Type material

Paratype (NMNH, 1).

New records

China: Sichuan, E’bian, 1900 m, 2.vi.1960, Huifen Yin, ex Schima superba (NMNH, 1).

Diagnosis

5.5 mm long (mean = 5.5 mm; n = 3); 2.12–2.2× as long as wide. This species is distinguished by the dense mesonotal mycangial tuft that extends laterally from the scutellum to striae 3; declivital posterolateral margin carinate to interstriae 5; elytral disc with a profound transverse saddle-like depression; declivital base with very large incurved spine on interstriae 3, interstriae 3 with four additional equally sized and spaced denticles; declivity broadly sulcate to interstriae 5; elytral disc sulcate anteriad to spine on interstriae 3; large body size; body shiny, appearing polished, largely glabrous, minutely punctate; declivital punctures confused; and pronotal asperities very broad, fine, widely spaced.

Similar species

Anisandrus auratipilus, A. hera, A. klapperichi, A. venustus, A. xuannu.

Distribution

China (Sichuan, Yunnan), Myanmar.

Host plants

Recorded from Schima superba (Theaceae).

Remarks

This species is transferred to Anisandrus because of the visible scutellum, pronotal base with a large, dense setal tuft (indicating a mesonotal mycangium), contiguous procoxae; antennal club type 1, taller than wide, and protibiae triangular.

Anisandrus sinivali sp. nov.

http://zoobank.org/FEDF5FDF-F95F-4201-A6FC-4BF2707A5FEC

Fig. 23G, H, L

Type material

Holotype , female, India: Bengal [West Bengal], Kalimpong, Samsingh, 7.x.1933, C.F.C. Beeson (NMNH).

Diagnosis

3.9 mm long (n = 1); 2.29× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc with a weak transverse saddle-like depression; declivity posterolateral margins rounded; elytral surface opalescent; declivital interstriae 2 armed with a blunt tubercle at summit, interstriae 3 armed by one or two denticles near declivital summit ventrad to tubercle on interstriae 2; declivital face convex, evenly rounded toward apex; and pronotal disc feebly asperate.

Similar species

Anisandrus apicalis, A. congruens, A. cristatus, A. geminatus, A. niger.

Description

(female). 3.9 mm long (n = 1); 2.29× as long as wide. Body dark brown. Legs and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, finely reticulate, sparsely finely punctate; punctures bearing a long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed Antennal scape regularly thick, longer than club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 longer than pedicel. Club longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 0.86× as long as wide. In dorsal view rounded, type 1, sides convex, rounded anteriorly; anterior margin with a row of five large serrations. In lateral view type 3, short and tall, disc as long as anterior slope, summit at midpoint. Anterior slope with densely spaced, large coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny, impunctate, feebly asperate, basal and lateral areas densely finely punctate, each puncture bearing moderate, erect, hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Mycangial tuft present along basal margin, tuft densely setose, approximately the width of scutellum. Elytra: 1.06× as long as wide, 1.24× as long as pronotum. Scutellum broad, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 1/2, then broadly rounded to apex; surface opalescent. Disc with a weak medial transverse saddle-like depression, striae not impressed, with small, shallow punctures separated by less than one diameter of a puncture, setose, setae as long as two punctures, recumbent, hair-like; interstriae flat, punctate, punctures strongly confused, setose, setae 2–3× width of interstriae 2, erect hair-like, unarmed by granules. Declivity occupying approximately 1/2 elytra, evenly rounded, declivital face convex; striae weakly impressed, strial punctures somewhat larger and deeper than those of disc; interstriae sparsely uniseriate punctate, setae 2–4× width of interstriae 2, erect, hair-like, interstriae 2 narrower than interstriae 3 at midpoint of declivity, declivital interstriae 2 armed with a blunt tubercle at summit, interstriae 3 armed by one or two denticles near declivital summit ventrad to tubercle on interstriae 2. Posterolateral margin rounded, unarmed by granules. Legs: procoxae contiguous. Protibiae obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with seven very large socketed denticles, their length much longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with at least five and seven large socketed denticles, respectively.

Etymology

Hindu mythology, Sinivali – goddess of fecundity. Pronunciation – Sinivālī. Noun in apposition.

Distribution

India (West Bengal).

Host plants

Unknown.

Remarks

The holotype is card mounted obscuring ventral characters, including mesotibial denticles.

Anisandrus ursulus (Eggers, 1923)

Fig. 24A, B, G

Xyleborus ursulus Eggers, 1923: 173.

Xylosandrus ursulus (Eggers): Wood and Bright 1992: 801.

Anisandrus ursulus (Eggers): Dole and Cognato 2010: 527.

Type material

Holotype (SDEI). Not examined.

New records

China: Guangdong, W of Qixing, Heishiding nature reserve, 27°27.9'N, 111°54.3'E, 190 m, forested stream valley, at light, 1–3.v.2011, M. Ficáček, J. Hájek (MNHP, 1). Guangxi A. R., Longsheng hot spring, 25°53.6'N, 110°12.4'E, 360 m, forested river valley, wet rocks, M. Ficáček, J. Hájek, J. Růžička (MNHP, 1). Jiangxi, Long Nan, 12.vii.2016, Lv-Jia, Lai, S-C., ex Cyclobalanopsis glauca (RABC, 1).

Diagnosis

4.3–4.9 mm long (mean = 4.5 mm; n = 5); 1.88–1.96× as long as wide. This species is distinguished by the mesonotal mycangial tuft the length of the scutellum; elytral disc convex; declivity obliquely truncate with lateral margins obliquely costate; declivity opalescent and unarmed; declivital striae not impressed; body stout and densely covered by erect dark brown pubescence.

Similar to Cnestus mutilatus and Hadrodemius species but declivity less steeply truncate, with posterolateral margins rounded, never carinate, procoxae contiguous and the mesonotal mycangial tuft the length of the scutellum.

Similar species

Anisandrus hirtus, Cnestus ater, C. mutilatus, Hadrodemius spp.

Distribution

China (Fujian, Guangdong*, Guangxi*, Jiangxi*), India (Nicobar Is, West Bengal), Indonesia (Bali, Batoe Is, Java, Maluku, Sulawesi, Sumatra), Laos, East & West Malaysia, New Guinea, Philippines, Solomon Islands, Thailand, Vietnam.

Host plants

The species is polyphagous (Browne 1961b).

Figure 24.

Dorsal, lateral and declivital view of Anisandrus ursulus, 4.3–4.9 mm (A, B, G), A. venustus holotype, 3.1 mm (C, D, H), and A. xuannu holotype, 4.0–4.15 mm (E, F, I).

Anisandrus venustus sp. nov.

http://zoobank.org/719C65BA-BB8D-4343-B729-2F4ED776C0AD

Fig. 24C, D, H

Type material

Holotype , female, Taiwan: Taichung, Heping Dist., 2.iv.2014, C.-S. Lin (TARI). Paratypes, female, as holotype (MSUC, 2; NHMUK, 1; NMNH, 1); Yilan Co., Chilan cypress forest trail, 12.5K, EtOH+pinene, 22.xii.2018, Liu, Lan-Yu (LLYC, 2; RABC, 1).

Diagnosis

3.1 mm long (mean = 3.1 mm; n = 4); 2.38× as long as wide. This species is distinguished by the dense mesonotal mycangial tuft that extends laterally from the scutellum to striae 3; declivital posterolateral margin rounded; elytral disc with a broad, weak transverse saddle-like depression; declivital summit with a small denticle on interstriae 2 and a minute denticle on interstriae 1, interstriae 3 unarmed; declivital strial punctures large, seriate, each bearing a recumbent seta, interstriae flat, minutely punctate, punctures strongly confused, setose, setae hair-like, erect; declivity opalescent; elytral disc shiny and finely punctate; body abundantly covered with long erect hair-like setae; and pronotal asperities large, coarse, moderately spaced.

Similar species

Anisandrus apicalis, A. auratipilus, A. hera, A. klapperichi, A. percristatus, A. xuannu.

Description

(female). 3.1 mm long (mean = 3.1 mm; n = 4); 2.38× as long as wide. Body bicolored with pronotal and elytral bases lighter than rest of body. Pronotal and elytral bases brown, remainder of elytra and head dark brown. Legs and antennae light brown. Head: epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, impunctate, median area with a broad diamond-shaped smooth, glabrous, strongly shiny area; lateral areas shagreened, weakly rugose, setose; each shallow ruga bearing a long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, shorter than length of club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 shorter than pedicel. Club longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 0.64× as long as wide. In dorsal view conical, type 0, sides convex, conical anteriorly; anterior margin with a row of 6–8 moderate serrations. In lateral view short and tall, type 3, disc as long as anterior slope, summit at midpoint. Anterior slope with moderately spaced, large, coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny with dense, large, fine punctures bearing short to moderate, erect hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Mycangial tuft present along basal margin tuft broad, densely setose, laterally extending to elytral striae 3. Elytra: 1.44× as long as wide, 2.23× as long as pronotum. Scutellum broad, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 3/4, then narrowly rounded to apex; surface shiny. Disc shiny, with a broad, weak transverse saddle-like depression behind declivital summit, striae not impressed, with moderate, shallow punctures separated by 1–2 diameters of a puncture, setose, setae as long as a puncture, recumbent, hair-like; interstriae flat, minutely punctate, punctures strongly confused, setose, setae 1–1.5× width of interstriae 2, erect hair-like, unarmed by granules. Declivity occupying approximately 1/3 of elytra, steeply rounded, declivital face flattened, opalescent; striae not impressed, strial punctures much larger and deeper than those of disc, and bearing setae 2× as long as those of disc; interstriae densely minutely punctate, punctures strongly confuses, setose, setae 1–1.5× width of interstriae 2, erect, hair-like, interstriae 2 as wide as interstriae 3 at midpoint of declivity, declivital summit with a small denticle on interstriae 2 and a minute denticle on interstriae 1, interstriae 3 unarmed. Posterolateral margin rounded, unarmed. Legs: procoxae contiguous; prosternal coxal piece short, inconspicuous. Protibiae obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with six large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with seven large, narrow socketed denticles.

Etymology

L. venustus = like Venus, lovely, beautiful, elegant, graceful. An adjective.

Distribution

Taiwan.

Host plants

Unknown.

Anisandrus xuannu sp. nov.

http://zoobank.org/4FFF4E2C-330F-4A01-9EC2-5496FC0A5B73

Fig. 24E, F, I

Type material

Holotype , female, 四川 : 峨眉山洪椿坪295 1964-V-12 采集者 : 黄复生 [China: Sichuan, Hongchunping, Emeishan Mt., 12.v.1964, Fusheng Huang, ex Fagaceae] (NMNH). Paratypes, female, China: Chongqing, Simian Shan, 7.v.2016, Tian-Shang, Lv-Jia (RABC, 1); Sichuan, Mt. Emei, 600–1050 m, 5–19.v.1989, L. Bocák (RABC, 1).

Diagnosis

4.0–4.15 mm long (mean = 4.08 mm; n = 3); 2.0–2.31× as long as wide. This species is distinguished by the dense mesonotal mycangial tuft that extends laterally from the scutellum to striae 3; declivital posterolateral margin costate to interstriae 5; elytral disc with a deep transverse saddle-like depression, depressed area sulcate; declivital summit with large incurved spine on interstriae 2, interstriae 3 unarmed; declivity moderately sulcate to interstriae 4; declivital strial punctures large, seriate, interstriae minutely biseriately punctate, setose, setae short erect bristle-like; moderate body size; declivity shagreened; elytral disc rugose; body abundantly covered with long erect hair-like setae; and pronotal asperities small, coarse, densely spaced.

Similar species

Anisandrus auratipilus, A. hera, A. klapperichi, A. percristatus, A. venustus.

Description

(female). 4.0–4.15 mm long (mean = 4.08 mm; n = 3); 2.0–2.31× as long as wide. Body bicolored with pronotal and elytral bases lighter than rest of body. Pronotal and elytral bases, head, legs, and antennae light brown, remainder of elytra red-brown. Head: epistoma entire, transverse, with a row of hair-like setae. Epistoma entire, transverse, with a row of hair-like setae. Frons weakly convex to upper level of eyes, impunctate, shagreened, weakly rugose, setose; each shallow ruga bearing a long, erect hair-like seta. Eyes shallowly emarginate just above antennal insertion, upper part smaller than lower part. Submentum large, distinctly triangular, slightly impressed. Antennal scape regularly thick, as long as club. Pedicel as wide as scape, shorter than funicle. Funicle 4-segmented, segment 1 as long as pedicel. Club longer than wide, obliquely truncate, type 1; segment 1 corneous, encircling anterior face; segment 2 narrow, concave, corneous on anterior face only; sutures absent on posterior face. Pronotum: 0.78× as long as wide. In dorsal view rounded, type 1, sides convex, rounded anteriorly; anterior margin with a row of six very large serrations. In lateral view type 3, short and tall, disc as long as anterior slope, summit at midpoint. Anterior slope with densely spaced, large coarse asperities, becoming lower and more strongly transverse towards summit. Disc subshiny, median area impunctate, reticulate, lateral areas with dense, small, shallow punctures bearing moderate, erect hair-like setae, some longer hair-like setae at margins. Lateral margins obliquely costate. Base transverse, posterior angles broadly rounded. Mycangial tuft present along basal margin tuft broad, densely setose, laterally extending to elytral striae 3. Elytra: 1.45× as long as wide, 1.86× as long as pronotum. Scutellum broad, large, linguiform, flush with elytra, flat, shiny. Elytral base transverse, edge oblique, humeral angles rounded, parallel-sided in basal 1/2, then broadly rounded to apex. Disc rugose, shiny, with a deep transverse saddle-like depression just behind declivital summit, depressed area sulcate; striae not impressed, with small, shallow punctures separated by two diameters of a puncture, setose, setae as long as a puncture, recumbent, hair-like; interstriae flat, punctate, punctures strongly confused, setose, setae 1× width of interstriae 2, erect hair-like, unarmed by granules. Declivity occupying approximately 1/2 elytra, evenly rounded, declivital face nearly flat, moderately sulcate to interstriae 4, shagreened; striae not impressed, strial punctures much larger and deeper than those of disc, and bearing setae as described for disc; interstriae minutely biseriately punctate, setose, setae short, erect, bristle-like, interstriae 2 as broad as interstriae 3 at midpoint of declivity, declivital summit with large incurved spine on interstriae 2, interstriae 3 unarmed; lateral margins of declivity densely setose with very long, erect hair-like setae 2–4× width of interstriae 2. Posterolateral margin costate to interstriae 5. Legs: procoxae contiguous. Protibiae obliquely triangular, broadest at apical 1/3; posterior face smooth; apical 1/2 of outer margin with six large socketed denticles, their length longer than basal width. Meso- and metatibiae flattened; outer margins evenly rounded with at least eight large socketed denticles.

Etymology

Chinese mythology, Xuannü “mysterious lady”- the goddess of fertility. Noun in apposition.

Distribution

China (Chongqing, Sichuan).

Host plants

Recorded from Fagaceae.

Remarks

The holotype is point mounted with an excessive amount of opaque glue which obscures the examination of ventral characters. Locality labels on the holotype are in Chinese and were translated by You Li. An English locality label has been placed on the specimen below the original locality labels.

Arixyleborus Hopkins, 1915

Arixyleborus Hopkins, 1915a: 59.

Xyleboricus Eggers, 1923: 212. Synonymy: Schedl 1952d: 162.

Type species

Arixyleborus rugosipes Hopkins, 1915a; original designation.

Diagnosis

1.35–5.2 mm, 2.0–3.5× as long as wide. Arixyleborus is distinguished by the elytra with distinctive deep strial furrows and interstrial ridges, ridges either granulate or carinate (three species without). Arixyleborus can be further diagnosed by the obliquely truncate antennal club with segment 1 almost covering the posterior face (type 2), club wider than long or as long as wide; protibiae slender or evenly rounded, posterior face flat and unarmed or inflated and granulate; scutellum variable either flush with elytra and flat, flush with elytra and medially impressed or flat and depressed below elytra; elytra from dorsal view typically angulate apically, rarely rounded; mycangial tufts absent; and procoxae contiguous.

Arixyleborus is similar to Stictodex with which it shares a broad antennal club but which lacks the distinctive elytral ridges and furrows. In addition, Arixyleborus has declivital striae 1 parallel to the suture while in Stictodex they are not parallel but undulating.

Similar genera

Cnestus, Pseudowebbia, Stictodex, Truncaudum, Webbia.

Distribution

Distributed throughout tropical Asia and Oceania.

Gallery system

An unbranched radial or curved entrance tunnel, sometimes with a few branches. As the larvae develop, their feeding activity extends part of the main gallery into a single longitudinal brood chamber usually approximately rectangular in shape, and the width of the main gallery (Browne 1961b).

Key to Arixyleborus species (females only)

1	Posterior face of protibiae inflated and granulate; scutellum flush with elytra and flat; lateral margin of pronotum costate or carinate	2
–	Posterior face of protibiae flat and unarmed; scutellum flush with elytra and medially impressed or depressed below level of elytra; lateral margin of pronotum oblique	15
2	Declivital posterolateral carina forming a circumdeclivital ring; lateral profile of declivity appearing truncate; pronotum from dorsal view type 8, with disc very long compared to anterior slope	*resecans*
–	Declivital posterolateral costa extending to interstriae 7; lateral profile of declivity appearing rounded or obliquely truncate; pronotum from dorsal view type 7, with disc as long or slightly longer than anterior slope	3
3	Anterior margin of pronotum viewed from above slightly angularly projecting, the asperities on the margin distinctly larger than those on the anterior slope, and separated from them by the height of a serration or more (Fig. 25C)	4
–	Anterior margin of pronotum viewed from above evenly rounded, the asperities on the anterior margin not distinctly larger than those on the anterior slope, and separated from them by the less than the height of a serration (Fig. 27E)	8
4	Smaller, 1.35–1.5 mm; dorsal profile of elytral apex rounded; elytral posterolateral costa denticulate	*tuberculatus*
–	Larger, 1.9–3.5 mm; dorsal profile of elytral apex angulate; elytral posterolateral costa carinate and unarmed	5
5	Larger, 3.2–3.5 mm; pronotal disc rugose; lateral margin of pronotum carinate	*grandis*
–	Smaller, 1.9–2.2 mm; pronotal disc punctate; lateral margin of pronotum costate	6
6	Declivital face without strial furrows and interstrial ridges below	*leprosulus*
–	Declivital face with strial furrows and interstrial ridges at least to midpoint	7
7	Declivital strial furrows at least 1.5× the width of interstrial ridges on disc; interstrial ridges denticulate, setose, setae recumbent, hair-like, as long as striae 2 with at declivital base; striae strongly impressed; declivity weakly shagreened, interstrial ridges almost appear shiny (Fig. 26E)	*malayensis*
–	Declivital strial furrows equal in width to interstrial ridges on disc; interstrial ridges finely tuberculate, glabrous or with minute setae no longer than 1/2 width of a strial furrow; striae moderately impressed; declivity strongly shagreened (Fig. 30E)	*yakushimanus*
8	Posterolateral declivital costa carinate and unarmed	9
–	Posterolateral declivital costa acute or not, armed with granules or denticles	10
9	Declivity with odd interstriae more strongly elevated than even interstriae; declivital interstriae minutely and equally denticulate	*minor*
–	Declivital interstriae 1 strongly elevated on apical 1/2, other interstriae similarly elevated; declivital interstriae 1 denticulate, denticles very large, denticles on remaining interstriae greatly reduced and less abundant	*suturalis*
10	Elytral strial furrows and interstrial ridges of striae and interstriae 1–3 anteriorly extending no further than apical 1/3 of disc (Fig. 26L)	11
–	Elytral strial furrows and interstrial ridges of striae and interstriae 1–3 anteriorly extending at least to midpoint of disc (Fig. 27K)	12
11	More elongate form, 2.9–3.3× as long as wide; more elongate pronotum (1.3 × longer than wide; declivity with short coarse setae	*mediosectus*
–	Less elongate form, 2.6–2.7× as long as wide; less elongate pronotum (1.1–1.2× longer than wide; declivity with fine hair-like setae	silvanus sp. nov.
12	Elytral strial furrows and interstrial ridges anteriorly extending to apical 1/4 of disc; interstriae densely setose with long hair-like setae and bristles	*rugosipes*
–	Elytral strial furrows and interstrial ridges anteriorly extending just beyond the midpoint of disc; interstriae lightly setose, nearly glabrous	13
13	Declivity interstriae 1–3 strongly and uniformly convex from base to apex	*nudulus*
–	Declivity interstriae 1–3 feebly convex, convexity variably decreasing from base to apex	14
14	Antennal club as wide as long; larger 2.2 mm; elytra 1.35× longer than pronotum	phiaoacensis sp. nov.
–	Antennal club wider than long; smaller, 2.0 mm; elytra 1.24× longer than pronotum	crassior sp. nov.
15	Elytral disc with a transverse saddle-like depression (Fig. 30B)	16
–	Elytral disc flat, without a transverse saddle-like depression (Fig. 29F)	17
16	Larger, 5.2 mm; scutellum depressed below level of elytra and flat	titanus sp. nov.
–	Smaller, 2.8–3.0 mm; scutellum flush with elytra and medially impressed	*granifer*
17	Striae and interstriae on disc never forming strial furrows (Fig. 29E)	18
–	Striae and interstriae on disc forming deep strial furrows and interstrial ridges (Fig. 28A)	20
18	Declivital interstrial granules large, widely spaced and uniseriate	*hirsutulus*
–	Declivital interstriae granules small, densely spaced and confused	19
19	Elytral interstriae bearing two rows of long thick semi-erect hair-like setae; shallow strial furrows on declivity	sittichayai sp. nov.
–	Elytral interstriae bearing one row of short erect black bristles and longer semi-erect hair-like setae; strial furrows never present on declivity	*granulifer*
20	Discal interstriae with tubercles larger than those on the declivity	*scabripennis*
–	Discal and declivital interstriae with multiple rows of confused tubercles of equal size	21
21	Discal striae deeply impressed; elytral interstriae with at least two rows of tubercles and long erect, fine hair-like setae, setae 2× the width of an interstria	*puberulus*
–	Discal striae weakly impressed; elytral interstriae with two rows of granules and long semi-recumbent fine hair-like setae, setae 1–1.5× the width of an interstria	22
22	Elytral vestiture comprised of only hair-like setae on both disc and declivity, setae long, fine, and semi-recumbent	*moestus*
–	Elytral vestiture comprised of hair-like setae and golden scales, long semi-recumbent fine hair-like setae on disc; declivital interstriae densely covered by two or three rows of dense, confused golden scales