A New Species of the Genus Macrorhyncolus Wollaston, 1873 (Coleoptera: Curculionidae: Cossoninae) from China

Zootaxa 4365 (5): 547–558 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4365.5.2 http://zoobank.org/urn:lsid:zoobank.org:pub:0A605C3C-A725-41CD-AB9A-F5255D2B6044

A new species of the genus Macrorhyncolus Wollaston, 1873 (Coleoptera: Curculionidae: Cossoninae) from China

YOUSSEF M. OMAR1,2, RUNZHI ZHANG1, 3,5 & STEVEN R. DAVIS4 1Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, No. 1 Beichen West Road, Chaoyang District, Beijing 100101, CHINA. 2Plant Protection Department, Faculty of Agriculture, Assiut University, Assiut, EGYPT 3State Key Laboratory of IMPIR of China, 25 Beisihuan Xilu, Beijing 100080, CHINA 4Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th St., New York, NY 10024-5192, USA 5Corresponding author. E-mail: [email protected]

Abstract

Following recent taxonomic study, Macrorhyncolus spinosus sp. nov. is described from China. Earlier, Macrorhyncolus included four species, M. crassiusculus, M. crassitarsis, M. littoralis and M. ventilaginis. A key is presented to all known species of Macrorhyncolus in the Palaearctic Region and China. Illustrations of diagnostic characters of the new species are provided. Macrorhyncolus spinosus is winged, has larger eyes, the rostrum is longer than wide and has a median furrow in the basal half, extending to the vertex, and striae 9 and 10 remain separate from base to apex.

Key words: Macrorhyncolus spinosus, new species, Rhyncolina, Rhyncolini, Cossoninae, taxonomy, weevils

Introduction

Following the classification of Alonso-Zarazaga & Lyal (1999), Macrorhyncolus Wollaston, 1873 belongs to the tribe Rhyncolini of the subfamily Cossoninae. The genus was erected by Wollaston (1873a) for M. crassiusculus Wollaston, 1873 from Japan. Wollaston (1873b) subsequently added M. crassitarsis Wollaston, 1873 from Ceylon (Sri Lanka). Broun (1880) described Eutornus littoralis from Tairua, New Zealand, but in his later “Revision of the New Zealand Cossonidae” he stated that a possible transfer of E. littoralis Broun, 1880, E. cylindricus Broun, 1893 and E. parvulus Broun, 1893 to different genera might be considered justifiable as their eyes are different from those of other Eutornus species (Broun 1909). Leech (1955) treated Eutornus littoralis as Macrancylus LeConte & Horn[?], 1876 and synonymized M. franciscanus Van Dyke, 1953 with M. littoralis. Marshall in Leech (1955) synonymized E. cylindricus and E. parvulus with M. littoralis. Kuschel (1969) transferred M. littoralis to Macrorhyncolus. Additional described species of Macrorhyncolus are M. ventilaginis Marshall, 1938 from Nakronda and Dehra Dun in India and M. sulcirostris Voss, 1957 from Japan, the latter of which was moved to the genus Sphaerocorynes Wollaston, 1873a (Hlaváč & Maughan 2013: 220). Macrorhyncolus protractus Horn, 1873 was listed in the WTaxa Database of Curculionoidea (http://wtaxa.csic.es/ index.aspx) but Anderson (1952) had returned M. protractus to its original genus, Rhyncolus Germar, 1817. So, these two last species were excluded from Macrorhyncolus. According to the Zoological Record (1864–2013) electronic database (https://apps.webofknowledge.com/ ZOOREC_GeneralSearch_input.do?product=ZOOREC&SID=Q2D6MGTWMEDBblSNq6L&search_mode=Gen eralSearch), WTaxa (http://wtaxa.csic.es), Fauna of Europe (http://www.faunaeur.org), the Coleopterorum Catalogus of Csiki (1936), the World Catalogue of Alonso-Zarazaga & Lyal (1999) and the list of Chinese Insects (Hua 2002), no additional Macrorhyncolus species have been described so far. In this paper, a new species is described from Yunnan Province, southern China, bringing the number of known species to five.

Accepted by D. Clarke: 29 Nov. 2017; published: 21 Dec. 2017 547 Materials and methods

Specimens were studied under a Zeiss Semi SV11 stereomicroscope. Photographs were taken with a Micropublisher 5.0 RTV digital camera, model MP5.0-RTV-CLR-10A-color, attached to a Zeiss Discovery V12 stereomicroscope, and z-stacks were combined using the software CombineZ. SEM photographs were taken from Au-Pd-coated specimens using a LEO 1550 FE scanning electron microscope. Line illustrations were made in Adobe Illustrator CS3 and SEM photographs were enhanced in Adobe Photoshop CS3. Measurements were taken using an ocular micrometer and are defined using the following abbreviations: ACL—antennal club length; ACW—antennal club width; AFL—antennal funicle length; AL—antennal length; ASL—antennal scape length; BL—body length (including rostrum); EL—elytral length; EWB—elytral width at base; EWW—elytral width at widest part; PL—pronotal length; PW—pronotal width (widest part); RL—rostral length; RWA—rostral width at apex; RWB—rostral width at base. Hind wing terminology follows Zherikhin and Gratshev (1995). The new species is assigned to the genus Macrorhyncolus after comparing it with high resolution photos of the holotype of Macrorhyncolus crassiusculus Wollaston, 1873 (holotype tag: Macrorhyncolus crassiusculus W. (handwriting), Japan; G. Lewis. 1910—320; NHMUK010874327) and also due to the consensus with the diagnostic characters of the genus established by Wollaston (1873a, 1873b), Morimoto (1973), Zhang (1992) and Anderson (2002).

Results

Taxonomic treatment

Macrorhyncolus Wollaston, 1873

Macrorhyncolus Wollaston, 1873a: 33. Type species: Macrorhyncolus crassiusculus Wollaston, 1873, by monotypy. = Macrorhycolus; Leng & Mutchler, 1914: 479 (misspelling).

Diagnosis. Body slightly depressed, sides parallel; glabrous. Eyes large, prominent; distance between them slightly less than width of rostrum at base. Rostrum as long as or slightly longer than head, longer than wide, subparallel, slightly expanded towards apex in dorsal view. Antennae inserted close to base of rostrum such that dorso-lateral margins of rostrum are distinctly emarginated behind antennae; scapes with apex extended beyond hind margin of eyes; funicles with seven segments. Pronotum oblong, longer than wide. Elytra not marginate at base, angular. Tibiae with large uncus at outer-apical angle, produced at inner angle into distinct mucro; tarsi with tarsomere 3 often shallowly bilobed and broader than 2; tarsomere 3 of hind legs slightly wider than 2, entire or weakly emarginated (Wollaston 1873 a, 1873b; Morimoto 1973; Zhang 1992; Anderson 2002). Distribution. Japan; Sri Lanka; India; China; New Zealand; Australia (including Tasmania); United Kingdom (Ireland; Kent, South East England). According to Alonso-Zarazaga & Lyal (1999), the genus has been introduced to Chile, North America and South Africa.

Key to the species of Macrorhyncolus Wollaston in the Palaearctic Region and China

1- Rostrum as long as wide; elytral stria 10 merging with 9 in middle; venter with small, fine, sparse punctures . M. ventilaginis - Rostrum longer than wide; stria 9 and 10 remaining separate to apex; venter with coarse, dense punctures ...... 2 2- Head longer than wide; rostrum lacking median furrow (entirely punctured) ...... M. crassiusculus - Head wider than long; rostrum with median furrow born in basal half and extending to vertex ...... M. spinosus sp. n.

Macrorhyncolus spinosus Omar & Zhang sp. n. (Figs 1–35)

Holotype: Female: China. Yunnan Province: Shidong Village, Huangping Town, Heqing County, [26°01′N, 100°26′E]; 28 II 2004; collected by Qiao Li. Paratypes. 2M# and 2F#, same data as the holotype. Both holotype

548 · Zootaxa 4365 (5) © 2017 Magnolia Press OMAR ET AL. and paratype specimens are deposited in the Institute of Zoology (IOZ), Chinese Academy of Sciences (CAS), Beijing, China. Diagnosis. Body 4.50–4.70 mm; rostrum with apical 2/3 flat, basal 1/3 convex, parallel-sided, with median furrow born in basal half and continuing until vertex; scrobes covering basal 2/3 of rostrum until slight distance before anterior margin of eyes; points of antennal insertion at basal 1/3 of rostrum; upper margin of scrobes same level as upper margin of eyes; eyes black, dull, convex, subcircular, with convex facets; pronotum dorsally flat, glossy; with moderately dense, deep, large and circular punctures on whole surface; punctures separated by approximately 2–4 x puncture diameter; elytral interstriae strongly elevated, wider than striae except at base where they are as wide as or slightly wider than striae; interstriae 1 and 2 not dilated beyond declivity; striae with deep, circular punctures; punctures separated by approximately 0.5 x puncture diameter or less, sometimes cutting interstriae borders; protibiae with uncus (along outer margin distal to tarsal groove) longer than premucro; uncus longer and more stout in males. Description. Measurements (female): BL: 4.50–4.70 mm; EL: 2.57–2.75 mm; EWB: 1.05–1.18 mm; EWW: 1.05–1.18 mm; PL: 1.15–1.19 mm; PW: 0.97–1.04 mm; RL: 0.64–0.68 mm; RWA: 0.36–0.39 mm; RWB: 0.35– 0.39 mm; AL: 1.06–1.10 mm; ASL: 0.50–0.49 mm; AFL: 0.36–0.39 mm; ACL: 0.20–0.22 mm; ACW: 0.15–0.17 mm. Measurements (male): BL: 4.25–4.40 mm; EL: 2.40–2.54 mm; EWB: 0.91–1.06 mm; EWW: 0.91–1.06 mm; PL: 0.95–1.08 mm; PW: 0.83–0.92 mm; RL: 0.60–0.55 mm; RWA: 0.34–0.34 mm; RWB: 0.33–0.33 mm; AL: 0.88–1.03 mm; ASL: 0.36–0.44 mm; AFL: 0.33–0.39 mm; ACL: 0.19–0.20 mm; ACW: 0.15–0.15 mm.

FIGURES 1–3. Macrorhyncolus spinosus. 1, dorsal view; 2, lateral view (scale bar applies to figs 1–2); 3, antero-dorsal aspect of head and pronotum.

A NEW SPECIES OF MACRORHYNCOLUS WOLLASTON, 1873 Zootaxa 4365 (5) © 2017 Magnolia Press · 549 FIGURES 4–9. Macrorhyncolus spinosus, SEM photographs of legs. 4, fore-leg; 5, enlargement of pro-tibia; 6, enlargement of pro-tarsus; 7, middle leg; 8, enlargement of meso-tarsus; 9, hind leg.

550 · Zootaxa 4365 (5) © 2017 Magnolia Press OMAR ET AL. FIGURES 10–13. Macrorhyncolus spinosus, mouthparts. 10, left maxilla dorsal view; 11, labium ventral view; 12, left mandible dorsal view; 13, right mandible dorsal view.

A NEW SPECIES OF MACRORHYNCOLUS WOLLASTON, 1873 Zootaxa 4365 (5) © 2017 Magnolia Press · 551 FIGURES 14–15. Macrorhyncolus spinosus hindwing. C = Costa, Sc = Subcosta, R = radius, rcm = margin of radial cell, rf = radial fold, 1rs-2rs = radial sclerites, rm = radiomedia, msc = medial sclerotization, rsc = radial sclerotization, R3 = 3rd radial st nd nd vein, pst = postradial stripe, mst = medial stripe, Cu = Cubital, af = apical fold of Cu, 1A2 = 1 anal vein 2 branch, 2A = 2 anal vein.

Body (Figs. 1–3) moderately dorso-ventrally flattened, oblong in cross-section; glossy; glabrous. Colour entirely dark reddish-brown; antennae and tarsi reddish-brown. Rostrum moderately long, curved at point of antennal insertion, apical 2/3 flat, basal 1/3 convex, parallel-sided, with dense elliptical punctures along basal half; punctures becoming more circular and smaller towards apex, punctures separated by 1–2 x diameter of puncture; median furrow born in basal half and extending to vertex. Mouthparts (Figs. 10–13). Labium (Fig. 11) with 3- segmented palpi; basal segment with two dorsal setae, middle segment with one dorsal seta, apical segment lacking setae or with one seta, with 1–2 apical sensilla; prementum with two dorsal setae (one at each dorso- lateral corner); ligula strongly sclerotized, acute; mandibles weakly falcate; left mandible (Fig. 12) with two incisors and molar region, outermost incisor larger, with one seta on outer dorsal margin; right mandible (Fig. 13) similar to left one, except with one incisor and extended crenulate molar region; maxilla (Fig. 10) with 3- segmented palpi, basal two segments each with one seta near outer apical margin; palpiger with one dorsal seta; galea with large, paddle-shaped setae along mesal margin; lacinia free of setae, almost rectangular. Scrobes well-defined, wide, deep, triangular, covering basal 2/3 of rostrum and extending to slight distance (~length of antennal pedicel) before anterior margin of eyes; point of antennal insertion at basal 1/3 of rostrum; dorsal margins flush with dorsal margin of eyes, ventral margins extending beneath eyes. Antennae chestnut brown; glossy; long, stout, robust; scape clavate, robust, glabrous, glossy, passing posterior margin of eye, slightly bent upwards at distal one-half, shorter than funicle and club combined, more than 1.5 times as long as diameter of eye, smooth along basal 1/2 and with small sparse punctures along distal 1/2; funicle with 7 compact antennomeres (including pedicel), with long, sparse, sub-erect, pale-yellowish setae, first antennomere (pedicel) robust, longer than wide, second antennomere small and short, antennomeres 3–7 sub-clavate, wider than long; club composed of 3 antennomeres and apical annulation (appearing as being with 4 antennomeres), ovate, covered with mixture of short, appressed, sub-erect and long, sparse, erect setae. Head oval, dorsally constricted behind eyes at distance of

552 · Zootaxa 4365 (5) © 2017 Magnolia Press OMAR ET AL. 1/2 x diameter of eye; large, deep, and densely punctured before constriction, with small, shallow, sparse punctures after constriction; vertex convex in lateral view; frons as broad as base of rostrum, slightly depressed, with median furrow extending from distal extension of antennal scrobe to just beyond posterior margins of eyes, as viewed dorsally. Eyes dull black, convex, sub-circular, widely separated dorsally, laterally located before base of rostrum. Pronotum longer than wide, constricted anteriorly forming broad collar; flat dorsally, glabrous, glossy, with moderately dense, deep, circular punctures on whole surface, punctures separated by approximately 2–4 x puncture diameter; basal margin sub-linear, lateral margins sub-parallel. Mesonotum (Fig. 16) with mesoscutum lacking defined posterior margin; axillary cord rounded; apex of mesoscutellum visible, rounded, with acute postero- medial margin. Scutellum oblong, dark brown, glossy, glabrous, longer than wide, convex. Metanotum (Fig. 17) with metascutum reaching postero-lateral margin of notum; scutellar groove reaching posterior margin of notum, anterior margin obtusely triangular; allocrista obtusely angled at antero-mesal angle; metascutellum large, each region on either side of scutellar groove triangular. Metendosternite (Fig. 19) with long, narrow, curved hemiductus, furcal arm narrow, apex shallowly bifid; anterior tendons inserted near base of furcal arms. Proventriculus as in Fig. 18. Elytra glossy; disc broadly convex, elongate; sides parallel until declivity, moderately tapered to apex; base sub-linear; striae with deep circular punctures; punctures separated by approximately 0.5 x puncture diameter or less, punctures larger and deeper at base, cutting borders of interstriae, becoming smaller and more sparse towards apex; stria 6 not reaching base; striae 1 and 2 extending until apex; stria 1 deeper than others, particularly at shoulder; stria 10 ending just before middle of elytra; intervals elevated, wider than striae, less so at base, sub-equal in width, widening slightly at declivity, with single row of fine punctures; intervals 4, 5 and 6 uniting at declivity; interval 9 elevated, connecting with interval 3 near apex; humeri truncate. Hindwings (Figs. 14–15) slender, lacking jugal area (anal lobe); rf mostly a light sclerotization, abbreviated apically, not reaching rcm; rcm fused into single uniform sclerotization, radial window absent; 1rs and 2rs triangular, approximately equal in size; R3 present, forming a narrow, sclerotized stripe; Cu distant from posterior margin of wing (af continuing to margin); rm partially present as a faint sclerotization; rms absent

(or at least indiscernible); 2A simple; 1A2 partially present, faint. Thoracic sterna glossy, with deep circular punctures on whole surface; prosternum with punctures separated by 0.5–1 x diameter of puncture; procoxae separated by approximately 0.5 x diameter of coxa; mesoventrite with punctures separated by 1 x diameter of puncture; mesocoxae separated by distance greater than diameter of coxa; metaventrite with punctures separated by 1–2 x diameter of coxa, median furrow along posterior one-half; metacoxae separated by distance less than diameter of coxa. Legs (Figs. 4–9). Femur longer than tibia, robust at distal two-thirds, slightly laterally compressed, sparsely punctured; tibiae with sparse elliptical punctures; uncus at outer apical angle approximately same length as tarsomere five (longer and more robust in protibia of males); with small mucro at inner apical margin; fore-tibia at inner distal surface before uncus with grooming device consisting of patch of long setae; mid- and hind-tibiae with long sub-erect setae along distal 1/3; tarsi with distal margins bearing pale, sub-erect setae; tarsomere 1 longer than wide; 2 normal; 3 slightly bilobed or notched; 5 long, clavate, smooth, glossy, slightly laterally compressed; pretarsal ungues (claws) simple, separated, curved. Abdomen. Tergites (Figs. 20–23). Tergite VII in female (Figs. 20–21) with rows of stridulatory plectra, in male (Figs. 22–23) bearing more developed stridulatory plectra. Ventrites with sparse, circular punctures; 1 and 2 wide, clearly separated with shallow furrow. Punctures on ventrites 1–2 separated by distance approximately 2–3 x puncture diameter; on 3 and 4 sub-equal in width, with row of punctures along inner basal margin. Ventrite 5 rounded at apex and with sparse punctures. Male terminalia and genitalia (Figs. 24–31). Spiculum gastrale of sternite IX (Figs. 29–30) broadly curved, with expanded, hood-like, flattened apex; base narrow, broadly bifid. Sternite VIII divided, large; with each sclerite bearing about 5 setae at apico-lateral margins; and with dense covering of short and long, acute, triangular cuticular spines on apical membranous area. Tegmen (Fig. 31) complete, narrow; dorsal parameroid lobes absent; manubrium short, 1/4–1/5 x length of tegmen and with central longitudinal rib bearing 2 elongate spines at posterior margin. Penis (Figs. 24–28) with median struts (temones) slightly more than 3 x length of median lobe; apical margin of median lobe bearing few setae; endophallus (internal sac) simple, lacking any sclerotizations. Female terminalia and genitalia (Figs. 32–35). Gonocoxites (Figs. 33–34) oblong; styli elongate, narrow; each stylus bearing about 6 apical setae. Spermatheca (Fig. 35) falciform; with globular base and strongly curved and narrowing apex. Sternite VIII (Fig. 32) elongate; spiculum (spiculum ventrale) long, narrow, about 3/4 of

A NEW SPECIES OF MACRORHYNCOLUS WOLLASTON, 1873 Zootaxa 4365 (5) © 2017 Magnolia Press · 553 length of base; base of spiculum bifurcate; arms of base extending nearly to posterior margin of sternite; posterior margin of sternite with many setae. Etymology: The specific epithet is an adjective derived from the Latin word spina, which means spine, and refers to the large uncus on the protibiae.

FIGURES 16–23. Macrorhyncolus spinosus. 16, mesonotum; 17, metanotum; 18, proventriculus (dissected open); 19, metendosternite arrow pointing to anterior tendon; 20, female tergum; 21, enlargement of female tergite 7 showing reduced stridulatory plectra (arrow); 22, male tergum; 23, enlargement of male tergite 7 showing developed stridulatory plectra (arrow).

554 · Zootaxa 4365 (5) © 2017 Magnolia Press OMAR ET AL. FIGURES 24–28. Macrorhyncolus spinosus, male terminalia. 24, aedeagus, dorsal view (photomicrograph); 25, ditto with tegmen removed; 26, aedeagus dorsal view; 27, aedeagus lateral view (photomicrograph); 28, ditto.

A NEW SPECIES OF MACRORHYNCOLUS WOLLASTON, 1873 Zootaxa 4365 (5) © 2017 Magnolia Press · 555 FIGURES 29–35. Macrorhyncolus spinosus. 29–31, male terminalia. 29, 8th and 9th sternites dorsal view; 30, ditto (photomicrograph); 31, tegmen dorsal view. 32–35, female terminalia. 32, 8th sternite; 33, coxites, styli and bursa copulatrix; 34, female terminalia (photomicrograph); 35, spermatheca.

556 · Zootaxa 4365 (5) © 2017 Magnolia Press OMAR ET AL. Host plant: Collected from under the bark of a dead pine tree from Yunnan Province. Distribution: Yunnan Province, southern China. Sexual dimorphism: The rostrum of the females is longer than that of the males, and the protibial uncus is longer and stouter in males. Ecological notes on other species: The genus Macrorhyncolus contains five species. Macrorhyncolus crassiusculus was collected by Wollaston among fir trees and fungi on the Japanese island Honshu. Iwata & Banno (1995) found it in a nest of Reticulitermes speratus (Kolbe, 1885) on a fallen pine tree among a small number of termites. They also reported that when the species and termites were released into a petri-dish, it remained calm at a certain distance from the termite crowd. Macrorhyncolus littoralis was first found on the underside of a log on the beach in Tairua, New Zealand. Kuschel (1972) found it to be extremely common at or near the seashore on driftwood throughout New Zealand, and he believed that it is highly likely that it can be found more broadly in Australia, not only in Victoria and Tasmania. Kuschel (1969) also found a single specimen on a piece of driftwood at Chepu, Chiloe, on an uninhabited stretch of the west coast of Chile. Telfer (2007) collected it under driftwood on the beach on the Kent coast, Ireland, and surmised that while it is a flightless species, colonies are likely capable of long-range dispersal by sea on driftwood. Dispersal in this manner has been suggested for cossonines by Oevering & Pitman (2001). Macrorhyncolus ventilaginis was collected from Ventilago calyculata Tulasne, Ficus glomerata Roxb. and an unidentified wood in Dehra Dun (Lachiwala, Nakronda), United Provinces, India.

Discussion

The genus Macrorhyncolus was previously known from four species: one from Japan, one from India, one from Ceylon (Sri Lanka) and one from New Zealand. In terms of biogeographical zones, two species were described from the Palearctic Region and one from the Oriental Region, and only one species from the Australian Region. The fifth species, described here from southern China, is considered the first record of the genus for China. The new species, M. spinosus, differs from other Palearctic species of the genus Macrorhyncolus as detailed in the key above as well as from M. littoralis and M. crassitarsis as follows: M. spinosus is winged and has larger eyes whereas M. littoralis is apterous (Welch 1990), but M. crassitarsis has small eyes, short scapes, the point of antennal insertion is at the middle of the rostrum, and the base of the pronotum is narrower than the elytral base. Also, the tarsi are short and stout in M. crassitarsis, and the 3rd tarsomere is simple (not bilobed).

Acknowledgments

We are sincerely grateful to Dr. Max Barclay, Keita Matsumoto and the Natural History Museum, London for providing high resolution photos of the holotype of Macrorhyncolus crassiusculus for comparison. Also, we are sincerely thankful to Prof. Christopher H. C. Lyal, PhD (Department of Entomology, The Natural History Museum, London) for providing constructive comments. A genuine gratitude is extended to Dr. Rich Leschen (Landcare Research, Crown Research Institutes, New Zealand) and Dr. Keita Matsumoto (Natural History Museum, London) for providing references and translation. The first author is thankful and grateful to Dr. Ning Liu (Institute of Zoology, Chinese Academy of Science) for her kind help and efforts with this study, and other laboratory staff kindly assisted in it as well. This research was supported by the Chinese Academy of Sciences President’s International Fellowship Initiative for Visiting Scientists (Grant No. 2015vbb042) and the Chinese Academy of Sciences visiting professorship for senior international scientists (Grant no.2009S2-30). Partial funding was provided by the University of Kansas Entomology Endowment, the US National Science Foundation grant DEB- 1110590 (to M.S. Engel, P. Cartwright, and S.R. Davis), and continued support was provided by the Gerstner and Kalbfleisch postdoctoral fellowships through the American Museum of Natural History and Richard Gilder Graduate School during the completion of this study.

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