Japanese Journal of Systematic Entomology, 21 (1): 35–58. May 30, 2015.

The Family (Coleoptera: Derodontoidea) of Japan and Taiwan

Hiroyuki YOSHITOMI1), Toshio KISHIMOTO2)* and Chi-Feng LEE3) 1)Ehime University Museum, Bunkyo 3, Matsuyama, 790-8577 Japan E-mail: [email protected] 2)Japan Wildlife Research Center, Koutoubashi 3-3-7, Sumida-ku, Tokyo, 130-8606 Japan * Present address: Museum of Natural and Environmental History, Shizuoka, Ooya 5762, Suruga-ku, Shizuoka, 422-8017 Japan 3)Applied Zoology Division, Taiwan Agricultural Research Institute, 189 Chung-Cheng Road, Wufeng 41362, Taichung, Taiwan E-mail: [email protected]

Abstract The Japanese and Taiwanese species of the genus is revised. The genus is subdivided into two subgenera: Nosodendron and Dendrodipnis. Four species are recognized, including two new species, Nosodendron (D.) ogasawaraense and N. (D.) taiwanense, described in this paper. The larvae of three species, N. (N.) asiaticum, N. (D.) coenosum, and N. (D.) ogasawar- aense, are described. Key words: new species, larvae, distribution, associated plants, filter feeding.

Introduction recorded (Háva, 2014), and the adults and larvae of these species had not been reviewed. In the present paper, we The adaptation to aquatic habitats by members of review the Japanese and Taiwanese species of the genus Coleoptera evolved independently in some clades, and Nosodendron, with biological notes and descriptions of the there are various degrees of adaptation (Jäch, 1998). The larval stages. families of water (= water adapted beetles) are known from three suborders: (all families), Materials and Methods (mainly Dytiscoidea), and (part of the family in , , , and so on). In General observations and dissections were made under the derivative group of Polyphaga (i.e., , a Leica MZ95 stereomicroscope. Microstructures of the Chrysomeroidea, and “Cucujioidea”), whose members are dissected parts were studied in pure glycerine under an mostly phytophagy, saprophagy, mycophagy, and detritophagy, Olympus BH-2 compound microscope. After observation, aquatic adaptation is rare except for aquatic plant feeders. the dissected parts were mounted on the same card with the The family Nosodendridae (wounded-tree beetles) specimen. Photographs (Figs. 1, 9, 10) were taken under a inhabits tree sap, and contains 82 species under three genera Leica MZ95 and combined in Helicon® Focus ver. 4.70.5 Pro from all over the world (Háva, 2005, 2014). The genus (Helicon Soft® Limited). Some structures were observed with Nosoglobulus Háva, 2003 includes 3 species from the Oriental a SEM (Hitachi S-225), after coating with gold (Figs. 2–6), Region, the genus Nosotetocus Scudder, 1892 includes 3 fossil and a digital microscope HiROX KH-1300, and images were species from the Miocene in Colorado, and the remaining captured with the 2D measurement software SHX-13M ver. Nosodendron is the most diverse genus (75 present and 1 2.9.0 (Fig. 11). fossil species) distributed in all zoogeographical Region. Morphological terminology follows Leschen & Beutel Considering their habitat (tree sap and slime flux), the family (2010), Lawrence (1991), and Kiselyova & McHugh (2006). adapts to a kind of aquatic habitat, but there are few studies of its behavior and morphology. BPBM: Bishop Museum, Honolulu, Hawaii The phylogenetic positions of the family Nosodendridae EUMJ: Ehime University Museum, Matsuyama, Japan and superfamily Derodontoidea are uncertain. Based on KUMJ: Kyushu University Museum, Japan the phylogenetic analysis of the adult morphology, Beutel NSMT: National Science Museum, Tsukuba (1996) showed this family is a sister group of the family TARI: Taiwan Agricultural Research Institute, Taichung, . Lawrence et al. (2011), analyzing numerous Taiwan characters of the adults and larvae, showed this family has TUAJ: Tokyo University of Agriculture, Atsugi, Japan Morphological abbreviations used in the measurements a sister group relationship with the superfamily Scirtoidea are as follows: EL — length of elytra in suture; EW — (excluding ), and this group plus Derodontidae are maximum width of elytra; HL — length of head; PL — situated in the basal clade of Tenebrionoidea + length of pronotum in median line; PW — maximum width + + Curculionidae. Also, molecular of pronotum; TL — total length (HL+PL+EL). The average is phylogenetic studies could not clarify the taxonomic status given in parentheses after the range. of Nosodendridae in the suborder Polyphaga. The family was situated in the basal-most taxon in Polyphaga except for Derodontoidea, Scirtoidea and Historoidea (Hunt et al., 2007), Systematics or showed a sister group relationship with Genus Nosodendron Latreille, 1804 (Bocak et al., 2014). From Japan and Taiwan, two Nosodendron species were Nosodendron Latreille, 1804: 146. ––– Háva, 2014: 14 [world

Ⓒ Japanese Society of Systematic Entomology 36 Yoshitomi, H., T. Kishimoto and C.-F. Lee

Fig. 1. Dorsal habitus of Nosodendron spp. A, N. (N.) asiaticum; B, N. (D.) coenosum; C, N. (D.) ogasawaraense (holotype); D, N. (D.) taiwanense (holotype). Scales = 1.0 mm.

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Fig. 2. SEM photographs of Nosodendron (N.) asiaticum. A–C, Elytra; D–F, pronotum.

catalogue]. Other synonyms see Zahradník & Háva convex dorsally, flattened ventrally, slightly to strongly (2014). Type species: Shaeridium fasciculare Olivier, shining. Coloration of body generally black. Dorsal surface 1790. covered with short erect setae or glabrous, with setal patches in the subgenus Nosodendron. Additional diagnosis. Close description of the family Gena with shallow concavity covered with short setae (Fig. Nosodendridae and the genus Nosodendron see Ivie (2002) 4A, C). Labrum (Fig. 8A, B) free, transverse, front margin and Leschen & Beutel (2010) for the adults, and Beutel (1996), straight (Fig. 8A) or shallowly concave (Fig. 8B). Mandibles Ge et al. (2007), Hayes & Chu (1946), Lawrence (1991) and (Fig. 8G, H) almost symmetrical, subtriangular, with pointed Leschen & Beutel (2010) for the larvae. apical tooth and small and obtuse subapical tooth. Maxillae Adults. Body hemispherical to oblong-oval, strongly (Fig. 8C, D) with 3-segmented short maxillary palpi. Labium

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Fig. 3. SEM photographs of Nosodendron (Dendrodipnis) taiwanense sp. nov. (A, B, D, E) and N. (D.) ogasawaraense sp. nov. (C, F). A, Pronotum and its punctures (D); B, C, elytra and its punctures (E, F).

(Fig. 8E, F) deeply excised at front margin, with 2-segmented Elytra punctate sparsely to closely, lacking any maculations labial palpi. Mentum (Fig. 6) large, trapezoidal, subparallel- and striae, with plectrum (Fig. 4D–F) in humeral part of inner sided (Fig. 6B–D) or contracted in apical 1/3 (Fig. 6A), surface. Hind wings well deveroped; venation as shown in covered with large and shallow punctures (Fig. 6E, F), with Fig. 7B, C; flight activity present (see biological notes of each special concavities in some species (e.g., Yoshitomi, 2013b). species) or absent (Leschen & Beutel, 2010). Mesoventrite Antennae (Figs. 4B, 7A, C) relatively short, 11-segmented; (Fig. 5F) with deep median impression for reception of segments IX–XI forming club, closely covered with prosternal process. Legs (Fig. 5E) short and flat. Fore legs recumbent short setae. Scutellar shield small, triangular. received to excavation of hypomeron; mid legs received to

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Fig. 4. SEM photographs of Nosodendron (Dendrodipnis) ogasawaraense sp. nov. A, Head in ventral view; B, antenna; C, genal concavity; D, humeral part of right elytron in ventral aspect; E, inner surface of elytron and its close up (F). depression in anterior part of epipleuron; hind legs received to Tergites IX–X slightly sclerotized, small. Sternite IX long. depressions in abdominal ventrites I–II. Abdominal ventrites Aedeagus trilobite type, rather flat dorsally and ventrally; (Fig. 5A–D) microcuticulate, closely covered with deep apical parts of parameres having 1–4 short setae (“apical punctures bearing short setae; anterior margin of each segment setae” in Yoshitomi, 2013b). arranged deep notches (Fig. 5B, D). Female genitalia. Sternite VIII well sclerotized, with long Sexual dimorphism in external feature indistinct. and slender speculum ventral. Tergite VIII well sclerotized, Male genitalia. Tergite VIII well sclerotized, semicircular. semicircular. Ovipositor with short or long paraproct. Sternite VIII moderately sclerotized, with V-shaped strut. Larvae. Body (Fig. 9) elongate, flattened in ventral part,

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Fig. 5. SEM photographs of Nosodendron (Dendrodipnis) ogasawaraense (A, B, E, F) and N. (D.) taiwanense sp. nov. (C, D). A, C, Abdominal ventrites and their close up (B, D); E, fore leg in dorsal aspect:; F, meso- and metaventrites. well sclerotized dorsally and slightly sclerotized ventrally, dorsal portion; spiracles of abdominal segment VIII moved covered with granules forming specific and irregular patterns to apex. Abdominal segment VIII (Fig. 10B, D, F) triangular, (Figs. 9, 10); most of granule bearing short setae (Fig. 11D); tapering posteriorly, with one or two pairs of projections in granules situated near spiracles bearing pectinate short setae dorsal or lateral portion. (Fig. 11E). Head (Fig. 10A, C, E) prognathous. Spiracles on Remarks. Six larvae (N. fasciculare, N. californicum, N. thorax situated in lateral portion; spiracles on abdomen I–VII unicolor, N. zealandicum, N. asiaticum, N. angelum) and one forming short tubular processes (Fig. 11C), situated in lateral/ possible larva (N. ovatum) have been reported (Hayes & Chu,

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Fig. 6. SEM photographs of mentum of Nosodendron spp. (A–D) and its close up (E, F). A, Nosodendron (N.) asiaticum; B, E, N. (D.) coenosum; D, F, N. (D.) ogasawaraense; C, N. (D.) taiwanense.

1946; Crowson, 1959; Hayashi, 1954; Costa et al., 1986; distinct setal patches (Fig. 3); mentum subparallel- Beutel, 1996), but the taxonomic status of the possible larva sided (Fig. 6B–D), closely/sparsely covered with large of N. ovatum is uncertain because of the presence of strange punctures, with sulcus or impression in some species; urogomphi-like processes in tergite VII (Crowson, 1959). phallobase longer than parameres and penis; ovipositor Costa et al. (1986) and Sharp (1902) reported the larvae short/long. Larvae (N. unicolor, N. zealandicum, N. probably emit sound by using the galea; however, we could angelum, N. coenosum, N. ogasawaraense): abdomi- not confirm the stridulating sound in both field and rearing nal segment VIII with two pairs of projections in lateral conditions. Oehme (1949) described a pair of gland-like portions; spiracles on abdominal segments II–VII situated organs present on the dorsal part of the mesothorax and in dorso-lateral portion, visible from dorsal aspect abdominal segment I of the adult of Nosodendron fasciculare (Fig. 9B, C)...... subgenus Dendrodipnis (also see Klausnitzer, 1971). We dissected and examined two Japanese species (N. asiaticum, N. coenosum), but could not Subgenus Nosodendron Latreille, 1804 find this organ. Nosodendron Latreille, 1804: 146. Type species: Shaeridium Key to subgenera fasciculare Olivier, 1790. Nosodendron (Nosodendron): Champion, 1923: 578. 1. Adults: antennomere III almost equal length in antenno- Nosodendron fasciculare species group: Háva, 2014: 14 [world meres I–II combined (Fig. 7A); pronotum and elytra with catalogue]. distinct setal patches (Fig. 2; sometimes fallen out in specimens condition); mentum immediately contracted in Remarks. Four species distributed in the Holarctic Region apical 1/3 (Fig. 6A), closely covered with large punctures; are recognized in the subgenus. phallobase shorter than parameres and penis; ovipositor distinctly short. Larvae (N. fasciculare, N. californicum, Nosodendron (Nosodendron) asiaticum Lewis, 1889 N. asiaticum): abdominal segment VIII with a pair of [Japanese name: Kemon-himetogemushi] projections in dorsal portion; spiracles on abdominal (Figs. 1A, 2, 6A, E, 7A, B, 8B, C, F, 9A, 10A, B, 11F, 12, 13, segments II–VII situated in lateral portion, invisible 20A, B) from dorsal aspect (Fig. 9A)…….subgenus Nosodendron -. Adults: antennomere III clearly longer than antennomeres Nosodendron asiaticum Lewis, 1889: 229. Hayashi, 1954: 23 I–II combined (Fig. 7C); pronotum and elytra without [description of larva].

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Fig. 7. Antennae (A, C) and hind wings (B, D) of Nosodendron spp. A, B, Nosodendron (N.) asiaticum; C, D, N. (D.) coenosum.

Specimens examined. Adults. 1 male (coating with gold Larvae (all EUMJ, preserved in 70% ethanol). 1 ex., and observed in SEM) & 12 exs. (EUMJ), “(HOKKAIDO) “Assabu, Hokkaido, 24. V. 2013, A. Nomura leg. (from Ulmus Kawayu 21~22. VII. 1970 S. Kinoshita”; 1 ex. (KUMJ), davidiana var. japonica)”; 1 ex., ditto, but “25. V. 2013”; 2 “Hokkaido Y. Nishio VI. 15. 1948 SAPPORO”; 1 ex. (KUMJ), exs., ditto, but “8. VI. 2013”; 1 ex., ditto, but “24. VI. 2013”. “Makoi, Shari-chô, Hokkaidô, Japan 13. VI. 2000 Y. Kida Description. Adults. Male. Body (Fig. 1A) oval, leg.”; 1 ex. (KUMJ), “Kamimuri, Maruseppu-chô, Hokkaidô, hemispherical, strongly convex dorsally, shiny. Coloration Japan 21. V. 2000 Maruyama M. leg.”; 5 exs. (EUMJ), “Mt. of body black; antennomeres I–VIII, mouth parts and legs NIYAMA Nanae Hokkaido 25 VII 1982 Y. Kusui leg.”; 4exs. blackish brown; antennomeres IX–XI closely covered with (EUMJ), “Assabu, Hokkaido, 12. V. 2013, A. Nomura leg. yellowish-brown short setae. (from Ulmus davidiana var. japonica)”; 10 exs., same locality, Head closely covered with coarse punctures, gently collector and host plant, but “24. VI. 2013”; 2 exs. (EUMJ), arcuate in front margin. Pronotum (Fig. 2D–F) punctate as same locality and collector, but “8. V. 2013 (from Betula in head; scaly setal patches present on mesal parts, indistinct platyphylla)”; 2 exs. (EUMJ), same locality, collector and host in some specimens; PW/PL 2.09–2.56 (2.33). Scutellar plant, but “10. V. 2013”; 1 ex. (KUMJ), “[JAPAN: FUKUI] shield triangular, microcticulate. Elytra (Fig. 2A–C) oval, Koike—Karikomi-no-ike 31. V. 1977 H. Sasaji”, “J017578”; well convex dorsally; punctures large and deep, uniformly 1ex. (EUMJ), “Nishidake-Path Togakushi Nagano-Pref. close and distinct, bearing short setae in anterior edge of 6 VII 2008 S. TSUYUKI leg.”; 1 ex. (EUMJ), “Hirakura each puncture, distance between punctures ca 0.5–1.0X its (600–700 m) Misugi Vill. Mie Pref., JAPAN 27. VII. 1985 diameter; scaly setal patches present, semicircular, about 0.2 Katsumi AKITA leg.”; 1 male, 1 female & 10 exs. (EUMJ), mm in size of individual patch, uniformly ordered by 7–8 X “Mt. Sara IYO 2. VII. 1953 T. Edashige”. 1 ex., “Mt. KOTSU 5 in each elytron, distance between patches ca 1.0–2.0X its TOKUSHIMA, P 29-IV 1966 Coll. M. SAKAI”; 1 ex. (EUMJ), diameter; EL/EW 1.12–1.26 (1.21); EL/PL 3.02–3.69 (3.44); “Mt. Kamegamori Ehime: JAPAN 16. VII. 1977 A. Oda leg.”; EW/PW 1.19–1.28 (1.22); TL/EW 1.77–1.95 (1.86). 4 exs. (EUMJ), “(SHIKOKU) Omogokei Omogo-mura 9. VI. Male. Tergite VIII (Fig. 12A) well sclerotized, 1978 A. Oda leg.”; 7 exs. (EUMJ), “[Ehime: Japan] Jyojyu semicircular, sparsely covered with short setae; sternite VIII Mt. Ishizuchi 29. VI. 2014 K. Sonaka leg.”; 1 ex. (EUMJ), (Fig. 12C) moderately sclerotized, short V-shaped, bearing “MIYAZAKI: JAPAN Gokase-cho 1100m 28. III. 2004 M. minute setae; tergite IX (Fig. 12B) lightly sclerotized, Matsumoto leg.”. subtriangular, bearing minute setae; sternite IX (Fig. 12D)

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Fig. 8. The adult mouth parts of Nosodendron spp. A, B, Labrum in dorsal aspects; C, D, maxillae in dorsal aspects; E, F, labium; G, H, mandibles in dorsal (G) and ventral (H) aspects. A, D, E, G, H, Nosodendron (D.) coenosum; B, D, F, N. (N.) asiaticum. lightly sclerotized, short and wide, bearing short setae, with with relatively short and stout spiculum ventrale. Ovipositor lateral struts; tergite X (Fig. 12B) membranous, fan-shaped, (Fig. 12I) short; styli oblong, bearing very long setae at closely covered with minute setae. Aedeagus (Fig. 12E, F) apices; distal lobe of gonocoxite oblong, 1.1 times as long about 0.6 mm, 1.9 times as long as sternite VIII; phallobase as proximal lobe; paraproct very short; approximate ratio of short and wide, arcuate in basal margin; parameres short, 1.6 stylus, gonocoxite, and paraproct (n = 1) as 1.0 : 5.3 : 3.9. times as long as phallobase, rather pointed at apices, with Measurements of adults (n = 20). TL 4.40–5.85 (5.12) three pairs of short and stout setae in apico-dorsal surfaces, mm; HL 0.65–0.98 (0.82) mm; PW 2.00–2.55 (2.26) mm; PL sparsely punctate; penis 2.4 times as long as phallobase, 0.80–1.10 (0.97) mm; EL 2.95–3.80 (3.33) mm; EW 2.43–3.15 obtuse at apex, wide, with long and stout struts. (2.76) mm. Female. Tergite VIII (Fig. 12G) moderately sclerotized, Larvae. Body length about 7.0 mm in fully expanded transverse, closely covered with minute spines, sparsely with specimen. Dorsal surface sparsely covered with granules short setae; sternite VIII (Fig. 12H) moderately scleortized, in dorsal surface. Head (Fig. 10A) transverse, with straight transverse, with straight posterior margin, bearing long setae front margin of clypeus. Labrum semicircular, arcuate in along posterior margin, closely covered with minute setae, front margin, bearing simple short setae, with complicated

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Fig. 9. Dorsal habitus of Nosodendron spp., larvae. A, N. (N.) asiaticum; B, N. (D.) coenosum; C, N. (D.) ogasawaraense. epipharynx (Fig. 13E). Mandibles (Fig. 13A, B) with three Subgenus Dendrodipnis Wollaston, 1873 teeth in subapical retinaculum. Maxillae (Fig. 13D) with simply pointed lacinia; galea slender; palpi shorter than galea. Dendrodipnis Wollaston, 1873: 33. Type species: Dendro- Hypopharynx (Fig. 13F) longitudinal; bridge sclerite simply dipnis coenosus Wollaston, 1873 (by the original curved, indistinct in anterior arm of bridge; suspensorial designation). sclerite long. Labium (Fig. 13C) relatively short, with Nosodendron (Dendrodipnis): Sharp, 1902: 670; Champion, long palpi. Thorax sparsely covered with granules forming 1923: 578. some irregular lines (Fig. 10A), projecting postero-laterally Nosodendron coenosum species group: Háva, 2014: 15 [world in postero-lateral corners. Abdomen with three pairs of catalogue]. longitudinal granules lines, with lateral projections projecting Remarks. This subgenus has been treated as a junior postero-laterally; spiracles on abdominal segments II–VII (Fig. synonym of the nominotypical subgenus by some authors (e.g., 11F) situated in ventral part of lateral projection, invisible DallaTorre, 1911; Háva, 2014). In the present paper, we treat from dorsal aspect; abdominal segment VIII (Fig. 10B) it as an independent subgenus based on the adults and larval widest at basal 1/4, then tapering posteriorly, with a pair of characteristics shown in the previous key. Most of the species projections in dorsal portion. in the genus (= 71 species treated in Nosodendron coenosum Distribution. Japan: Hokkaido, Honshu, Shikoku, Kyushu; species group in Háva, 2014) should belong to this subgenus, Russia: Kuril Islands (after Háva, 2014). but its monophyly must be reviewed. Biological notes. Both the larvae and adults of some species were collected from tree sap (see Table 1). This species is distributed in the plains to the mountains in the Nosodendron (Dendrodipnis) coenosum northern part of Japan (e.g., Hokkaido), but are limited to (Wollaston, 1873) [Japanese name: Kuro-himetogemushi] the mountainous areas in the southern part (e.g., Honshu to (Figs. 1B, 6B, 7C, D, 8A, D, E, G, H, 9B, 10C, D, 11D, E, Kyushu). 14–16, 20C, D) Remarks. Lewis (1889) described this species based on Nantaizan (in Nikko), Ichibosayama (probably incorrect Dendrodipnis coenosus Wollaston, 1873: 34. spelling of “Ichibusayama” in Kyushu), and other mountains. Nosodendron coenosum: Lewis, 1889: 229. Nomura & Kamezawa, 2014: 21 [SEM observation].

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Fig. 10. Head (A, C, E) and abdominal segments IV–VIII of Nosodendron spp., larvae. A, B, N. (N.) asiaticum; C, D, N. (D.) coenosum; E, F, N. (D.) ogasawaraense.

Specimens examined. Adults. 4 exs. (EUMJ), “Rinshi- “Is. Miyake (Tokyo to 8-VIII-1956) H. Yamazaki”; 2 males no-mori Park Koyamadai Shinagawa-ku, Tokyo [Honshu, & 2 females, “[Shimane: St. 62] Daimanji-san, Okinoshima, JAPAN] N35°37'26'' E139°42'12'' 5. V. 2013, S. Nomura Japan, N36 15.409 E133 19.831, ca 618m, 10. X. 2014, leg.”; 2exs. (EUMJ), ditto but 29. ix. 2013; 1 male (TUAJ), H. Yoshitomi leg.”; 2 males, 1 female & 9exs (EUMJ, 1

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Fig. 11. Larvae of Nosodendron (Dendrodipnis) ogasawaraense (A–C), N. (D.) coenosum (D, E), and N. (N.) asiaticum (F). A, Thoracic surface; B, lateral part of abdominal sengemns I–III; C, spiracle on abdominal segment I; D, granules on abdominal segment II; E, granule situated near spiracle; F, lateral part of abdominal segments I–III in ventral aspect. Scales = 500 μm. male specimen observed by SEM), “[EHIME: JAPAN] Matsuyama City 5. V. 2002 Jirô Ogawa leg.”; 1 ex. (EUMJ), Ishitegawa Park Nakamura Matsuyama City 18. VI. 2007 S.-T. “EHIME : JAPAN Near Rakanana Nomura Town 18. VI. Hisamatsu leg.”; 6 exs. (EUMJ), “(SHIKOKU) Side of Ishite 1959 N. Hamada leg.”; 1 ex. (EUMJ), “Hiraodai 27. VII. 1979 Riv. Matsuyama City 27. IX. 1983 K. Fujita leg.”, “at sap of Y. Takakura”; 1 ex. (KUMJ), “[Kyushu] Ohata-ike (Hyuga) Aphananthe aspera”; 14 exs. (EUMJ), “[Ehime-pref.: Japan] 10. Ix. 1952 Nakao, Ogata, & Yamamoto”; 1 ex. (KUMJ), “Mt. Takusen-cho Matsuyama-shi 12. VII. 2002 T. Kurihara leg.”; Tatsuda Kumamoto City Kumamoto Pref. 24. IV. 1976 Isao 2 exs. (EUMJ), same locality but 14. VII. 2003, T. Kitano Ohtsuka”; 1 ex. (KUMJ), “Oigami Hitoyoshi City 22. II. 1957 leg.; 1 ex. (EUMJ), same locality but 20. V. 2006, T. Kitano Isao Ohtsuka”. leg.; 1 female (EUMJ), “EHIME: JAPAN Mt. Shiroyama Larva. 9exs. (EUMJ), “[Shimane: St. 62] Daimanji-san,

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Fig. 12. Male (A–F) and female (G–I) genitalia of Nosodendron (N.) asiaticum. A, G, Tergite VIII; B, tergite IX–X; C, H, sternite VIII; D, sternite IX; E, F, aedeagus in dorsal (E) and ventral (F) aspects; I, ovipositor.

Okinoshima, Japan, N36 15.409 E133 19.831, ca 618m, pronotum; EL/EW 1.16–1.30 (1.24); EL/PL 3.19–3.71 (3.46); 10. X. 2014, H. Yoshitomi leg.”; 1 ex. (EUMJ, preserved in EW/PW 1.15–1.26 (1.21); TL/EW 1.76–1.94 (1.86). permanent slide), “Ishitegawa, Matsuyama, -. IV. 2013 S.T. Male. Tergite VIII (Fig. 14A) well sclerotized, transverse, Hisamatsu leg.”. densely covered with short setae, bearing curved setae Description. Adults. Body (Fig. 1B) oblong-oval, in lateral portions; sternite VIII (Fig. 14C) moderately strongly convex dorsally, shiny. Coloration of body black; sclerotized, V-shaped, bearing minute setae; tergite IX (Fig. antennomeres IX–XI closely covered with yellowish-brown 14B) lightly sclerotized, U-shaped, sparsely punctate; sternite short setae. IX (Fig. 14D) lightly sclerotized, expanded apically, bearing Head closely covered with coarse punctures bearing short short setae and spines in apical portion, with lateral struts; setae, arcuate in front margin. Mentum (Fig. 6B, E) flat, tergite X (Fig. 14B) membranous, bearing short spines. covered with large punctures. Pronotum punctate as in head; Aedeagus (Fig. 14, E, F) about 0.9 mm, 1.5 times as long as PW/PL 2.15–2.50 (2.32). Scutellar shield triangular, sparsely sternite VIII; phallobase short, subparallel-sided in posterior covered with fine punctures. Elytra punctate as in head and half, arcuate in basal margin; parameres long, straight in

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Fig. 13. Larval mouth parts of Nosodendron (N.) asiaticum. A, B, Mandibles in dorsal (A) and ventral (B) aspects; C, labium; D, maxilla; E, epipharynx; F, hypopharynx. ab, Anterior branch of suspensoria; bg, bridge sclerite of hypopharynx; ss, suspensorial sclerite. lateral margins, 3.3 times as long as phallobase, truncate at Female. Tergite VIII (Fig. 14G) moderately sclerotized, apices, with two pairs of short setae in apico-dorsal surface, semicircular, closely bearing short curved setae; sternite VIII sparsely punctate; penis 3.2 times as long as phallobase, (Fig. 14H) moderately sclerotized, transversal oblong, bearing obtuse and thick at apex, with long struts. long setae along posterior margin, closely covered with short

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Fig. 14. Male (A–F) and female (G–I) genitalia of Nosodendron (D.) coenosum. A, G, Tergite VIII; B, tergite IX–X; C, H, sternite VIII; D, sternite IX; E, F, aedeagus in dorsal (E) and ventral (F) aspects; I, ovipositor. setae, with long and slender spiculum ventrale. Ovipositor transverse, with straight front margin of clypeus. Labrum (Fig. 14I) long; styli thick, bearing long and short setae in shallowly concave in front margin. Mandibles (Fig. 15A, B) apical portions; distal lobe of gonocoxite obtuse at apices, 1.2 with two teeth in subapical retinaculum. Maxillae (Fig. 15D) times as long as proximal lobe; approximate ratio of stylus, with tridentate lacinia; galea wide; palpi longer than galea. gonocoxite and paraproct (n = 1) as 1.0 : 5.3 : 6.2. Hypopharynx (Fig. 15E) relatively wide; bridge sclerite with Measurements of adults (n = 20). TL 5.45–7.00 (6.30) anterior arm; suspensorial sclerite short. Labium (Fig. 15C) mm; HL 0.80–1.10 (0.89) mm; PW 2.47–3.12 (2.81) mm; PL relatively short; palpomeres II nipple-like. Thorax (Fig. 10C) 1.05–1.35 (1.22) mm; EL 3.60–4.60 (4.20) mm; EW 3.10–3.70 slightly projecting postero-laterally in postero-lateral corners. (3.39) mm. Abdomen with short lateral projections projecting postero- Larvae. Body length about 10.0 mm in fully expanded laterally; spiracles on abdominal segments II–VII (Fig. 11B) specimen. Dorsal surface closely covered with granules except situated in dorso-lateral part of each tergite, visible from for glabrous areas forming irregular patterns. Head (Fig. 10C) dorsal aspect; abdominal segment VIII (Fig. 10D) straightly

May 30, 2015, JJSE 21 (1) 50 Yoshitomi, H., T. Kishimoto and C.-F. Lee

Fig. 15. Larval mouth parts of Nosodendron (D.) coenosum. A, B, Mandibles in dorsal (A) and ventral (B) aspects; C, labium; D, maxilla; E, hypopharynx. aa, Anterior arm of bridge sclerite; ab, anterior branch of suspensoria; bg, bridge sclerite of hypopharynx; pme, proximal median element of hypopharyngeal sclerite; ss, suspensorial sclerite.

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20 18 larvae mature larvae immature 16 adults 14 12 10 8 6 4 2 Fig. 16. Number of Nosodendron (Dendrodipnis) coenosum attacking to 0 5/16 6/5 7/10 8/10 9/30 10/21 11/6 12/6 1/17 2/21 3/19 4/7 4/10 5/13 sap of one tree of Aphananthe aspera at Ishitegawa, Matsuyama-shi (Fig. 20D) 2013 2014 in 2013 to 2014. tapering posteriorly, with two pairs of projections in lateral N. coenosum; mentum (Fig. 6D, F) with shallow longitudinal margin. concavity in basal 1/2; PW/PL 2.13–2.37 (2.22); elytra (Fig. Distribution. Japan: Hokkaido (after Sakai, 1985), 3C, F) covered with large punctures with short setae; EL/EW Honshu, Izu Isls. (Miyake-jima), Okinoshima, Shikoku, 1.11–1.24 (1.20); EL/PL 3.11–3.42 (3.22); EW/PW 1.18–1.24 Kyushu. (1.21); TL/EW 1.72–1.88 (1.83). Biological notes. Both the larvae and adults of some Male. Tergite VIII (Fig. 17A) well sclerotized, semicircular, species were collected from tree sap (see Table 1). Hibernation densely covered with short spines, bearing short curved setae in is occurred in the adult stage and possibly also the larval stage lateral portion; sternite VIII (Fig. 17C) moderately sclerotized, in the soil under the sap (6 Dec. to 19 Mar. in Fig. 16). The V-shaped, bearing minute setae; tergite IX (Fig. 17B) lightly length of one generation is uncertain, but it is probably more sclerotized, U-shaped, sparsely punctate; sternite IX (Fig. 17D) than one year. Because both larvae and adults can be observed lightly sclerotized, expanded apically, bearing short setae and through the year (Fig. 16), old adult individuals (elytral spines in apical portion, without lateral struts; tergite X (Fig. surface is worn down and its setae are almost removed) are 17B) bearing short setae. Aedeagus (Fig. 17E, F) about 0.7 also seen in most seasons, and several instar larvae can be mm, 1.6 times as long as sternite VIII; phallobase short, arcuate obtained in the same season (Yoshitomi, 2015). This species in basal margin; parameres long, straight in lateral margins, 3.0 is distributed mainly in the plains in the southern part of Japan times as long as phallobase, truncate at apices, with two pairs (e.g., Honshu to Kyushu). of short setae in apico-dorsal surface, closely punctate in apical Remarks. Háva (2005, 2014) recorded this species from portions; penis 2.6 times as long as phallobase, obtuse at apex, Taiwan, but these records indicate that the following species with short struts. was probably misidentified. Female. Tergite VIII (Fig. 17G) moderately sclerotized, This is the first record of this species from the Izu Isls. semicircular, with weakly straight in posterior margin, closely covered with minute spines, sparsely bearing short Nosodendron (Dendrodipnis) ogasawaraense sp. nov. curved setae; sternite VIII (Fig. 17H) moderately sclerotized, [Japanese name: Ogasawara-himetogemushi] transverse, bearing minute setae in apical portion, with long (Figs. 1C, 3C, F, 4, 5A, B, E, F, 6D, F, 9C, 10E, F, and slender spiculum ventrale. Ovipositor (Fig. 17I) long; 11A–C, 17, 18, 20E) styli oblong, bearing long and short setae in apical portions; distal lobe of gonocoxite obtuse at apex, 1.67 times as long as Type series. Holotype (EUMJ), male, “[OG12] Sekimon, proximal lobe; approximate ratio of stylus, gonocoxite, and Hahajima, Ogasawara Isls., Japan, N26 41.018 E142 paraproct (n = 1) as 1.0 : 6.0 : 8.13. 09.631 30. IX. 2012, H. Yoshitomi leg”. Paratypes (EUMJ, Measurements of adults (n = 11). TL 4.65–5.77 (5.22) NSMT): 3 males, 3 females & 6 exs., same data as for the mm; HL 0.60–0.85 (0.74) mm; PW 2.18–2.55 (2.36) mm; PL holotype; 1 ex., ditto but 29. IX. 2012; 1 female, “[OG6] 0.95–1.20 (1.06) mm; EL 3.00–3.85 (3.42) mm; EW 2.70–3.10 Koumoridani, Hahajima, Ogasawara Isls., Japan, N26 39.203 (2.86) mm. E142 09.295, 25, 28, 30. IX. 2012, H. Yoshitomi leg.”; 1 ex., Larvae. Externally (Fig. 9C) closely similar to N. “[OGASAWARA] Okiko Hahajima I 17. V. 1972 Y. Kusui coenosum; body length about 7.0 mm; abdominal segment leg.”. VIII (Fig. 10F) longer; maxillary and labial palpi shorter. Larval specimens examined. 4 exs. (EUMJ, preserved in Distribution. Japan: Ogasawara Isls. (Haha-jima). 70% ethanol), “[OG12] Sekimon, Hahajima, Ogasawara Isls., Biological notes. The adults and larvae were collected Japan, N26 41.018 E142 09.631 30. IX. 2012, H. Yoshitomi from the sap of Hibiscus glaber (Matsum. ex Hatt.) Matsum. leg”. ex Nakai in Sekimon. One specimen was collected using a Description. Adults. Externally (Fig. 1C) closely similar to light trap setting in Koumoridani. For the habitat photos of the

May 30, 2015, JJSE 21 (1) 52 Yoshitomi, H., T. Kishimoto and C.-F. Lee

Fig. 17. Male (A–F) and female (G–I) genitalia of Nosodendron (D.) ogasawaraense. A, G, Tergite VIII; B, tergite IX–X; C, H, sternite VIII; D, sternite IX; E, F, aedeagus in dorsal (E) and ventral (F) aspects; I, ovipositor.

Ogasawara Isls., see the website https://sites.google.com/site/ but 6~7. V. 2006; 3 exs. (TARI), ditto but 8. XI. 2005; 1 ex. waterbeetlesofjapan/home/field-investigation/. (TARI), ditto but 23. IV. 2006; 4 exs. (TARI), ditto but 27. Remarks. Nosodendron (D.) carolinense Yoshitomi, 2013 VIII. 2005; 1 ex. (EUMJ), “[Taiwan] Wulu Alt. ca. 700m is known from the Caroline Isls., Micronesia, but differs Haiduan Township Taitung Country 17–18. VI. 2011 J. Yamasako leg.”; 1 ex. (Yoshitomi, 2013a). (BPBM), “TAIWAN (Formosa) Puli (Hori) July, 1954 Native collector”; 14 exs. (TARI), “Taiwan: Taichung Pahsienshan Nosodendron (Dendrodipnis) taiwanense sp. nov. 18. III. 2014. leg. C.-F. Lee”; 7 exs. (TARI), “Taiwan: Taitung [Japanese name: Taiwan-himetogemushi] Chihpen 13.IV.2014. leg. T.- (Figs. 1D, 3A, B, D, E, 5C, D, 6C, F, 19, 20F) T. Chung”; 1 ex. (EUMJ), “[TAIWAN] Siangyang, Alt. ca. 2,100–2,300m, Haiduan Township, Taitung Country 7–9. VI. 2013, J. Yamasako leg.”; 1 299; Háva, 2014: 15. ex. (EUMJ), “TAIWAN Siangyang, Alt. ca 2,100–2,300m, Haiduan Township, Taitung Country (Light) 8. VI. 2013, K. Sonaka leg.”. Heshe (Tongfu) 7~9. IV. Additional specimens (non-type series). 5 Males & 1 2006, leg. C.F. Lee”. Paratypes, 1 Male, 1 female & 2 exs. female (EUMJ), “Omoto, Ishigaki-jima, Okinawa, 24-28. (TARI), same data as for the holotype; 4 exs. (TARI), ditto X. 2014, R. Okano leg., by banana trap”; 2 exs. (TARI),

May 30, 2015, JJSE 21 (1) Nosodendridae of Japan and Taiwan 53

Fig. 18. Larval mouth parts of Nosodendron (D.) ogasawaraense. A, B, Mandibles in dorsal (A) and ventral (B) aspects; C, labium; D, maxilla; E, epipharynx; F, hypopharynx.

May 30, 2015, JJSE 21 (1) 54 Yoshitomi, H., T. Kishimoto and C.-F. Lee

Fig. 19. Male (A–F) and female (G–I) genitalia of Nosodendron (D.) taiwanense. A, G, Tergite VIII; B, tergite IX–X; C, H, sternite VIII; D, sternite IX; E, F, aedeagus in dorsal (E) and ventral (F) aspects; I, ovipositor.

“TAIWAN: Taitung Lanyu island 7.IV.2015, leg. Y.-T. Wang”. sclerotized, slightly expanded apically, bearing short setae Description. Adults. Externally (Fig. 1D) closely similar in apical portion, without lateral struts; tergite X (Fig. 19B) to preceding two species; mentum (Fig. 6C) flat, covered with membranous, bearing short spines. Aedeagus (Fig. 19E, F) large punctures; pronotum (Fig. 3A, D) shallowly punctate; about 0.9 mm, 2.1 times as long as sternite VIII; phallobase PW/PL 2.21–2.50 (2.34); elytra (Fig. 3B, E) covered with short, tapering basally in lateral margins, shallowly concave shallow punctures with short setae; EL/EW 1.20–1.30 (1.25); in basal margin; parameres very long, gently tapering apically, EL/PL 3.21–3.77 (3.52); EW/PW 1.16–1.25 (1.21); TL/EW 4.17 times as long as phallobase, rounded at apices, with two 1.80–1.98 (1.88). pairs of short setae in apico-dorsal surfaces, closely punctate; Male. Tergite VIII (Fig. 19A) well sclerotized, semi- penis 3.6 times as long as phallobase, bearing short spines in circular, densely covered with short setae, bearing curved dorso-apical portion, with long and straight struts. setae in lateral portions; sternite VIII (Fig. 19C) moderately Female. Tergite VIII (Fig. 19G) moderately sclerotized, sclerotized, V-shaped, bearing minute setae; tergite IX (Fig. semicircular, closely covered with minute spines and short 19B) slightly sclerotized, U-shaped, projecting in lateral setae; sternite VIII (Fig. 19H) moderately sclerotized, portions, bearing minute setae; sternite IX (Fig. 19D) slightly transverse, bearing minute setae in apical portion, with long

May 30, 2015, JJSE 21 (1) Nosodendridae of Japan and Taiwan 55

Fig. 20. Adult (A) and larva (B) of Nosodendron (N.) asiaticum, at Assabu, Hokkaido, photo by A. Nomura; C, D, larvae and adults (C) and its habitat (D) of N. (D.) coenosum, at Matsuyama, Ehime Pref., photo by HY; E, adults of N. (D.) ogasawaraense, at Sekimon, Ogasawara, photo by HY; F, adult of N. (D.) taiwanense, at Taiwan, photo by CL. spiculum ventrale. Ovipositor (Fig. 19I) long; styli oblong, Distribution. Taiwan, Lanyu Island, Japan: Ishigaki-jima. bearing long and short setae in apical portions; distal lobe Biological notes. The type series were collected from the of gonocoxite rather pointed at apices, 1.55 times as long as sap of Trema orientalis and Zelkova serrata, and Ishigaki proximal lobe; approximate ratio of stylus, gonocoxite, and specimens were collected using a banana trap. paraproct (n = 1) as 1.0 : 3.1 : 6.5. Remarks. The specimens collected from Ishigaki-jima are Measurements of adults (n = 20). TL 5.47–7.05 (6.28) slightly different from the Taiwanese specimens in the shapes mm; HL 0.80–1.15 (0.94) mm; PW 2.40–3.10 (2.76) mm; PL of the apices of the parameres and the basal margin of the 1.00–1.40 (1.18) mm; EL 3.62–4.68 (4.16) mm; EW 2.80–3.68 phallobase is very shallowly concaved, but we treat them as (3.33) mm. the same species in this paper. Larvae unknown. Nosodendron coenosum was recorded from Taiwan

May 30, 2015, JJSE 21 (1) 56 Yoshitomi, H., T. Kishimoto and C.-F. Lee

Table 1. The associated plants of Nosodendron spp. Host, related plants subgenus Species Reference family species Nosodendron californicus Pinaceae Abies concolor Reichardt (1976), Ives (2002) Abies grandis Reichardt (1976), Ives (2002) Pseudotsuga menziesii Ives (2002) Fagaceae Quercus kelloggii Reichardt (1976) asiaticum Ulmaceae Ulmus davidiana var. japonica Hayashi (1954), Sakai (1985) Salicaceae Populus tremula var. sieboldii present study Hippocastanaceae Aesculus turbinata present study Betulaceae Betula platyphylla present study Alnus japonica Hayashi (1954) Pinaceae Abies firma present study Dendrodipnis unicolor Ulmaceae Ulmus americana Reichardt (1976) Ulmus sp. Ives (2002) Aceraceae Acer saccharum Reichardt (1976) Fagaceae Quercus alba Reichardt (1976) Salicaceae Populus sp. Reichardt (1976), Ives (2002) Salix sp. Reichardt (1976), Ives (2002) Betulaceae Alnus sp. Ives (2002) angelum Lauraceae Nectandra sp. Reichardt (1976) Casuarinaceae Casuarina sp. Costa et al. (1986) testudinum Musaceae banana stumps Reichardt (1976) coenosum Ulmaceae Aphananthe aspera Sakai (1985), present study Cannabaceae Celtis sinensis Sakai (1985), present study Lauraceae Cinnamomum camphora Otsuka (2003) Pinaceae Abies firma Saito & Suzuki (2013) Salicaceae Idesia polycarpa Nomura & Kamezawa (2014) hispidum Fabaceae Acacia sp. Yoshitomi (2013) nomurai Fabaceae Acacia sp. Yoshitomi (2013) taiwana Ulmaceae Trema orientalis present study Zelkova serrata present study Musaceae banana trap present study ogasawaraense Malvaceae Hibiscus glaber present study zealandicum Podocarpaceae Dacrydium cupressinum Crowson (1959) by Háva (2005, 2014) and Steinhammer (2007). It was ...... ….. N. (D.) ogasawaraense, sp. nov. misidentified and actually belongs to the present new species since both species are very similar in general morphology and Larvae* all available material collected from Taiwan were found to be 1. Abdominal segment VIII with two pairs of projections this new species. in lateral portions. Spiracles on abdominal segments II– VII situated in dorso-lateral portion, visible from dorsal Key to species of the genus Nosodendron aspect…..subgenus Dendrodipnis…...... ….2 in Japan and Taiwan -. Abdominal segment VIII with a pair of projections in Adults dorsal portion. Spiracles on abdominal segments II– 1. Body oval. Mentum immediately contracted in apical 1/3. VII situated in lateral portion, invisible from dorsal Elytra with paches of scaly erected setae. TL about 4.40– aspect…….subgenus Nosodendron……..N. (N.) asiaticum 5.85 mm. …...... …….N. (N.) asiaticum 2. Abdominal segment VIII as long as that of width…… -. Body oblong. Mentum subparallel-sided. Elytra without ...... ….N. (D.) coenosum paches of setae. TL about 4.65–7.05 mm. ………..2 -. Abdominal segment VIII longer than that of wid 2. Parameres truncate at apices; phallobase arcuate in basal th……...... …N. (D.) ogasawaraense, sp. nov. margin; distributed in Japan……...... ….3 *: The larva of N. (D.) taiwanense unknown. -. Parameres rounded at apices; phallobase shallow- ly concave in basal margin; distributed in Taiwan and Discussion Ishigaki-jima...... ,...... N. (D.) taiwanense, sp. nov. Feeding habit 3. Mesal part of mentum lacking concavity; interior Both the larvae and adults of Nosodendron are attracted corners of parameres right-angled; penis long, 3.2 to tree sap and dwell in slime fluxes. Table 1 shows the host times as long as phallobase; distributed in Hokkaido plant relationship based on the reported records. Up to the to Kyushu……...... ……N. (D.) coenosum present, thirteen families of trees have been recorded as the -. Mesal part of mentum with shallow longitudinal concavity host plants. in basal part; penis short, 2.6 times as long as phallobase; The feeding habitat is uncertain. Hayashi (1954), Sokoloff distributed in Ogasawara Isls. …………......

May 30, 2015, JJSE 21 (1) Nosodendridae of Japan and Taiwan 57

(1959), and Costa et al. (1986) wrote that the larvae of Champion, G. C., 1923. A revision of the eastern species of Nosodendron are carnivorous, and feed mainly on dipteran Nosodendridae (Coleoptera). Annals and Magazine of Natural larvae and also probably other small creatures (e.g., Nematodes, History, 12: 578–591. Costa, C., S. A. Casari-Chen & É. P. Teixeira, 1986. Larvae mites, Collembola, Corrodentia, etc.) living in and around tree of Neotropical Coleoptera XVI. Nosodendridae. Revista sap. On the other hand, Lawrence (1989), Kovac et al. (2007), Brasileira de Entomologia, 30: 291–297. and Leschen & Beutel (2010) suggested that their main source Crowson, R. A., 1959. Studies on the Dermestoidea (Coleoptera), of food is not such small creatures, but fungal hyphae and with special reference to the New Zealand fauna. Transactions conidia. Certainly, the pointed mandibles in both the larvae and of the Royal Entomological Society of London, 111: 81–94. adults seem to indicate they are predators, but judging from Dalla Torre, K. W., 1911. Pars 33: Nosodendridae. In: Junk, W. its host plant specificity and complicated hypopharynx and & Schenkling, S. (eds): Coleopterorum Catalogus, W. Junk, epipharynx, the nosodendrid is probably a filter-feeder of Berlin, 96 pp. Ge, S-Q., R. G. Beutel and X-K. Yang, 2007. Thoracic yeast or fungus, and its carnivorous habit is accidental. morphology of adults of Derodontidae and Nosodendridae and its phylogenetic implications (Coleoptera). Systematic Respiratory habit Entomology, 32: 635–667. Kovac et al. (2007) reported that Amphicrossus japonicus Háva, J., 2000. Distributional notes on some Oriental Nosodendr- Reitter, 1873 (Coleoptera, Nitidulidae) inhabits tree sap, and idae (Coleoptera), with descriptions of five new species. Acta is adapted for aquatic habitats, i.e., the larvae have strongly Musei Moraviae, Scientiae biologicae (Brno), 85: 57–65. prolonged specular tubes and adults wear an air sheath at Háva, J., 2005. World catalogue of the Nosodendridae (Coleoptera), with new faunistic records and descriptions of three new their ventral surface using their antennae. In the family species. Entomological Problems, 35: 75–83. Nosodendridae, both the larvae and adults inhabit similar Háva, J., 2014. Updated world catalogue of the Nosodendridae microhabitats of A. japonicus, but the method of respiration is (Coleoptera: Derodontoidea). Heteropterus Revista de different. The larvae of Nosodendridae have well-developed Entomología, 14: 13–24. spiracles on the dorsal (subgenus Dendrodipnis) and ventral Hayashi, N., 1954. Notes on Nosodendron asiaticum Lewis (Studies (subgenus Nosodendron) surface, and project their spiracles on Coleopterous larvae II). New Entomologists, 3 (4): 22–25. from the surface of sap and slime fluxes. In addition, granules (In Japanese, with English title.) bearing pectinate short setae are scattered near the spiracles Hayes, W. M. P. & Chu, H. F., 1946. The larvae of the genus Nosodendron Latr. Annals of the Entomological Society of (Fig. 11E), and they probably play the part of a plastron. America, 39: 69–79. On the other hand, the adults frequently open up narrowly Hunt, T., J. Bergsten, Z. Levkanicova, A. Papadopoulou, O. St. between the elytra and abdomen, and take air from its opening John, R. Wild, P. M. Hammond, D. Ahrens, M. Balke, M. S. to under part of the elytra. This behavior occurs on the surface Caterino, et al., 2007. A comprehensive phylogeny of beetles of slime fluxes. reveals the evolutionary origins of a superradiation. Science PNU, 1913–1916. Acknowledgements Ivie, M. A., 2002. 67. Nosodendridae Erichson, 1846, pp. 224–227. In.: Arnett, R. H., Thomas, M. C., Skelley P. E. & Frank J. Field research and collection in Hahajima was done H.: American Beetles. Volume 2. Polyphaga: Scarabaeoidea with the permission of the Japan Environmental Agency, through Curculionidea. 861 pp., CRC Press, Washington. Ogasawara Board of Education, Tokyo Metropolis and the Jäch, M. A., 1998. Annotated check list of aquatic and riparian/ littoral beetle families of the world (Coleoptera), pp. 25–42. - Agency for Cultural Affairs. We thank Dr. Shuhei Nomura In Jäch, M.A. & Ji, L. (eds.): Water Beetles of China, Vol. II. and Mr. Hiroshi Kamezawa (both NSMT), Drs. Masahiro - Wien: Zoologisch-Botanische Gesellschaft in Österreich and Sakai and Sadatomo Hisamatsu, and Mr. Ryosuke Okano and Wiener Coleopterologenverein, 371 pp. Mr. Kotaro Sonaka (all EUMJ), Mr. Shepherd Myers (BPBM), Kiselyova, T. and J. V. McHugh, 2006. A phylogenetic study Dr. D. K. Young, Mr. Akihide Nomura (Assabu-cho), Mr. of Dermestidae (Coleoptera) based on larval morphology. Katsumi Akita (Tsu-shi) for supplying the materials used in Systematic Entomology, 31: 469–507. this paper, Dr. Rafal Ruta (University of Wrocław) and Dr. Jiri Klausnitzer, B., 1971. Zur Biologie einheimischer Kaferfamilien. 6. Háva for giving useful suggestions, and Mr. Dennis Murphy Nosodendridae. Entomologische Berichte, Berlin, 15: 71–73 Kovac, D., Jelínek, J., Hashim, R., and D. Wiwatwitaya, 2007. (The United Graduate School of Agricultural Sciences, Transition from bamboo sap to water:Aquatic habits in the Ehime University) for his critical reading of the draft. Part of Amphicrossus japonicus (Coleoptera: Cucujoidea: this study is supported by the KAKENHI (24510333; head Nitidulidae). European Journal of Entomology, 104: 635–638. investigator: H. Kojima). Latreille, P. A., 1804. Genera crustaceorum et insectorum. A. Koenig, Paris, 280 pp. References Lawrence, J. F., 1989. Mycophagy in the Coleoptera: feeding strategies and morphological adaptations. In Wilding, N., Beutel, R. 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