Curculioninae Latreille, 1802 305 Schultze, A. (1902): Kritisches Verzeichniss der bis 3.7.10 Curculioninae Latreille, 1802 jetzt beschriebenen palaearctischen Ceuthor- rhynchinen. – Deutsche Entomologische Zeitschrift Roberto Caldara , Nico M. Franz, and Rolf 1902: 193 – 226. G. Oberprieler Schwarz, E. A. (1894): A “ parasitic ” scolytid. – Pro- ceedings of the Entomological Society of Washington 3: Distribution. The subfamily as here composed (see 15 – 17. Phylogeny and Taxonomy below) includes approx- Scudder, S. H. (1893): Tertiary Rhynchophorous Coleo- ptera of the United States. xii + 206 pp. US Geological imately 350 genera and 4500 species (O’ Brien & Survey, Washington, DC. Wibmer 1978; Thompson 1992; Alonso-Zarazaga Stierlin, G. (1886): Fauna insectorum Helvetiae. Coleo- & Lyal 1999; Oberprieler et al. 2007), provisionally ptera helvetiae, Volume 2. 662 pp. Rothermel & Cie., divided into 34 tribes. These are geographically Schaffhausen. generally restricted to a lesser or larger degree, only Thompson, R. T. (1973): Preliminary studies on the two – Curculionini and Rhamphini – being virtually taxonomy and distribution of the melon weevil, cosmopolitan in distribution and Anthonomini , Acythopeus curvirostris (Boheman) (including Baris and Tychiini only absent from the Australo-Pacifi c granulipennis (Tournier)) (Coleoptera, Curculion- region. Acalyptini , Cionini , Ellescini , Mecinini , idae). – Bulletin of Entomological Research 63: 31 – 48. and Smicronychini occur mainly in the Old World, – (1992): Observations on the morphology and clas- from Africa to the Palaearctic and Oriental regions, sifi cation of weevils (Coleoptera, Curculionidae) with Ellescini, Acalyptini, and Smicronychini also with a key to major groups. – Journal of Natural His- extending into the Nearctic region and at least tory 26: 835 – 891. the latter two also into the Australian one. Styph- – (1996): The species of Phaenomerus Sch ö nherr lini are restricted to the Palaearctic region and the (Coleoptera: Curculionidae: Zygopinae) of the Aus- seemingly related Itini and the monobasic Acen- tralian region. – Invertebrate Taxonomy 10: 937 – 993. trusini to only its western part. Ancylocnemidini , Ulmer, J. B., Duncan, R. E., Prena, J. & Pe ñ a, J. E. Microstylini , Nerthopini , and Ochyromerini are (2007): Life history and larval morphology of Eurhi- largely African but extend into the Oriental region, nus magnifi cus Gyllenhal (Coleoptera: Curculion- the last extensively so and just reaching into Aus- idae), a new weevil to the United States. – Neotropi- tralia, whereas the small tribes Diabathrariini , and cal Entomology 36: 383 – 390. Ulomascini are entirely African. Nine tribes only Urban, C. (1925): Der Veilchenr ü ß ler. – Entomologische Bl ä tter 21: 139 – 141. occur in the New World, Camarotini , Ceratopo- Wagner, H. (1936): Vorstudien zu einer monogra- dini, Otidocephalini, Piazorhinini , Thecesternini , phischen Bearbeitung der Ceuthorrhynchinen Erodiscini , Prionobrachiini , Pyropini, and Sphaeri- Mitteleuropas (Col. Curcul.). – Entomologische Blä t- opoeini, the last four restricted to the Neotropical ter 35: 161 – 170. region. Cranopoeini , Cryptoplini , Storeini , and – (1938): Monographie der pal ä arktischen Ceuthor- Viticiini are limited to the Australo-Pacifi c region, rhynchinae (Curcul.). – Entomologische Bl ä tter 34: and the last two reaching north into Indonesia and 145 – 172. southern Japan. Only two tribes appear to have a Weng, J.-L., Nishida, K., Hanson, P. & Lapierre, L. Gondwanan distribution, Eugnomini occurring (2007): Biology of Lissoderes Champion (Coleoptera, in Australia and South America and Derelomini in Curculionidae) in Cecropia saplings inhabited by Africa and South to Central America, but the com- Azteca ants. – Journal of Natural History 41: 25 – 28. position of these tribes as including genera from Wibmer, G. J. & O ’ Brien, C. W. (1986): Annotated these disparate regions needs verifi cation. checklist of the weevils (Curculionidae sensu lato ) of South America (Coleoptera: Curculionoidea). Biology and Ecology. Adults of Curculioninae – Memoirs of the American Entomological Institute 39: are usually found on fl owers, fruits, leaves, and i – xvi, 1 – 563. branches of many and various herbaceous plants – (1989): Additions and corrections to annotated and trees in nearly all habitat types, ranging from checklists of the weevils of North America, Central low plains to high mountains and from deserts to America, and the West Indies, and of South Amer- tropical pluvial forests. The larvae are predomi- ica. – Southwestern Entomologist Suppl. 13: 1 – 49. nantly endophytic in reproductive plant parts such Zherikhin, V. V. & Egorov, A. B. (1991): Shuki-dol- gonosiki (Coleoptera, Curculionidae) dalnego vostoka as fl ower and fruit buds, seeds, and fruits, but some SSSR. 164 pp. Akademija Nauk SSSR, Dalne- develop in shoots (in which some incite galls) or vostochnoe Otdelenie, Biologo-pochvennyj Insti- mine in leaves. The slimy, slug-like larvae of Cio- tut, Vladivostok [1990]. nini are exceptional in the subfamily for being Zherikhin, V. V. & Gratshev, V. G. (1995): A compara- ectophytic. No species, however, live in aquatic tive study of the hind wing venation of the super- habitats. The biology of the subfamily varies con- family Curculionoidea, with phylogenetic implica- siderably between the tribes, and sometimes also tions. Pp. 634 – 777 in Pakaluk J. & Slipinski, S. A. between their genera, but several tribes are associ- (eds.). Biology, Phylogeny, and Classifi cation of Coleo- ated with only one or a few plant families and some ptera: Papers Celebrating 80th Birthday of Roy A. Crow- genera can be highly host-specifi c on a single plant son . Volume 2. Muzeum i Instytut Zoologii PAN, genera or even species. Patterns of host associa- Warszawa. tion are, however, often confused by the uncertain 306 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler composition of many of the tribes and by the unre- (blotched), with the pattern often intricate and spe- liability of some published host records. Curcu- cies diagnostic (Scherf 1964). Generally there is only lionine weevils may often be found on fl owers one mine per leaf, but several larvae of the small or foliage of plants on which their larvae do not Rhamphus species may mine in the same leaf (Scherf develop, and such fortuitous occurrences must be 1964). Pupation occurs either in the mines (Rham- distinguished from true host associations in which phini, Styphlini) or in litter on the ground or in soil the plants serve as substrates for oviposition and (Storeini). The larvae of Anoplus Germar and some larval development. The lists of plants on which Rhamphus and Rhynchaenus species cut out pieces of weevils were only collected, without established leaf, which they stick together and in which they evidence of a reproductive interaction, may portray fall to the ground and pupate (Scherf 1964). Larval false patterns of host association that are perpetu- habits associated with leaf mining are probably as ated over time and can have serious implications varied and intricate in other leaf-mining groups. for applied fi elds of research, such as biological The induction of galls appears to be more com- control. The known biology and host associations mon in Curculioninae than in other subfami- of the various tribes are summarized under their lies, with the exception of the Ceutorhynchini treatment below, but some of these associations are (Conoderinae). Korotyaev et al. (2005) compiled a subject to verifi cation. useful summary of galling habits in the subfamily, Two particular lifestyles are more common in discussing also ecological and evolutionary aspects Curculioninae than in other subfamilies, namely that seem to make this habit more prevalent in mining in leaves and inciting galls. The bionom- Curculioninae than in other subfamilies. Galling ics of leaf-mining insects was reviewed by Hespen- appears to occur more commonly on herbaceous, heide (1991), who mentioned some aspects of the short-lived plants and in rapidly growing organs biology of the best-known curculionine leaf min- (mainly young, slender stems but also roots), attri- ers, the Rhamphini , but did not cover the habit butes characteristic of nonwoody plants growing comprehensively. The knowledge of leaf mining in temperate or arid regions with short growing among European weevils is well covered by Scherf seasons. The lifestyle seems to be derived from stem (1964), who recorded it to occur in the tribes Ano- boring (no transition from leaf-mining known) and plini, Rhamphini, and Styphlini, whereas May enables a species to exploit very slender stems and (1993) provided an account of known leaf-mining similar plant organs that it could not utilize other- genera of Storeini in New Zealand. The habit fur- wise. The larvae of some Smicronyx Schoenherr, on ther occurs in the small South American tribes completion of their development, leave their galls Camarotini , Piazorhinini , and Prionobrachiini on the stems of parasitic dodder ( Cuscuta ) and tun- and in the Australo-Pacifi c Viticiini , although it nel into the stem of its Vernonia host for pupation is poorly documented in these groups. Nothing (Anderson 1970). Generally, the closest relatives of appears known about this habit in Africa. Leaf- gall incitants develop in fl owers or seed capsules of mining weevil larvae tend to be fl at, with the head the same plant species, and no instances are known prognathous, deeply excised posteriorly and with of closely related species inducing galls on different reduced numbers of setae, and the body tapered host organs. Galling thus enables a greater number caudally and with dorsal and ventral or lateral of related species to exploit the same host but does ambulatory ampullae and/or covered with fi ne not form a defi nite feeding niche in any group. In asperities that enable the larva to crawl in its mine. central Europe, approximately 10% of the weevils May (1993) sketched a progression of increasing whose larval habits are known incite galls (Scherf specialization in New Zealand leaf-mining storeine 1964), and weevils (as Coleoptera as a whole) never genera, from the largely unmodifi ed larva (with a form a major component of gall-forming faunas. normal head capsule) of Phorostichus linearis (Broun) Some weevils also live as inquilines in galls induced via Peristoreus fl avitarsis (Broun) and Hypotagaea con- by other insects, such as cecidomyid fl ies and color (Broun), in which the head is weakly emargin- cynipid and tenthredinid wasps, in which they may ate, to that of Neomycta rubida Broun, in which the live as commensals or pseudoparasites. The exam- epicranium is deeply excised except for the median ples of curculionine gallers provided by Korotyaev endocarina. A similar head capsule occurs in the et al . (2005) largely comprise Palaearctic species of genus Geochus Broun (see Molytinae: Phrynixini ) the tribes Mecinini , Smicronychini , and Tychiini , as well as in Rhamphini ( Ixalma Pascoe, Rhamphus but galling is also known to occur in other tribes Clairville, Rhynchaenus Clairville; van Emden 1938; and in the southern hemisphere, for instance, in Scherf 1964; Lee & Morimoto 1996), in which a Anthonomini , Camarotini , Cryptoplini , Curcu- comparable cline of increasing morphological lionini, Eugnomini , Otidocephalini, Rhamphini , regression appears to occur (May 1993). The fl at Storeini , Thecesternini , and Sidomenia Laporte, larva of the undescribed Australian viticiine genus as detailed in the sections on these tribes below. Austrocis is similarly adapted to leaf mining, the Whether galling may be as common in tropical head also deeply V-shaped posteriorly (but without and in southern temperate regions as is it northern a protruding endocarina) and the body segments temperate ones remains to be investigated. each carrying a lateral pair of strong ampullae. Leaf Curculionine weevils are important pollina- mines are generally of two types, linear or round tors of several plants, in particular palms and basal Curculioninae Latreille, 1802 307 angiosperms (magnoliids), in which cantharophily & Morimoto (1995 a) described a species of Endae- is the dominant pollination system. Weevil pol- nidius from fl owers of Knema in Malaysia. In Win- lination of palms has been reported or indicated teraceae, the genus Exospermum in New Caledonia for the genera Acanthococos, Acrocomia, Astrocaryum, is exclusively pollinated by a species of Palontus Attalea, Bactris, Butia, Catoblastus, Chamaerops, Cocos, Kuschel (Pellmyr et al. 1990), also currently placed Cryosophila, Elaeis, Hydriastele, Phoenix, Phytele- in Storeini (Kuschel 1990 a), but the taxonomy of phas, Rhapidophyllum, Rhapis, Sabal, Salacca, Serenoa, all these weevil pollinators needs attention. Wee- Socratea, Syagrus, and Wettinia (Henderson 1986; vils have also been reported to be involved in the Silberbauer-Gottsberger 1990; Meekijjaroenroj & pollination of Amborellaceae (Thien et al . 2003), Anstett 2003) and, given the close association of the most basal lineage of angiosperms, but their Derelomini with several other palm genera (Franz identity (given as “ Cryptorhynchinae ” ) is in need of & Velente 2005), is probably more widespread verifi cation; they more likely also belong to a cur- in the family. In Africa and America, the wee- culionine tribe such as Ochyromerini or Storeini. vils involved in palm pollination belong to the At least in Eupomatia and Myristica, the association tribe Derelomini, whereas those in Asia and the between plant and weevil seems to be a nursery pol- Australo-Pacifi c region all seem to be Acalyptini lination mutualism, the pollinator breeding solely (Kojima & Morimoto 2005). The best-known case in the ovules of the host and thus being as depen- of weevil pollination of palms is that of the African dent on the plant, as the latter is on the weevil for oil palm, Elaeis guineensis, which is pollinated by the its pollination; also dubbed a system of pollination derelomine Elaeidobius kamerunicus (Faust), which by deceit (Armstrong 1997), this is similar to weevil Q: “deceipt” was introduced to Asia in 1981 to improve seed set pollination in cycads and palms. was changed in oil palms planted there (Krantz & Poinar 2004). Due to their generally high host specifi city, cur- to “deceit”. Weevils also pollinate several species of Cyclan- culionine weevils also constitute important con- Correct? thaceae and some Araceae (Franz 2003 a, b, 2004, trol agents of weedy plants throughout the world. 2006, 2007 a, b; Franz & Valente 2005). Weevil pol- O ’ Brien (1995) summarized the role of weevils gen- lination in magnoliids has been shown to occur in erally in the biological control of weeds but used the families Annonaceae, Eupomatiaceae, Myristi- examples mainly from the subfamilies Brachycer- caceae, and Winteraceae and is likely to occur also inae, Conoderinae, and Molytinae, as species of in other basal angiosperm families. In Annonaceae, especially the fi rst group have achieved singularly Momose et al. (1998) reported species of Endaenidius spectacular control of major aquatic weeds in sev- Morimoto and Endaeus Schoenherr (Ochyromerini) eral regions of the world. The various species of to visit fl owers of Uvaria (Cyathostemma ) in Malay- Curculioninae currently used to control weeds, as sia, evidently pollinating these plants (Zhou et al. enumerated by Julien & Griffi ths (1998), mostly 2009), and Ratnayake et al. (2006, 2007) similarly belong to the tribes Mecinini , Smicronychini , and found Polyalthia and Xylopia species in Sri Lanka Storeini . Mecinini make very successful biocon- to be pollinated by weevils identifi ed as Endaeus . trol agents of European species of toadfl ax (Linaria , Kojima & Morimoto (1995 a) described species of Plantaginaceae) in North America. The seed-feed- Endaeus and Endaenidius from fl owers of Polyalthia ing Rhinusa antirrhini (Paykull) and R. neta (Ger- in Malaysia and further species of Endaenidius from mar) were accidentally introduced from Europe those of Enicosanthum and Fissistigma, of the same into Canada and the United States in the early 20th family, whereas similar undescribed species of century and exert some biological control of yel- Ochyromerini are indicated to pollinate the annona- low toadfl ax (Linaria vulgaris) and Dalmatian toad- ceous vine Melodorum leichhardtii in Australia. In fl ax (L. genistifolia). Their effort was boosted in the Japan, species of Endaeus and Endaenidius also occur early 1990 s by the release in both countries of the in fl owers of Magnolia (Magnoliaceae) and Illicium stem-boring Mecinus janthinus Germar and the root- (Schizandraceae) (Kojima & Morimoto 1995 d), galling Rhinusa linariae (Panzer), the former now indicating that Ochyromerini may be involved well established but the latter less so. In contrast, in the pollination of several basal angiosperm Smicronychini have proved less successful in weed families. The monobasic family Eupomatiaceae, control. Smicronyx roridus Marshall and S. rufovitta- restricted to eastern Australia and New Guinea, tus Anderson failed to establish on dodders (Cus- and the apparent sister group of Annonaceae are cuta, Convolvulaceae) in the Bahamas, Barbados, pollinated exclusively by weevils of the genus and Bangladesh (Julien & Griffi ths 1998), whereas Elleschodes Blackburn, currently placed in the tribe the stem-galling S. guineanus Voss and S. umbrinus Storeini (q.v. ) (Armstrong & Irvine 1990), the fl ow- Hustache have more of an impact on the parasitic ers emitting fragrances consisting of short-chain witchweed Striga hermonthica (Scrophulariaceae) in fatty esters that attract the pollinators (Bergströ m Africa (Anderson & Cox 1997) and S. dorsomaculatus et al . 1991). In the nutmeg family Myristicaceae, Cox similarly so on Alectra vogelii , although they are Myristica insipida in Australia is also pollinated by not able to control these weeds suffi ciently (Sau- weevils (Armstrong 1997), and apparently several erborn & M ü ller-St ö ver 2009). The Mexican Smi- species are involved (Armstrong & Irvine 1989 a, b), cronyx lutulentus Dietz, established in a small area but none of them are described and their taxo- in Australia on Parthenium hysterophorus (Astera- nomic affi nities are as yet unclear, whereas Kojima ceae), has also not had a major impact on this weed 308 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler

(Dhileepan & Strathie 2009). Similarly, Sibinia fasti- ending before anterior margin of eyes, very short giata Clark ( Tychiini), released from 1997 to 2002 in Rhamphini , almost vertical in Microstylini. Eyes against Mimosa pigra (Fabaceae) in northern Austra- usually large, rounded to elliptical, fl at to distinctly lia, failed to establish (Ostermeyer & Grace 2007), convex, sometimes nearly contiguous dorsally (e.g., whereas Anthonomus santacruzi Hustache ( Anthono- Ancylocnemidini : Ancylocnemis) or ventrally (e.g., mini) is deemed to be the most promising biocon- Acentrusini , Nerthopini : Nerthops Schoenherr, trol agent of Solanum mauritianum (Solanaceae) in Smicronychini), frons slightly wider than rostrum South Africa (Olckers 2009). Species of Storeini at base to very narrow (e.g., Cionini , Rhamphini, released in South Africa to control the invasive Aus- Ochyromerini). Antennae usually inserted near tralian shrub Hakea sericea (Proteaceae) have also middle of rostrum but often more basally in female been highly successful. The seed-feeding Erytenna than in male (at base of rostrum in Rhamphini); consputa Pascoe, released in 1972, caused a reduction funicles fi ve- (e.g., Cionini, Mecinini) to seven-seg- in seed production of up to 90%, whereas the shoot- mented, usually with fi rst two segments distinctly boring Cydmaea binotata Lea, released in 1979, ini- longer than others. Mouthparts generally phanero- tially established but failed to have any long-term gnathous (e.g., Ting 1936; Franz 2006). Mandibles impact and appears to have largely died out (Gor- without deciduous processes; sometimes moving don 1999). A third species, the fl owerbud-feeding vertically (some Curculionini); typically with two Dicomada rufa Blackburn, was released in 2005 to or three teeth; outer margin with or without setae. augment the biological control of the plant (Kluge Maxillae with galea, lacinia, and palpifer fused; & Gordon 2004). A recent comparison of Hakea seri- outer margin with one or more “ lacinial teeth ” , cea infestation levels in 1979 and 2001 (Esler et al . palps two- or three-segmented. Labium variously 2010) revealed that, together with the seed-feeding shaped, with or without setae along lateral and moth Carposina autologa Meyrick and mechanical apical margins; ligula absent to projecting beyond eradication, Erytenna consputa caused a 64% decrease labial palps, palps two- or three-segmented. in the distribution of this weed in South Africa and Pronotum slightly to distinctly convex, some- a signifi cant decrease in its overall density. When times tuberculate (e.g., some Anthonomini , Cio- the impact of an introduced plant is ambivalent, nini), at base slightly to distinctly narrower than weevils may function both as biocontrol agents and elytral bases; sides more or less curved, usually as pests. Thus, the seed-feeding Ochyromera ligustri widest at basal half, rarely carinate (some Dere- Warner (Ochyromerini) is regarded as harmful to lomini), rarely with ocular lobes (some Storeini ), Chinese privet ( Ligustrum sinense , Oleaceae) in North sometimes abruptly restricted at apex to form a America, where the plant is used in landscaping, narrow neck or at base to become pedunculate but benefi cial in southeastern states where it is con- (Erodiscini , Otidocephalini). Prosternum with api- sidered an invasive weed (Cuda et al . 2008). cal margin regularly transverse, sometimes more or less concave (e.g., Cionini, Storeini sensu lato ) or Morphology, Adults (Fig. 3.7.10.1 – 3.7.10.4). with weak to deep canal into which the rostrum Length 1 – 12 mm (rostrum excluded). Body broadly fi ts in repose (e.g., Smicronychini, Storeini sensu ovate to globular (e.g., Anthonomini , Camarotini , stricto , some Cionini, Mecinini, and Rhamphini), Cionini , Derelomini : Anchylorhynchus Schoenherr, in this case, fore coxae separated, walls of canal Microstylini, Ochyromerini) to narrowly elongate in Storeini sensu stricto excised to form a cleft of (e.g., Erodiscini , Mecinini , Otidocephalini), slightly “ peep-hole ” (Fig. 3.7.10.2 A and B), through which fl attened to markedly convex. Integument black to the weevil may be able to see when the rostrum is brown to reddish to pale yellowish, rarely metallic folded down. Scutellum usually clearly visible but blue (Cionini: Nanomicrophyes Pic, Mecinini: Mecinus sometimes indistinct (e.g., Smicronychini, some Germar, Pyropini: Pyropus Schoenherr), covered Storeini). Elytra with interstriae usually wider than with more or less adpressed, recumbent to erect, striae, fl at to moderately convex, sometimes at hair-like to rounded scales, which vary in color least in part distinctly convex (e.g., Cionini, Dere- (white, yellowish, reddish, brown); color uniform lomini), rarely with tubercles (some Anthonomini, in some taxa; species of other taxa with contrast- Cionini, Cryptoplini, Storeini); striae usually well ing scale patterns, sometimes with metallic sheen, defi ned and with deep punctures (obscure in some or scales regularly arranged in one to two series on Cionini), sutural fl anges unequal. Hind wings usu- each elytral interstria. ally well developed, rarely reduced [e.g., Styph- Head with rostrum mostly long to very long lini, Cionini: Nanomicrophyes , and Tychiini : Tychius and slender, in lateral view more or less curved and (Apeltarius Desbrochers)]. Metanepisternal sutures narrowed from base to apex, at base dorsally some- without sclerolepidia. Legs with femora weakly to times concave (e.g., Anthonomini , Curculionini , distinctly clavate, often with weak to very robust Ochyromerini) to distinctly set off from head tooth, hind femora with apodeme for jumping (Smicronychini ), mostly sexually dimorphic with in Rhamphini; tibiae with apex dorsoventrally female rostrum longer, narrower, and smoother, oblique, usually without spurs (except Nothofagi- rarely short, broad, fl attened to fl ared (e.g., Dia- nus Kuschel), fore and middle tibiae at inner apex bathrariini, Microstylini , Ulomascini ); scrobes usually clearly imperfectly uncinate (Thompson oblique to parallel to long axis of rostrum and 1992) or mucronate, especially in male, fore tibiae Curculioninae Latreille, 1802 309

Fig. 3.7.10.1 Ventrites. A, Anthonomus sp. (Anthonomini), B, Cionus sp. ( Cionini); C, Curculio caryae (Horn) ( Curculionini); D, Perelleschus carludovicae (G ü nther) ( Derelomini); E, Ellescus ephippiatus (Say) (Ellescini); F, Mecinus pyraster (Herbst) ( Mecinini); G, Sicoderus sp. ( Erodiscini); H, Pyropus pusillus Pascoe (Pyropini); I, Myrmex sp. (Otidocephalini); J, Smicronyx sp. ( Smicronychini); K, Sibinia sp. ( Tychiini); L, Ulomascus marshalli Ghesqui é re ( Ulomascini). Not at the same scale. 310 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler

Fig. 3.7.10.2 Structural details. A, Storeus sp. ( Storeini sensu stricto); cleft wall (arrow) of prosternal canal, lateral view; B, Aporotaxus kanalensis Perroud (Storeini sensu stricto); setiferous hole (arrow) in wall of prosternal canal, lat- eral view; C, Leucomelacis quadrinotatus Lea (Storeini sensu stricto), left hind tibia, apical view; D, Sellechus tibialis Lea (Storeini sensu lato ), foreleg of male, caudal view; E, Cratoscelocis foveicollis Lea ( Cranopoeini), left hind leg, ventral view; F, Micraonychus casuarinae Lea (Storeini sensu lato ), tarsus, dorsal view. Not at the same scale.

sometimes sexually dimorphic (Fig. 3.7.10.2 D), Abdomen (Fig. 3.7.10.1) with posterior mar- hind tibiae with uncus often small or absent, espe- gin of ventrites 2, 3, and 4 straight to concave, cially in female (in some Cionini and Derelomini sometimes laterally extended caudad (angled) unci absent on all tibiae) (Fig. 3.7.10.2 C, E, and (2 – 4 in Cionini , Mecinini , 2 – 3 in Ellescini , Ochy- F), uncus sometimes displaced outward and inner romerini, Rhamphini , Tychiini : Lignyodina , only apical angle then with praemucro fl anked by tufts 2 in Tychiina), 5 sometimes with median fringe of of long setae (e.g., Mecinini, Microstylini ); tarsi hair-like scales and/or fovea (Mecinini, Tychiini) with tarsomere 3 usually bilobate and wider than and seldom with teeth (Mecinini: Miarus Schoen- 2, onychium (tarsomere 5) usually long, some- herr). Tergites VII and VIII usually hidden by ely- times short, reduced, or fully lost (some Anoplini , tra but sometimes clearly visible as pygidium (e.g., Diabathrariini , Storeini, Viticiini ); claws simple Acalyptini , Curculionini , Mecinini, Microstylini ; to appendiculate to toothed or bifi d, divergent males only in some Derelomini, and Tychiini); to divaricate, mostly separate but often connate tergite VII in some taxa with plectrum consisting at base (e.g., Cionini, most Mecinini and Smicro- of one or more pairs of raised, setiferous tubercles nychini, various other genera) to one claw reduced scraping against inner apical fi le on underside of (Cionini, some Mecinini) or fully fused (single) elytra (elytro-tergal stridulation system 1 of Lyal & (some Cionini, Cryptoplini: Haplonyx Schoenherr) King 1996), as far as studied present in both sexes to variously reduced or fully lost (e.g., Cryptoplini: in Camarotini : Prionomerina , Ellescini: Ellescus Cryptoplus Erichson, Viticiini: Aviticis Kojima & Dejean, Erodiscini , Otidocephalini, Rhamphini, Morimoto). Proventriculus generally well devel- and Smicronychini but apparently only in males in oped, with eight strong basal plates and paired Acentrusini (Acentrus Desmarest), Ellescini: Doryto- rows of internal brushes, but weakly developed in, mus Germar, Prionobrachiini , Storeini , Tychiini, e.g., Rhamphini and absent in Cryptoplini. and Pachytychius Jekel (Lyal & King 1996), system 2 Curculioninae Latreille, 1802 311

Fig. 3.7.10.3 Male terminalia of Cyclanthura laticola Franz ( Derelomini). A, Tergum VIII, ventral view; B, sternum VIII, ventral view; C, Spiculum gastrale, ventral view; D, tegmen, ventral view; E, aedeagus, ventral view (with dark ampullate sclerite); F, aedeagus, lateral view. Scale bars 0.1 mm. 312 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler

Fig. 3.7.10.4 Female terminalia of Cyclanthura laticola Franz ( Derelomini). A, Tergum VIII, ventral view; B, tergum IX, ventral view; C, sternum VIII, ventral view; D, gonocoxites + styli, ventral view; E, spermatheca + gland, lateral view. Scale bars 0.1 mm. Curculioninae Latreille, 1802 313

(positions of plectrum and fi le reversed) reported sensillum sometimes absent; epipharynx with in Anoplini, and Derelomini ( q.v.) . Aedeagus sub- three to six pairs of anteromedian setae ( ams ), four cylindrical to fl attened (strongly modifi ed in pairs of median spines ( msp ), two to three pairs of Mecinini: Miarus), usually symmetrical (distinctly anterolateral setae ( als); labral rods (tormae) short asymmetrical in some Rhamphini and Tychiini: to long, separated to united posteriorly. Antennae Lignyodina), with long apodemes (temones) usu- with membranous basal segment bearing a coni- ally fused to aedeagus (articulated in Tychiini), cal, blunt to elongate sensorium and several min- endophallus without or with sometimes complex ute sensory processes. Mandibles short to elongate, internal sclerites, occasionally with more or less more or less triangular, brown, each with two api- long fl agellum. Tegmen usually with complete cal teeth, one pair of mandibular setae and one sen- ring (clearly incomplete in Lignyodini and Tychi- sillum. Maxillary palps two-segmented, segments ini), with distinct apodeme and basal plate usually variable in length, bearing one to two sensilla, basal consisting of two lateral sclerotized parts. Ster- segment with one short lateral seta, apical segment nite VIII of female with long apodeme, with basal with minute papillae. Labium with more or less plate usually sclerotized at sides. Bursa copulatrix complete premental sclerite, anterior and posterior membranous, rarely sclerotized (e.g., in Mecinini: median extensions mostly pronounced; premen- Miarus ). Spermatheca usually falciform, sometimes tum with one pair of long setae, two pairs of short with body distinctly globose (e.g., Anthonomini , ligular setae and two pairs of sensilla; postmentum Cionini), spermathecal duct usually membranous with two or three pairs of more or less long setae; (partially sclerotized, e.g., in Mecinini: Gymnetron palps one- or two-segmented. Schoenherr), base of spermathecal gland sclero- Thorax narrower than or as wide as abdomen. tized (funnel-shaped) in Cryptoplini, and Tychiini: Thoracic spiracles unicameral or bicameral. Pro- Sibinia Germar ( Microtychius Casey) (various sources, notum with 9 – 11 pairs of minute to long setae, supplemented by the authors’ studies of relevant prodorsum, and epipleural lobes of mesothorax characters of numerous curculionine taxa). and metathorax each with one pair of short to long setae. Pleural areas of prothorax with two to Morphology, Larvae (Fig. 3.7.10.5). Body slen- fi ve pairs of short to long setae, of mesothorax and der to robust, elongate, slightly to strongly curved; metathorax each with one pair of long setae. Ster- integument usually white, rugulose to smooth; num of each thoracic segment with two to four tuberculate to spinose asperities usually distrib- pairs of setae. uted over entire body, sometimes more conspicu- Abdomen with eight pairs of lateral unicam- ous on thorax and abdomen. Head free, pale yellow eral or bicameral spiracles, spiracle VIII located on to dark brown to black; usually wider than long, elongated dorsal protuberances in Ochyromerini. broadly rounded or narrowing dorsally or ven- Segments I – VII, each with three dorsal folds, some- trally, posteriorly emarginate in leaf-mining taxa; times segment I with only two folds (e.g., Mecinini , anterior stemmata present, black to transparent Rhamphini ); segment VIII, if present, always with (lacking pigmentation); posterior stemmata ves- two folds; all segments with one short to long pair tigial to absent. Frontal sutures usually distinct of antedorsal setae and two to fi ve pairs of post- throughout length, closed anteriorly by fronto- dorsal setae. Epipleuron with one long seta and epicranial bridge (bracon). Epicranial suture about one short seta. Pedal areas each with one long seta. half as long as head capsule, short when head pos- Eusternum with two long setae. Segment IX with teriorly emarginate. Median endocarina almost fi ve to seven pairs of short setae. Anus subterminal always distinct, sometimes widest at apex, occa- to terminal, anal folds with several pairs of minute sionally very short or absent (e.g., Eugnomini). setae (Ahmad & Burke 1972; Anderson 1973; Clark Epicranium with four to fi ve pairs of dorsal setae et al. 1978; Cussigh 1992; May 1993, 1994; Mar- (des), des1 often short, des2 and des4 sometimes min- valdi 2003). ute to absent, des3 and des5 longest, with one pair of sensilla behind des1 and another between des4 Morphology, Pupae (Fig. 3.7.10.5). Color white and des5; laterally usually with one or two pairs of to occasionally yellowish, sometimes with several short to long setae ( les ), one pair of sensilla usually darker regions. Head with distirostral setae absent in front of les 1; ventrally with two pairs of short to or arranged in one or two pairs, short, two or three moderately long setae ( ves); posteriorly with two to pairs of basirostral setae and one pair of interorbital four pairs of minute setae ( pes ) and one pair of sen- setae; pair of supraorbital setae present or absent; silla. Frons with three to fi ve pairs of frontal setae frons with one or two pairs of setae. Pronotum (fs), fs1 and fs2 sometimes absent, fs3 minute, fs4 with one pair of anteromedian setae, two to three long, fs5 long or short to vestigial; with two pairs (rarely one) pairs of anterolateral setae, one pair of of frontal sensilla, one laterally or posteriorly of discal setae, one pair of posteromedian setae, one to fs3 and the other between fs3 and fs4 . Clypeus four pairs of posterolateral setae. Mesonotum and wider than long, with two pairs of short setae ( cls ) metanotum with anteronotal setae usually absent, and two sensilla located between or just in front of with one to three pairs of posteronotal setae. Abdo- cls . Labrum with three pairs of setae (lms ) of vari- men with or without one pair of anterotergal setae, ous lengths, paired sensilla present but median one to fi ve pairs of discotergal setae, one pair of 314 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler

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Fig. 3.7.10.5 Pre-imaginal stages. A, Curculio sp. (Curculionini), larva, lateral view; B, Curculio sp. (Curculionini), larva, head, anterior view; C, Cryptoplus sp. (Cryptoplini), larva, lateral view; D, Cryptoplus sp. (Cryptoplini), larva, head, anterior view; E, Myossita banksiae Lea ( Eugnomini), larva, lateral view; F, Myossita banksiae (Eugnomini), larva, head, anterior view; G, Daeneus annae Oberprieler (Ochyromerini), larva, lateral view; H, Daeneus annae (Ochyromerini), larva, head, anterior view; I, Elleschodes sp. ( Storeini sensu lato ), larva, lateral view; J, Elleschodes sp. (Storeini sensu lato ), larva, head, anterior view; K, Haplonyx maximus Lea (Cryptoplini), pupa, ventral view; L, Daeneus annae (Ochyromerini), pupa, ventral view; M, Storeus albosignatus Blackburn (Storeini sensu stricto ), pupa, ventral view (G, H, and L after Oberprieler 1993, all others after May 1994). Scale bars 1 mm except 0.5 mm for B, D, F, H, and J. Curculioninae Latreille, 1802 315

laterotergal setae on tergites I – VII, sometimes also dorsally by a distance similar to or less than its basal on VIII; tergum IX with or without a pair of short width. Moreover, the Curculiones longirostres possess to long, more or less sharply pointed, contiguous or phanerognathous mouthparts, antennal scapes not widely separated posterior processes (urogomphi). passing the anterior margin of the eyes, the tibiae Abdomen lacking ventral setae (Burke 1968; Clark lacking spurs, and the body length seldom exceed- et al . 1978; Cussigh 1992; May 1993, 1994). ing 10 mm. Even though this concept lacked a clear defi nition (autapomorphic characters), it was an Phylogeny and Taxonomy. In its various current important fi rst step in gathering the various taxa of taxonomic circumscriptions, the subfamily Cur- “ fl ower weevils ” into a larger group. culioninae is a morphologically and biologically In the fi rst morphological phylogenetic analysis highly diverse group of weevils, without any clear of all higher weevil groups, Kuschel (1995) went concept yet of it representing a natural (monophy- further and formally subsumed all the taxa of letic) taxon either in its widest sense (Kuschel 1995) Thompson’ s Curculiones mediirostres and Curculiones or in a narrower one as adopted here. No attempt longirostres as tribes in a larger subfamily Curcu- has yet been made to subject a meaningful number lioninae. This concept, however, also included the of curculionine tribes to a modern phylogenetic Erirhininae, a group possessing the ancestral pedo- analysis with adequate taxon sampling, and also, tectal type of male genitalia and therefore treated as molecular analyses spanning all subfamilies of a separate family by Thompson (1992) and included Curculionidae (McKenna et al. 2009) so far have not in the subfamily Brachycerinae by Oberprieler et al. shown clear monophyly of and between tribes of (2007). However, even without the erirhinines, a Curculioninae. Considerably larger sets of taxa and subfamily Curculioninae in Kuschel’ s (1995) wide characters need to be analyzed to begin the under- sense is also not demarkated by any autapomorphic standing of proper tribal delineations and relation- characters. ships. As a result, the classifi cation of the subfamily In their catalogue of weevil genera, Alonso- and its tribes remains tentative and may change Zarazaga & Lyal (1999) adopted an eclectic system- signifi cantly in future. atic arrangement of curculionid subfamilies and The traditional concepts of most curculionine tribes, based in part on Thompson’ s (1992) classi- groups date back to Lacordaire’ s (1863, 1866) clas- fi cation and composing a subfamily Curculioninae sifi cation system, which was very comprehensive of 30 tribes. Although this catalogue was intended at its time but refl ects the poor contemporary essentially as a nomenclatural work, its classifi ca- knowledge of especially the taxa of the southern tion system has also become a standard taxonomic hemisphere. As a major revision of these concepts, reference. Anderson (2002) thus adopted Alonso- Morimoto (1962 b) published a tentative higher- Zarazaga & Lyal’ s (1999) concept of Curculioninae level phylogenetic reconstruction of weevils based in his treatment of the North American weevil mainly on differences in the shape of the metendo- genera, yet concluding that “ this subfamily [ … ] sternite and ovipositor. His scheme posits six main is now a large conglomerate of taxa of question- groupings within Curculioninae (i.e., Acalyptini , able relationships” . He reviewed a suite of possi- Anthonomini , Curculionini , Mecinini , Rham- bly distinctive features for the subfamily, i.e., apex phini, and Tychiini ), whereas two other tribes, of hind tibiae with small or no tooth (uncus); eyes Cionini and Smicronychini , were placed in a sepa- rounded; rostrum usually elongate and cylindri- rate subfamily Hyperinae. Being focused on the cal, distinctly longer in female than in male; anten- fauna of Japan, Morimoto’ s system did not, how- nae with scape at least reaching anterior margin of ever, consider the various curculionine groups the eye and inserted more basally in female than in more or less restricted to Africa, Australia, and male. However, none of these characters are pres- South America. ent in all tribes of Curculioninae, and all of them From a study of some key morphological char- occur also in other curculionid subfamilies. Indeed, acters, Thompson (1992) assembled many of the the Curculioninae are diagnosed in four different tribes here included in Curculioninae in an infor- parts of Anderson’ s (2002) key to the Nearctic cur- mal group named Curculiones longirostres , which culionid subfamilies. Despite the current absence spanned the former subfamilies Anthonominae, of unambiguous and unreserved diagnostic fea- Ceutorhynchinae, Cioninae, Curculioninae, Dere- tures, it is apparent that many curculionine tribes lominae, Gymnetrinae, Hyperinae, Rhamphinae, are more closely related to each other than to mem- Smicronychinae, Storeinae, Tychiinae, and Xiphas- bers of tribes placed in other subfamilies such as pidinae. He distinguished this group from the Cur- Conoderinae and Molytinae (see Oberprieler et al. culiones mediirostres (here subfamilies Molytinae, 2007), lending a measure of support to the present Cossoninae, and Conoderinae) by the tibiae being arrangement. imperfectly uncinate and from the Curculiones brevi- Molecular data and analyses have to date not pro- rostres (here subfamilies Entiminae and Cyclominae vided any resolution of this dilemma. Marvaldi et but also Eugnomini and Ulomascini ) by the head al. (2002) published a phylogeny of the superfamily being relatively small, subspherical, the eyes rela- Curculionoidea based on both molecular (18S ribo- tively large, ovate, forwardly placed, contiguous somal RNA) and morphological characters (ima- with the base of a slender rostrum and separated ginal and larval) but including representatives of 316 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler

only six curculionine tribes (Anthonomini , Cama- (1985) illustrated and keyed the larvae of 15 species rotini, Curculionini , Derelomini , Mecinini , Rham- of Curculio Linnaeus. More comparative and phylo- phini). The analysis of the combined data placed genetic studies of curculionine larvae are needed to these taxa on different branches of the cladogram elucidate the monophyly and relationships among and the molecular data alone did not resolve any tribes in the subfamily on larval characters (see also relationships among the “ higher ” Curculion idae. May 1993). More recent molecular analyses (Hundsdö rfer et al . Curculioninae are poorly represented in the fos- 2009; McKenna et al. 2009) based on more genes and sil record, most specimens having been described including also more curculionine tribes (11 and 13, from Eocene deposits at Florissant and Green River Q: Please respectively) did not fare much better, although the in the United States (e.g., Scudder 1890, 1893, check if the latter recovered a well-supported clade including Wickham 1911, 1912, 1917) and Havighorst in Ger- sentence has Curculioninae together with Molytinae, Conoder- many (Zherikhin 1995) and from Miocene deposits been correctly inae, Cossoninae, and Scolytinae. It is evident that at Shanwang in China (Zhang 1989) and the Simni- reworded: “although the analyses of relationships in Curculioninae and cel Valley in Romania (Marinescu & Proches 1999). latter recovered these other subfamilies of “ higher ” Curculionidae Most were placed in the extant genera Anthonomus , a well-sup- need to be based on much larger sets of taxa and Curculio ( “Balaninus ” ), and Tychius , but their true ported clade... characters. identities are generally unclear and all of them are The larvae of Curculioninae are similarly diffi - in need of critical assessment. A much older fos- cult to diagnose. Van Emden (1938) provided diag- sil, Arariperhinus monnei , was recently described noses of the larvae of Mecinini and Rhamphini and from the late Lower-Cretaceous Crato Formation their European genera and species. Scherf (1964), in the Araripe Basin, Brazil, and interpreted as in a comprehensive treatment of the morphol- belonging to Curculioninae (and tentatively to ogy and ecology of the larvae of central European the tribe Anthonomini ) (Santos et al. 2011), based weevils, distinguished the genera and species of on its strongly convex body, rounded eyes, slen- Curculioninae in a key based largely on their host- der rostrum, and absence of a prosternal canal and plants, developmental sites, and feeding pattern, tibial spurs. However, these characters do not dis- with morphological differences occasionally in tinguish the fossil from extant South American support but made no attempt to diagnose the lar- stenopelmine Brachycerinae such as Argentinorhyn- vae of higher (tribal) groups. May (1993, 1994) pro- chus Br é thes, Cyrtobagous Hustache and Onychylis vided diagnoses of and keys to the larvae of some LeConte, and it may equally well represent this Australo-Pacifi c tribes – Cryptoplini , Eugnomini , subfamily. Storeini (as Curculionini) – but in most cases, these Given the uncertain concept of a subfamily Cur- were based only on few representatives. In a key to culioninae, the taxon is here necessarily treated the South American larvae of Curculionoidea, Mar- in a pragmatic sense, largely following Alonso- valdi (2003) placed Curculioninae near Lixinae, Zarazaga & Lyal (1999), Alonso-Zarazaga (2002), from which they were distinguished by the reduc- Korotyaev (2002), Marvaldi & Lanteri (2005), Elgu- tion in overall size and the number of cephalic setae, eta & Marvaldi (2006), Oberprieler et al . (2007), in addition to a posteriorly emarginate head (not Oberprieler (2010), and Bouchard et al . (2011), who in all taxa) and often distinctive feeding styles (in considerably restricted its scope in comparison fl owers, fruits, seeds, etc.). According to van Emden with classifi cations proposed by earlier authors, (1938) and Marvaldi (2003), only larvae of the tribes such as Hoffmann (1954; as Calandrinae), Kuschel Mecinini, Rhamphini, and Anthonomini are read- (1995), and Marvaldi & Morrone (2000). Most of its ily diagnosed, the fi rst two by having abdominal tribes as considered here were assigned subfamily segments with only two (instead of three) dorsal status in the past (e.g., Morimoto 1962 b; Thomp- folds and expanded pedal lobes and the last by son 1992; Marvaldi et al . 2002), but very few of having four (instead of six) epipharyngeal sensilla them as currently composed (e.g., in the nomen- and by lacking the setal pair 2 on the frons. In their clatural catalogue of weevil genera by Alonso- treatment of Anthonomini, Ahmad & Burke (1972) Zarazaga & Lyal 1999) can be regarded as natural. presented the most complete study of larvae of any The treatment of the tribes below corrects some large tribe of Curculionoidea, identifying the fol- of the most glaring misplacements of genera and lowing diagnostic features: (1) most abdominal thus attempts to provide a more robust taxonomic segments with three dorsal folds, (2) frontal setae foundation of the various taxa for future studies, 2 absent, (3) anus subterminal. Burke (1968) and but the delimitations of essentially all the tribes Ahmad & Burke (1972) have described and illus- are in need of critical assessment. A number of gen- trated the larvae of 63 species and the pupae of 58 era remain of uncertain placement, although the species of Anthonomini, covering 12 and 15 gen- majority of the ones so listed by Alonso-Zarazaga era, respectively. Clark et al. (1978) discussed the & Lyal (1999) can be provisionally accommodated immatures of nearly 15 species of Tychius , conclud- in Storeini (q.v. ). ing that they seem to support some groupings pre- Acalyptini (Fig. 3.7.10.6 A and B ). This tribe viously postulated on adult morphology, but that includes a number of palm fl ower weevils, but its the phylogenetic signal was not strong. Gibson composition remains unclear despite some recent Curculioninae Latreille, 1802 317

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Fig. 3.7.10.6 Habitus, adults. A, Acalyptus carpini (Schoenherr) (Acalyptini); B, Eudelodes bicolor (Faust) (Acalyptini); C, Acentrus histrio (Schoenherr) ( Acentrusini); D, Anoplus roboris Suffrian ( Anoplini); E, Anthonomus rectirostris (Linnaeus) ( Anthonomini); F, Camarotus sp. ( Camarotini); G, Cionus olivieri Rosensch ö ld ( Cionini); H, Cerocranus extremus Kuschel ( Cranopoeini); I, Sigastus fuscodorsalis (Heller) ( Cryptoplini); J, Curculio glandium Marsham ( Curculionini); K, undescribed genus from Australia with horizontal mandibles, seven funicles, divaricate claws, and trochanteral setae (Curculionini); L, Derelomus piriformis (Hoffmann) ( Derelomini). Scale bars 1 mm. 318 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler studies. Kojima & Morimoto (2005) distinguished Derelomorphus , Meredoloides , Meredolus , Nodocnemus , the predominantly Australo-Asian genera of Parimera , Tithene , and several undescribed genera Acalyptini from the African Derelomini , based in Australia occur on palms (Arecaceae), which they crucially on a unique confi guration of the pygid- are indicated to pollinate (Essig 1973; Mogea 1978; ium of the males in the former (representing ter- Cock 1985; Henderson 1986), although the larvae gite VII, with tergite VIII hidden under it) and also and life cycles have not been studied in detail. on their non-carinate prothorax, two-segmented Acentrusini (Fig. 3.7.10.6 C ). Hoffmann (1958) labial palps, unarmed tibiae, simply truncated and Alonso-Zarazaga (2005) provided a diagnosis elytra, absence of wing folding and elytro-tergal of this monobasic tribe, the latter author regard- stridulation structures, and some genital differ- ing it to be related to Styphlini but without iden- ences. Most of these features are, however, homo- tifying specifi c shared characters. The distinctive plasious or plesiomorphic with respect to the New characters of the tribe are: body densely covered World lineages included in Derelomini by Franz with broad scales; funicles seven-segmented; (2006). Based on a morphological cladistic analy- eyes almost contiguous ventrally; frons as wide as sis of 115 taxa and 155 characters, Franz (2006) rostrum at base; prosternum with rostral canal; included Acalyptini as a subtribe in Derelomini femora unarmed; tibiae with small uncus; claws (but the name of the latter is the junior one; see simple, free, divaricate. Alonso-Zarazaga 2007). The analysis yielded a The larvae of Acentrus histrio (Fig. 3.7.10.6 C), the number of homoplasious but nevertheless unam- single representative of the tribe (Alonso-Zarazaga biguously optimized morphological features for 2005), tunnel in the roots and basal parts of the Acalyptini in such an expanded sense as well as stems of Glaucium (Papaveraceae), pupating inside for less inclusive clades incorporating the tradi- the tunnels or in the soil (Hoffmann 1958). tionally recognized acalyptine genera (Acalyptus Ancylocnemidini . Comprising the single genus Schoenherr, Amorphoidea Motschulsky, Nodocne- Ancylocnemis Marshall (7 species), this tribe is also mus Marshall, Parimera Faust), involving inter alia of uncertain affi nity and taxonomic status. Ancy- losses mostly of setae in mouthpart structures. locnemis occurs mainly in Africa, from South Africa However, the confi guration of the pygidium was to Ethiopia, but two species have been described not included as a character in the analysis, and the from India and at least one undescribed species antennal club is four-segmented in all acalyptine occurs in southern China (M. Alonso-Zarazaga, genera (rather than this condition representing a personal communication). Marshall (1920) origi- reversal in only Nodocnemus and Parimera ). There is nally described the genus in Anthonomini but therefore no strong evidence supporting a merger later transferred it to the molytine tribe Trachodini of Acalyptini and Derelomini into a single tribe (Marshall 1943, 1948). Such a relationship was, (Acalyptini sensu lato ). In its narrow sense (Kojima however, disputed by Voss (1962) and Zimmerman & Morimoto 2005), it currently includes 14 gen- (1964), the former proposing a separate tribe for it, era, Acalyptus (ca. 4 species), Amorphoidea (ca. 14), placed near Rhamphini . The character of laterally Derelomorphus Marshall (1), Epamoebus Blackburn angled ventrites 2 – 4 might support a relationship (4), Eudela Pascoe (1), Eudelodes Zimmerman (2– 3), to this and allied tribes such as Ochyromerini and Imera Pascoe (4), Meredoloides Kojima (1), Meredo- Tychiini , but the exact relationships of the genus lus Marshall (1), Niseida Pascoe (1), Nodocnemus (2), are as yet unclear. Its salient characters are the fol- Orsophagus Roelofs (2), Parimera (ca. 6), and Tithene lowing: body densely squamose, with tufts of elon- Pascoe (11), ranging from the Palaearctic to the gate scales on pronotum and odd elytral interstriae; Oriental and Australo-Pacifi c regions. A suitable rostrum narrow, subcylindrical, straight; funicles defi nition of the tribe in this sense and an over- seven- or (in subgenus Epimerogrypus Heller) six- view of the genera (except Epamoebus and Meredo- segmented, thus club four- or fi ve-segmented; loides ) were provided by Kojima & Morimoto eyes moderately large, frontally positioned, and (2005). However, a plethora of undescribed gen- dorsally almost contiguous; fore coxae contigu- era and species occurs at least in Australia, mostly ous or separate; prothorax without ocular lobes; associated with palm fl owers, and further study of mesoventrite steeply declivous (almost vertical); the group is necessary. middle coxae close together; femora dentate, Kojima & Morimoto (2005) summarized the hind femora often very strongly so, the fl attened information available on the biology of Acalyptini . tooth bearing a row of curved setae at its poste- The larvae of Acalyptus develop in fl ower catkins of rior edge; tibiae imperfectly uncinate, internally Salix (Salicaceae) and those of the closely related with median tooth and externally with apical and Orsophagus and Epamoebus in fl owers of Melia azeda- sometimes additional subbasal tooth; tarsal claws rach (Meliaceae) and Acacia (Fabaceae), respectively. divaricate, slightly to strongly appendiculate; ven- Amorphoidea lives in fl owers of Malvaceae such as trites 2 – 4 laterally usually slightly angled; tegmen Hibiscus, Thespesia, and Gossypium (cotton), several dorsally membranous, without parameroid lobes. species becoming pests on the latter, and also, Eude- Some seemingly related undescribed small South lodes breeds in fl owers of Hibiscus tiliaceus along the African species agree with Ancylocnemis in most of east coast of Australia and the Torres Strait islands. these characters but have large, round erect setae, Curculioninae Latreille, 1802 319 the fore coxae broadly separated by a pectoral canal Combretaceae (Terminalia ), Geraniaceae (Geranium/ ending in a shallow receptacle on the metaventrite, Pelargonium), and Penaeaceae ( Penaea), but again, the femora adentate and the tibiae bearing a two- nothing is recorded about its larvae and their hosts pronged uncus. (Oberprieler 1993). Ancylocnemis appears associated with Cappara- Anthonomini (Fig. 3.7.10.6 E ). As currently ceae, Marshall (1943) recording A. sternalis Marshall composed (Clark 2010), this tribe includes 43 gen- from Capparis sepiaria (Wild Caper Bush) in India era and 826 species, the majority of species (558, of and some Ancylocnemis and the undescribed species which 395 are Neotropical, 121 Nearctic, and 58 with a pectoral canal living on Boscia foetida in South Palaearctic) classifi ed in the large and widespread Africa (R. Oberprieler, personal observation). The genus Anthonomus Germar. Almost all other gen- larvae probably develop in the fruits (Zimmerman era comprise fewer than ten species, except for 1964). Atractomerus Duponchel & Chevrolat (45 species), Anoplini (Fig. 3.7.10.6 D). The concept of this Bradybatus Germar (20), Huaca Clark (26), and Pseu- tribe is unclear. In the catalogue of Alonso-Zaraz- danthonomus Dietz (35). The classifi cation of the aga & Lyal (1999), it includes only two genera, tribe remains partially unresolved, as the constitu- the Palaearctic Anoplus Germar (5 species) and the ent genera vary signifi cantly in their morphologi- Madagascan Paranoplus Hustache (1), characterized cal characters and the composition of the tribe has by the absence of the onychium of the tarsi. How- changed considerably over the past 20 years. Dieck- ever, this loss has occurred independently numer- mann (1968) last revised the western Palaearctic ous times in Curculioninae and other subfamilies genera, whereas Clark (1988, 1989, 1990) clarifi ed and cannot be used as a tribal characterization. the phylogenetic relationships among several gen- Zherikhin & Egorov (1991) and Kojima & Morim- era of Neotropical anthonomines and provided a oto (2000) also placed the eastern Palaearctic genus wealth of taxonomic information on known and Sphinxis Roelofs (ca. 15) in Anoplini , and Kojima & new species. Kojima & Idris (2004) diagnosed the Morimoto (2002) added the mainly African Tham- tribe as follows: eyes more or less convex; rostrum nobius Schoenherr (ca. 10), which they recorded cylindrical; procoxal cavities generally contiguous; from the Oriental region but which is evidently a tibiae usually uncinate (at least on fore and middle composite genus (Oberprieler 1993). Several other legs); tarsal claws not widely divergent, generally Afrotropical genera, specifi cally the Madagascan sharply bifi d, and with one seta each; posterior Cionesthes Fairmaire, Lepidoops Hustache, Parendoe- margin of abdominal ventrites 2 – 4 straight, not lat- opsis Hustache, and Semicardius Hustache but also erally angled; aedeagus narrowed or emarginate at the continental Acalyptops Hartmann, Neosphinc- base in lateral aspect and with apodemes articulat- tocraerus Hustache and Pansmicrus Schoenherr, ing at ventral surface; sternite VIII of male paired, appear to belong to this tribe as well (Kojima & devoid of setae along caudal margin; sternite IX of Morimoto 2002), and its proper composition and male bilobed, with apodeme more or less curved defi nition are in need of study. Alonso-Zarazaga and oblique to axis of aedeagus; spermatheca with (2002) placed the tribe in Molytinae, without com- lateral lobe indefi nite, spermathecal gland long, ment or supporting evidence. The detection in Ano- linear. plus, Sphinxis, and Thamnobius of an elytro-tergal Anthonomini utilize more than 40 plant fami- stridulation system 2 (Kojima & Morimoto 2000, lies as hosts (H. Burke, personal communication), 2002), otherwise recorded only in a few Molyt- including both herbaceous groups such as Astera- inae and Cossoninae (Lyal & King 1996), may lend ceae, Euphorbiaceae, Malvaceae, and Solanaceae support to such a placement, but the feature was and woody ones such as Rosaceae, Sapindaceae, and interpreted as independently evolved in Acalyptini Ulmaceae (Scherf 1964; Dieckmann 1968; Burke by Kojima & Morimoto (2000) and has recently 1976; Anderson 1993; Clark & Burke 1996). Indi- also been reported to occur in Derelomini (Kojima vidual species, however, tend to have much more & Morimoto 2005). Elytro-tergal stridulation restricted host ranges, spanning a small number of systems in general are in need of wider study in species in one plant genus or a few closely related Curculioninae. genera (e.g., Scherf 1964; Burke 1976; Burke et al. Little is known about the biology of Anoplini. 1986). Nearly all Palaearctic and several North The larvae of Anoplus mine in leaves of Alnus and American species of Anthonomus live on Rosaceae, Betula (Betulaceae); on completion of their devel- causing serious damage to fruit trees in the gen- opment, they cut out pieces of leaf to form a pupal era Cerasus and Prunus and to shrubs of the genera covering, in which they drop to the ground for Fragaria (strawberries) and Rubus (blackberries, pupation (Scherf 1964). Zherikhin & Egorov (1991) dewberries) (Burke 1976). The North American recorded Sphinxis to be associated with Polygonum cotton boll weevil, A. grandis Boheman, is a serious (Polygonaceae), but Kojima & Morimoto (2000) pest of cotton plants (e.g., Cross et al . 1975) and one doubted this because Sphinxis generally inhab- of the most extensively studied species of weevils. its broad-leaved trees of the families Elaeocar- Scataglini et al. (2006) inferred that South American paceae, Lauraceae, and Sapindaceae. Thamnobius in populations of A. grandis represent multiple ancient Africa has been collected on Anacardiaceae ( Rhus ), and more recent invasions from the United States. 320 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler

In contrast to such pest species, the South Ameri- it also pupates in a spherical cocoon protected can Anthonomus santacruzi Hustache, which devel- in a pupation chamber. A Costa Rican species of ops in fl ower buds of Solanum mauritianum , has been Camptocheirus induces galls on Cinnamomum cinna- identifi ed as the most promising biocontrol agent momifolium (Lauraceae), where it is parasitized by a of this seriously invasive weed in South Africa braconid wasp (Fortier & Nishida 2004). (Olckers 2009). It is noteworthy that two Palaearc- Ceratopodini . Another small Neotropical tribe tic species of Anthonomus, A. phyllocola (Herbst) and of uncertain status and affi nity, the Ceratopodini A. pinivorax Silfverberg, and also Brachonyx indigena presently include only three genera, Aetiomerus Schoenherr and Macrobrachonyx gounellei Pic live Pascoe (1 species), Catiline Champion (3), and Cera- on conifers of the genera Picea and Pinus (Pinaceae) topus Schoenherr (ca. 20). The Madagascan Anthom- (Scherf 1964; Dieckmann 1968; Alonso-Zarazaga, elus Hustache (8) and the New Guinean Stelechodes personal communication). Larvae develop mainly Faust (1), as listed in the tribe by Alonso-Zarazaga in fl ower buds or fl ower heads and also in leaf buds, & Lyal (1999), almost certainly do not belong in it, fruits, and seed capsules, in which they also pupate although their proper placements are in need of (Burke 1976), but some species live as inquilines in assessment. Marvaldi & Lanteri (2005) regarded galls produced by fungi, nematodes, and several Ceratopodini as closely related to Camarotini , insect orders, sometimes in obligate relationships Piazorhinini , and Prionobrachiini , with which (see Gates & Burke 1972). In Africa, the genera cur- they share the large, triangular, often serrate tooth rently placed in Anthonomini live on Acacia (Faba- of the fore femora. A comprehensive revision of all ceae) and Ziziphus (Rhamnaceae) (Oberprieler 1993). these groups is needed. Camarotini (Fig. 3.7.10.6 F). This small Neo- Pakaluk & Carlow (1994) reported the larvae of tropical tribe currently includes six genera (for the Ceratopus to develop in fruits of Ficus and Helicostylus Australian Meripherinus Lea see Eugnomini ) and is (Moraceae). generally divided into two subtribes, Camarotina Cionini (Fig. 3.7.10.6 G). This tribe currently and Prionomerina (Alonso-Zarazaga & Lyal 1999). comprises seven genera occurring predominantly However, the prionomerines previously included in the Palaearctic region. The largest genus is also Piazorhinus Schoenherr, often split off into its Cionus Clairville (ca. 100 species), which is also own higher taxon ( Piazorhinina ) (e.g., O ’ Brien & represented in southern Africa, whereas the Wibmer 1982), whereas Kuschel (in Wibmer & other genera include fewer than 10 species (Cleo- O ’ Brien 1986) proposed a larger concept of Cama- pus Dejean, Stereonychus Suffrian) or are monoba- rotini that included not only Prionomerina and sic ( Cionellus Reitter, Nanomicrophyes Pic, Patialus Piazorhinina but also Prionobrachiina . Lyal & King Pajni, Kumar & Rose, Stereonychidius Morimoto). (1996) similarly included both the latter taxa in Based on the Palaearctic species, Caldara & Koro- Prionomerinae (thus in Camarotini), and it appears tyaev (2002) delimited the tribe as a monophyletic that all these taxa may represent a single lineage, group identifi ed by eight synapomorphies, i.e., characterized by mostly strongly infl ated fore fem- frons between eyes narrower than half the width of ora carrying a large, serrate tooth on the anterior the rostrum and narrower posteriorly than anteri- side, a short broad rostrum with short, compact orly; funicular segment 2 as long as or slightly lon- antennae, and leaf-mining larvae. Reichardt (1971) ger than 1; median process of mesoventrite at least revised the genus Camarotus Germar (40 species), half as wide as a coxa; mucro on all tibiae absent broad, fl at weevils characterized by having weakly in female; ventrites 1 and 2 much longer (at least geniculate antennae, widely separated fore coxae, 2.6 times) than 3 and 4; spermatheca with body strongly infl ated fore femora with a large, serrate markedly globose and ramus very short. Caldara tooth and appendiculate claws. Reichardt (1971) & Korotyaev (2002) also analyzed the phylogenetic recognized three species groups in Camarotus , based relationships of the genera, based mainly on the on differences in the structure of the elytra. The fi ve presence of a rostral canal and tibial mucrones genera of Prionomerina, Camptocheirus Lacordaire and on features of the mesoventrite. A revision of (6), Ectyrsus Pascoe (3), Odontopus Say (44), Pristimerus the entire tribe is in progress, preliminary results Schoenherr (2), and Themeropis Pascoe (9), have not indicating that most of the characters currently been studied recently, Ectyrsus also having been used to distinguish the seven genera are in need of moved from Camarotini to Anthonomini by Wib- reassessment with respect to the extra Palaearctic mer & O’ Brien (1986). species (Caldara & Kos ˇ t ´á l, in preparation). The larvae of Camarotus mine in parenchymous The larvae of Cionini are ectophagous, living tissue of leaves of Melastomaceae and other dicoty- externally on their host-plant or hidden between ledonous angiosperms (Reichardt 1971, based on adjoining leaves and feeding on fl owers and leaves. Bondar 1945) and also the larvae of several Priono- Although legless, they move nimbly on plant sur- merina are leaf miners on various plant families, faces because they are covered with viscid mucus, e.g., Odontopus on Annonaceae, Hamamelidaceae, which is secreted by a retractile tubular organ on Lauraceae, and Meliaceae (Anderson 1993). Rosado- the last segment. On completion of their develop- Neto & Lima (2002) recorded the larva of Odontopus ment, the larvae secrete a substance containing brevirostris (Hustache) to feed on the parenchyma chitinous microfi bers from their midgut (Tris- between the epidermal layers of the leaf, where tram 1978), with which they construct a spherical Curculioninae Latreille, 1802 321

translucent cocoon generally on the underside of onychium being narrow and the claws simple, fi ne, leaves or branches for pupation. Cionus and Cleopus subparallel (Menechirus ) to the onychium short and live mainly on Scrophulariaceae (but see McNeill et largely withdrawn into the lobes of tarsomere 3 al. 2005), but some species of Cionus and also Stere- (Sigastus ) to the claws being single, rarely bifi d at onychus occur on Oleaceae and Cionellus on Plantag- apex (Haplonyx ) to the onychium being minute and inaceae (Hoffmann 1958; Scherf 1964; Gumovsky hidden in the narrow cleft of tarsomere 3 (Crypto- 1997; Caldara & Korotyaev 2002). The only spe- plus ) to being completely absent (Zeopus ). Other cies of the Indian Patialus , P. tecomella Pajni, Kumar unusual features of Cryptoplini are the complete & Rose, lives on Marwar Teak, Tecomella undulata undifferentiation of the proventriculus, lacking (Bignoniaceae), and may seriously damage this eco- the normal plates and paired brushes found in nomically important tree (Rose & Sidhu 1994; Gaur most higher curculionids and the presence of a & Ahmed 1998). dorsal plate (tectum) on the penis. An undifferen- Cranopoeini (Fig. 3.7.10.6 H ). This tribe was tiated proventriculus is a feature of more primitive only recently established by Kuschel (2009) for the weevil groups (Calder 1989), and the only other Cranopoeus complex of genera that he had earlier Australian genera of Curculion idae so far known recognized in Molytinae (Kuschel 1987). The pre- for this condition are Strongylorhinus Schoenherr viously known genera, Cranopoeus Marshall (6 spe- and Atelicus , formerly classifi ed in Diabathrariini cies), Cratoscelocis Lea (1), Fergusoniella Zimmerman (Oberprieler et al. 2007; Oberprieler 2010). How- (1), Spanochelus Marshall (4), and Swezeyella Zim- ever, a weakly developed proventriculus occurs in merman (1), were included in Cryptoplini in the some other curculionid genera as well (see Calder catalogue of Alonso-Zarazaga & Lyal (1999), but 1898). A dorsal plate on the penis (the pedotectal Kuschel (2009) detailed a range of character differ- type of adeagus) is also characteristic of more prim- ences between the two groups and added another itive weevils, in Curculionidae, occurring only in six genera to the new tribe: Blastobius Kuschel (1), the subfamily Brachycerinae. The dorsal plate of Cerocranus Kuschel (1), Cranoides Kuschel (2), Docolens Cryptoplini is, however, basally broadly fused to Kuschel (1), Enneaeus Kuschel (1), and Onychomerus the pedon and the temones show no sign of a dorsal Kuschel (1). The Cranopoeini are most diverse in (tectal) arm at their bases, as is the condition in the the Pacifi c region (8 of the 11 genera, 6 only in New typical pedotectal aedeagus. The phylogenetic sig- Caledonia) but also present in Australia ( Cranopoeus, nifi cance of this character in Cryptoplini remains Cratoscelocis, Enneaeus, and Fergusoniella ) and New to be investigated. Other characters of the tribe are Guinea (undescribed). The tribe was well charac- the following: body densely squamose; rostrum terized by Kuschel (2009), the most conspicuous stout and straight, sometimes apically fl attened; feature being the waxy secretions of the thorax and prementum mostly transverse; maxillary and elytra that harden to form solid structures, which labial palps three-segmented; scrobes apically not reach an extreme form in the huge, curved, hollow visible from above; funicles seven- or (in Cryptoplus thoracic “ horn ” or “ helmet ” of Cerocranus extremus and Zeopus ) six-segmented; clubs short, four- or Kuschel (Fig. 3.7.10.6 H). fi ve-segmented; fore coxae contiguous or narrowly Nothing precise is known about the biology of separate, middle coxae closer together than hind Cranopoeini , but Kuschel (2009) recorded Eucalyp- ones; all femora dentate; tibiae two-pronged, with tus and Syzygium (Myrtaceae), Glochidion (Euphor- inner uncus and praemucro above it; endophallus biaceae), Nothofagus (Nothofagaceae), Sloanea sometimes with large basal sclerite; gonocoxites (Elaeocarpaceae), and Lythraceae (as Sonneratia- without styli; base of spermathecal gland funnel- ceae) as probable hosts. Enneaeus bituberculatus (Lea) shaped sclerotized (Kuschel 2009). in Australia is found on Eucalyptus and Melaleuca . Cryptoplini are mostly associated with Myrta- Cryptoplini (Fig. 3.7.10.6 I). This small Aus- ceae. The larvae of Haplonyx, and Cryptoplus develop tralo-New Guinean tribe comprises fi ve genera, but in fl ower and fruit buds of Eucalyptus, Melaleuca , its exact composition and distribution need fur- and Leptospermum , the adult female generally sev- ther study. The genera Cryptoplus Erichson (ca. 30 ering the stem of a fl owering or fruiting terminal species), Haplonyx Schö nherr (ca. 64), Zeopus Pascoe so that it falls to the ground, where the larvae feed (1), and Sigastus Pascoe (2 – 3) are confi ned to Austra- and pupate in the wilting buds. Some species of lia, whereas Menechirus Hartmann (1), a close rela- Haplonyx have been reared from fl eshy and woody tive of Sigastus , occurs from northeastern Australia galls on Eucalyptus and Leptospermum , in which to New Guinea and Seram. The other genera listed they may develop both as incitants and inquilines. in the tribe by Zimmerman (1994, as Haplonych- Nothing is recorded of the biology of Zeopus , but ini) and Alonso-Zarazaga & Lyal (1999) belong to it probably also lives on Eucalyptus or other Myrta- a different tribe, the Cranopoeini (q.v. ), except for ceae. The larvae of Sigastus develop in fl owers and Brendamaya Alonso-Zarazaga & Lyal, whose name fruits of Syzygium but have also been reared from is a synonym of Atelicus Waterhouse (see Platyurus woody homopteran galls on Eucalyptus and Alloca- in Zimmerman 1994), a genus tentatively placed suarina (Casuarinaceae) (Juniper & Britton 2010), in Cyclominae: Aterpini by Oberprieler (2010). whereas an undescribed species has become a pest The tarsi of Cryptoplini characteristically have the in fruits (nuts) of planted Macadamia (Proteaceae) onychium and claws reduced, ranging from the in northern Australia (Fay et al . 2001). Sigastus 322 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler fuscodorsalis (Heller) (Fig. 3.7.10.6 I), originally Curculio and allies. Similar vertically articulating described in Menechirus , was imported in fruits of mandibles occur in Belidae: Oxycoryninae ( Oxyco- Syzygium suborbiculare from Darwin in Australia to rynus Chevrolat, Parallocorynus Voss, and Rhopalotria the Philippines (Heller 1921). Menechirus has been Chevrolat) and in Brentidae: Antliarhinini (females reared from Syzygium fruits but also collected from of the seed-feeding Antliarhinus zamiae (Thunberg); Rhodomyrtus macrocarpa in Australia. mandibles articulating obliquely in other Antlia- Curculionini (Fig. 3.7.10.6 J and K ). This cosmo- rhinus species) and appear to be convergent adapta- politan tribe contains approximately 15 genera, tions for drilling very fi ne oviposition holes. Other although a number of others may belong in it as characters of Curculionini include a long to very well. The almost cosmopolitan genus Curculio Lin- long, slender rostrum (especially in the female), naeus (ca. 500 species) is the largest, several smaller seven-segmented funicles (six-segmented in Erga- ones – Archarius Gistel (ca. 10), Carponinus Heller (ca. nia and Pimelata), ten-striate elytra without stridu- 50), Ergania Pascoe (17), Indocurculio Pajni, Singh & latory fi le and mostly narrowly impressed around Gandhi (10), Koreoculio Kwon & Lee (2), Labaninus the scutellum, often large and ascending mesepim- Morimoto (14), Pagumia Kwon & Lee (1), Pimelata era, contiguous fore coxae, dentate femora, broad Pascoe (1), Pseudoculio Pelsue & O’ Brien (7), and Shi- fl at tarsi, divaricate and appendiculate claws (sim- gizo Morimoto (14) – occurring in the Palaearctic ple in Timola ; parallel but not connate in Ergania, and Oriental regions. In the east, the distribution Pimelata , and Pycnochirus ), often exposed pygidium of the tribe extends southward to New Guinea and (especially in male), and ventrite 5 in male generally Australia, from where approximately 16 species of with median to apical depression fl anked by tufts Curculio are known as well as Pycnochirus Berg (ca. or rows of dense, long setae. Ergania, Pimelata, Pycno- 2), a genus classifi ed in Trigonocolini by Alonso- chirus, and some other genera (Fig. 3.7.10.6 K) also Zarazaga & Lyal (1999) but agreeing with Curcu- retain a strong trochanteral seta and appear to be lionini in most characters, especially the structure generally less derived than the other genera. After of its tarsi, venter, and pygidium. It was originally Heller (1925, 1927), the tribe is generally divided described from New Guinea but redescribed from into three subtribes, Curculionina, Pseudobalani- Australia as Balanerhinus Lea (Heller 1925; Rhein- nina, and Timolina, but Pelsue & O ’ Brien (2011) heimer 1994). Several undescribed genera occur in recently proposed three additional ones, Archariina Australia (Fig. 3.7.10.6 K) and New Guinea. Three for Archarius , Koreoculio , and Pagumia , Erganiina for genera are known from the Afrotropical region, Ergania and Pimelata, and Labaninina for Labaninus, Timola Pascoe (ca. 30) from continental Africa and Pseudoculio , and Shigizo . However, none of the dis- Aviranus Fairmaire (ca. 20) and Pseudobalaninus tinguishing characters of these subtribes appear to Faust (ca. 40) from Madagascar, but some other constitute suitable synapomorphies for defi ning Madagascan genera may belong in the tribe as well. them as monophyletic groups, and the entire tribe Pelsue & O ’ Brien (2011) recently described a genus is in need of revision and phylogenetic analysis to Megaoculus based on two South American species, determine its proper delineation and the relation- but it does not convincingly fi t into Curculionini ships between the genera. and may represent a different tribe. Gibson (1969, The biology of many species of Curculionini is 1977) revised the North and Central American spe- relatively well known. Their host-plants mostly cies of Curculio, whereas Morimoto (1960 a, b, 1981) belong to the families Betulaceae, Fagaceae, Jug- treated the Japanese and Pelsue & Zhang (2000 a, landaceae, Moraceae, and Salicaceae (Hoffmann b, 2002, 2003) treated the Chinese ones. An explor- 1954; Scherf 1964; Gibson 1969, 1977; Rasplus et atory molecular phylogeny of 22 species of Curculio al . 2003). The larvae generally develop in flower (Hughes & Vogler 2004) supported two previously buds or fruits and pupate in the soil, but some, recognized European species groups but remained especially in the genus Archarius , incite galls or live inconclusive with regards to the monophyly of the as inquilines in galls produced by Hymeno ptera American groups. Pelsue (2004, 2005, 2007, 2009) (Hoffmann 1954; Scherf 1964; Cussigh 1992). revised the Oriental genera Indocurculio, Labaninus , The pecan weevil, Curculio caryae Horn, is the and Shigizo . most extensively studied and economically by far Curculionini are traditionally characterized the most important species of the tribe (see Neel by their smoothly triangular, vertically moving 1985). In Japan, the rostrum length of Curculio mandibles, but this character only occurs in the camelliae Roelofs is closely linked to the pericarp derived genera such as Curculio , Carponinus , Indocur- thickness of its host, Camellia japonica (Theaceae), culio , Koreoculio , Labaninus , and Shigizo . In Ergania , across multiple varying populations, yielding a Pimelata , and some Timola species, the mandibles pattern of adaptation suggestive of a co-evolu- articulate in a more or less oblique plane, the sock- tionary arms race (Toju & Sota 2006). In Austra- ets positioned lateroventrally, and a gradual tran- lia, most species of Curculio appear to develop in sition in the plane of the mandibles can be traced figs (Moraceae) (Zimmerman 1994), but Howden from a horizontal position as in Pycnochirus , and (1995) recorded C. niveopictus Lea ovipositing in some undescribed forms (Fig. 3.7.10.6 K) via the ripe fruits of Syzygium tiernayanum (Myrtaceae), oblique one as in Ergania to a fully ventral one in whereas Pycnochirus fuscovarius (Faust) has been Curculioninae Latreille, 1802 323 collected from fruits of Alectryon coriaceus (Sap- female detaches a staminode with her mandibles, indaceae) (Rheinheimer 1989). In Africa, Curcu- falls to the ground with it, transports it to a site in lio also develops mostly in figs (Marshall 1907; leafl itter using her frontal spine as a point of resis- Perrin 1992), whereas Timola is associated with tance against the substrate and then oviposits in it. Ebenaceae, T. posticata Marshall developing in Meanwhile, males associate with the moving sta- fruits of Diospyros lycioides, and other species liv- minode and fi ght each other with their forelegs, ing on, e.g., D. austro-africana and Euclea species executing fast blows until their tibial teeth cling (R. Oberprieler, personal observation). and dislodge competitors from the staminode. A Derelomini (Fig. 3.7.10.6 L). In a comprehensive phylogeny of Staminodeus hypothesizes that female cladistic analysis of more than 30 genera and 90 transporting behavior evolved before male fi ghting species in the in-group, Franz (2006) diagnosed the behavior (Franz 2003). Other Neotropical lineages Derelomini as follows: segments 1 and 2 of maxil- with potentially more than 100 species remain lary palps lacking mediolateral setae; tibial apices almost entirely undescribed yet constitute the pri- unarmed; adults feeding and reproducing only on mary pollinators of Cyclanthaceae (Franz 2007 a). angiosperm fl owers and infl orescences, typically Pollinating members of the related phyllotrogine acting as pollinators; and predominantly associated genus Cyclanthura Franz, in turn, have shifted onto with infl orescences of Arecaceae. Franz (2006) pro- species of Araceae, in particular Anthurium (Franz posed fi ve subtribal concepts to accommodate the 2003, 2007 b). major lineages in this morphologically and biologi- Diabathrariini (Fig. 3.7.10.7 A ). As limited by cally heterogeneous tribe, including the predomi- Oberprieler et al . (2007) and Oberprieler (2010), this nantly Old World (Asian) Acalyptina, which were small group only includes three African genera, nested within Derelomini (though see Acalyptini Aphanonyx Marshall (2 species) and Diabathrarius above). Members of Anthonomini , Ellescini , and Schoenherr (4) in southern Africa and Madagascar Trypetidini were identifi ed as the most immedi- and Onychogymnus Quedenfeldt (ca. 14) in central ate out-group, the last taxon, however, now con- Africa and Madagascar. All have entirely lost the sidered part of the molytine tribe Petalochilini (see onychia, a condition rare among African weevils Chapter xx.x). Kojima & Morimoto (2005) identi- but more common in Australia. The weevils are fi ed the unusual elytro-tergal stridulation system stocky with short, stout legs; densely covered in 2 (Lyal & King 1996) occurring in typical Derelo- broad, fl uted, overlapping scales; the rostrum short mini (the Old World Deleromus Schoenherr and and broad, sometimes apically strongly fl ared, fi t- Elaeidobius Kuschel), whereas the New World genus ting into a shallow depression or deep excavation Anchylorhynchus Schoenherr has the more common (in Aphanonyx) of the prosternum; the scrobes nar- system 1 (Lyal & King 1996). row, deep, and curving onto the venter of the head Derelomini , generally known as “ palm fl ower before the eyes; the funicles seven-segmented and weevils ” or “ derelomine fl ower weevils ” (Franz the clubs short; the maxillary and labial palps large, 2006), are primarily associated with the reproduc- three-segmented; the femora unarmed; the tibiae tive organs of monocotyledons belonging to the with a strong uncus and smaller praemucro above families Arecaceae and Cyclanthaceae (Henderson it; the proventriculus strongly developed; the ely- 1986; Franz & Valente 2005). According to Franz tra without stridulatory fi le; the pointed aedeagus & Valente (2005), certain lineages inhabiting tem- with two pairs of apical setae and the tegmen with perate regions or higher-elevation habitats have long parameres. The group is in need of revision, secondarily shifted onto dicotyledons, includ- and its taxonomic affi nities are unclear. ing members of the Acalyptina (Dieckmann 1988; Little is known about the biology of Diabath- Kojima & Morimoto 2005) and the genus Phyllotrox rariini, but Diabathrarius variegatus Boheman has Schoenherr in the Phyllotrogina. Larvae vary con- been reared from seedpods of Schotia afra in South siderably in life habits because they may be detri- Africa (S. Neser, personal observation), and Apha- tivorous or herbivorous, often feeding on parts of nonyx setulosus Marshall is there regularly collected the fl oral structures that are dispensable for the on Pterocarpus rotundifolius (R. Oberprieler, personal hosts (e.g., Celetes Schoenherr; Valente 2005) but observation) and may develop in its fruits as well sometimes destroying the seeds (e.g., Systenote- (both hosts Fabaceae). lus Anderson & G ó mez 1997). In certain lineages, morphological adaptations and reproductive strat- Ellescini (Fig. 3.7.10.7 B and C). Generally divided egies are closely correlated with those of their host- into subtribes Ellescina and Dorytomina, this tribe plants (Franz 2003 a, 2004). Notable derelomine was included in Tychiini (as a subfamily) by Clark lineages include the Staminodeina, in particular et al. (1977) based on the latero-posteriorly angled Staminodeus Franz, whose seven species are asso- ventrites and the direction of the pronotal scales. ciated with the staminodes of the infl orescences However, its composition and status are unclear. of cyclanths (Franz 2003 a). The males have a row The genera Minyrus Schoenherr and Ontoctetorus of 6– 10 teeth along the margin of the fore tibiae, Faust, listed in Ellescina by Alonso-Zarazaga & Lyal whereas the females have a small, curved spine (1999), are now considered to belong in Anthono- on the frons. Upon arriving at an infl orescence, a mini (Clark 2010), the Australian Sellechus Lea is 324 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler

Q: Figure has been resized - please confi rm.

Fig. 3.7.10.7 Habitus, adults. A, Aphanonyx latesetosus (Hustache) ( Diabathrariini); B, Dorytomus longimanus (Forster) ( Ellescini); C, Ellescus bipunctatus (Linnaeus) (Ellescini); D, Ludovix fasciatus (Gyllenhal) (Erodiscini); E, Ita chavanoni Meregalli & Borovec (Itini); F, Gymnetron beccabungae (Linnaeus) ( Mecinini); G, Mecinus janthinus Germar (Mecinini); H, Galloisia infl ata Hustache (Microstylini); I, Microstylus rufus Schoenherr (Microstylini); J, Nerthops guttatus Schoenherr (Nerthopini); K, Acallopistus sp. (Nerthopini); L, Teinomphasus robustus Oberprieler (Ochyromerini). Scale bars 1 mm. Curculioninae Latreille, 1802 325 currently best classifi ed in Storeini ( q.v.), and the in weevils overall) by possessing an elytro-femoral placement of the Asian Bathrorygma Marshall and stridulation system, consisting of a row of lateral Minyrophilus Voss in Ellescini is questionable. No elytral tubercles and a sharp keel on the inner face genera of Ellescina have been recently studied, but of the hind femora. many species of the Holarctic genus Ellescus Dejean Vanin (1986) and Anderson (1999) summa- are well known. The species of Dorytomus Germar rized the available information on the biology in North America, Japan, and Siberia were taxo- and host associations of the tribe. Adults are often nomically revised by O’ Brien (1970), Morimoto & found on infl orescences, particularly of aquatic or Enda (1962) and Korotyaev et al. (1976, 1996), and semi-aquatic plants, but the larvae appear to tun- Colonnelli (2003), respectively, synonymized all nel in branches of various plants. Ludovix fasciatus the subgenera as listed by Alonso-Zarazaga & Lyal (Gyllenhal) (Fig. 3.7.10.7 D) is seemingly unique (1999) with Dorytomus because they were shown to among weevils for being predacious, both adults be polyphyletic. and larvae feeding on eggs of the grasshopper Ellescini , in particular Dorytomus (ca. 100 species) genus Cornops (Acrididae) deposited inside stems and Ellescus (7), are generally associated with Populus of water hyacinth, Eichhornia crassipes (Pontederia- and Salix (Salicaceae) (Hoffmann 1958; Morimoto ceae) (Zwö lfer & Bennett 1969). Judging from the & Enda 1962; Scherf 1964; O’ Brien 1970; Dieck- even more extremely elongated rostrum of some mann 1988; Fjellberg & B ö cher 2006). The biology other species of Erodiscini, Vanin (1986) speculated of some Palaearctic species of these genera is well that entomophagy may be more common in this known. The larvae develop in the axis of mainly tribe. female catkins and penetrate the ovaries and pollen Eugnomini . In the catalogue of weevil genera masses, which causes the catkins to atrophy early (Alonso-Zarazaga & Lyal 1999), this tribe contains in their development and drop to the ground, the 23 genera, mainly from Australia and New Zealand larvae pupating in the soil (Hoffmann 1958; Scherf but also occurring in New Caledonia and South to 1964). The larvae of Dorytomus appear earlier in the Central America. Its precise composition and dis- season in Salix catkins than those of Ellescus (Scherf tribution, however, remain in contention. Kojima 1964). & Morimoto (1993) transferred the Oriental Api- Erodiscini (Fig. 3.7.10.7 D). This small Neo- onodes Marshall from Anthonomini to Eugnomini , tropical tribe was comprehensively revised by but Alonso-Zarazaga & Lyal (1999) placed it back Vanin (1986), who recognized 8 genera and 98 in Anthonomini (as a synonym of Pseudopoophagus species in it, and proposed a genus-level phylog- Voss). There are, however, also other Eugnomini in eny with two principal clades (exluding Sicoderus Indochina, similar to the undescribed Australian Vanin). Anderson (1999) added three further spe- genus pictured by Zimmerman (1992, pl. 511) (H. cies to Sicoderus , which comprises more than half Kojima, personal communication). Alonso-Zaraz- (59) of all species of the tribe. The other genera aga & Lyal (2002) also referred the New Zealand are much smaller, Ecnomorhinus Vanin (1), Erodis- Oropterus White to Eugnomini, whereas Leschen cus Schoenherr (5), Hammatostylus Champion (9), et al . (2003) listed this genus in Storeini and two Lancearius Vanin (2), Ludovix Laporte (2), Pimelero- other New Zealand genera in Eugnomini, Colabo- dius Vanin (12), and Prosicoderus Vanin (11). The telus Broun and Philacta Broun, which were placed Indonesian Pseuderodiscus Heller, listed in Erodis- in Erirhininae by Alonso-Zarazaga & Lyal (1999). cini by Alonso-Zarazaga & Lyal (1999), belongs Franz (2006) transferred the American genus to the molytine tribe Trachodini (H. Kojima, per- Pedetinus Faust from Derelomini to Eugnomini sonal communication). Erodiscini appear closely and presented four putative synapomorphies and related to Otidocephalini (Vanin 1986; Marvaldi two additional diagnostic features for the three & Lanteri 2005), and the two groups are some- American genera of the tribe, although this out- times combined (e.g., Anderson 1993). Erodiscini come was not a focus of the study. In addition, the are characterized by the following features (after Australian Meripherinus Lea belongs in Eugnomini Vanin 1986): body narrow, elongate; prothorax rather than in the New World tribe Camarotini , posteriorly distinctly constricted; integument and several further, undescribed genera occur in shiny, often with bronze sheen, largely glabrous Australia and New Guinea. The group was last with sparse long setae, rarely with scales; head studied by Cawthra (1966), who detailed its dis- hemispherical; eyes fl at; rostrum mostly strongly tinguishing characters and provided a defi nition elongate (up to twice the body length), down- for it, although no single character can be used curved; prementum quadrate to strongly bilobed, to unequivocally identify all genera as belonging labial palps three- to one-segmented; antennae to it. Typical features are the generally long and long, slender, funicles seven-segmented, seg- stout rostrum; often exodont mandibles; fl ex- ment 2 often elongate, clubs long, slender; elytra ible maxillae with long palps; large eyes; elongate 10-striate; ventrites 1 and 2 fused; legs long and head with distinct neck; conical, contiguous fore slender, femora clavate, ventrally usually dentate, coxae; large, fl at, smooth tooth on at least the hind tibiae with uncus and praemucro, tarsi long with femora; mucronate tibiae (except the American strongly elongate tarsomere 1, claws free, simple, genera; Franz 2006); free, simple to appendiculate or toothed. Erodiscus is exceptional in the tribe (as tarsal claws; and ventrites with straight posterior 326 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler margins. The Australian genera Meriphus Erichson, the larvae, although these have not yet been found. Myossita Pascoe, and Orpha Pascoe possess extraor- Geranorhinus (ca. 16) similarly occurs in Palaearc- dinary mouthparts, with the epistome produced tic halophytic habitats, from Algeria to Mongolia, into a triangular process extending over the man- although its species live on Tamarix (Tamaricaceae), dibles, which are fl at with two large, exterior teeth the larvae apparently forming galls and construct- and can open far outward and backward in a hori- ing cocoons for pupation. zontal plane. The function of these outwardly cut- Mecinini (Fig. 3.7.10.7 F and G ). This tribe ting mandibles is not clear. comprises six genera, Cleopomiarus (ca. 40 species), Cawthra (1966) and May (1993) summarized Gymnetron (100), Mecinus (50), Miarus (20), Rhinumia- what is known about the biology of Eugnomini. rus Caldara (4), and Rhinusa Stephens (40). Caldara The adults are generally found on fl owers, feeding (2001) performed a phylogenetic analysis of the on pollen and probably nectar, whereas the larvae genera, with Cionini , Ellescini , and Tychiini as mostly develop under bark of dead branches and out-group taxa. This analysis suggested a sister logs of a variety of plant species. A few, however, group relationship existing between Mecinini develop in living plant tissues and appear to be and Cionini, based on the shared characters of a more host specifi c. In New Zealand, Gonoropterus fi ve-segmented funicle, only partially visible tar- Broun develops in fruits of Nestegis (Oleaceae), Nyx- sal sockets, and basally connate claws, and yielded etes Pascoe in galls on Clematis (Ranunculaceae), three synapomorphies for Mecinini: ventral mar- Hoheria (Malvaceae), and also Nestegis , the latter gin of scrobes in dorsal view convex and slightly seemingly in symbiotic relationships with mealy- protruding; posterior margin of ventrites 3 and 4 bugs (Coccidae), and Oreocalus May in stems and generally laterally angled; tergite VII more or less galls of Hebe (Scrophulariaceae) (May 1993). In Aus- uncovered. Phylogenetically informative charac- tralia, Myossita breeds in the confl orescences and ters between the genera were found to occur on the infructescences ( “ cones ” ) of Banksia (Proteaceae) rostrum (position of antennal insertion), antennae (Scott 1982), whereas Omoides Boheman in Argen- (number and length of funicular segments), pro- tina and Chile is associated with several plant sternum (presence of a rostral canal), elytra, claws, families, mainly Flacourtiaceae (Morrone & Roig tergites (presence of a pygidium), aedeagus, and Juñ ent 1995), and South American species of Udeus spermatheca (shape, location of gland). Vahtera & Champion may occur on palms (Arecaceae), as some Muona (2006) confi rmed the monophyly of the species of this genus have been misplaced in Celetes closely related genera Miarus and Cleopomiarus from (N. Franz, personal observation). COI and ITS2 sequence data. Miarus, Gymnetron , Itini (Fig. 3.7.10.7 E ). This small tribe was and Rhinusa have also been subjected to phyloge- included in Curculioninae by Alonso-Zarazaga & netic analyses (Caldara 2007, 2008; Caldara et al . Lyal (1999) but was transferred to Molytinae by 2010), with particular attention given to the dis- Alonso-Zarazaga (2002). However, in a recent revi- junct Mediterranean-South African distribution sion Meregalli & Borovec (2011) again placed the of Gymnetron and also Cleopomiarus Pierce (Caldara tribe in Curculioninae, based mainly on the differ- et al. 2008). The Afrotropical species of Gymnetron , ent structure of the apex of the hind tibiae and of currently placed in 13 apparently endemic groups, sternite VIII in the males of Ita Tournier, and they seem very closely related to the Palaearctic species regarded this genus to be closely related to Styph- (distinguishable only by few subtle differences), lini, from which it differs in having very short but it is as yet unclear whether they represent scrobes and the antennae inserted near or before the same or a different lineage (Caldara 2008). the middle of the rostrum (at the apex in Styphlini). The Afrotropical species of Cleopomiarus, in con- These characters also occur in Geranorhinus Chevro- trast, form three groups based on characters of lat, which Meregalli & Borovec (2011) consequently their genitalia, two groups including also all the (but tentatively) transferred from Styphlini to Itini. Palaearctic species but the third being endemic to In contrast, Legalov (2011) placed Geranorhinus in the Afrotropical region (Caldara 2005). In Rhinusa , Hyperini: Coniatina, and it appears that the affi ni- ten groups assembled in three clades could be iden- ties of this genus are in need of further study. tifi ed (Caldara et al. 2010), one clade including all Ita (15 species) occurs in stable halophytic habi- species living on Scrophulariaceae and divided tats around the Mediterranean Sea, from Macaro- into two groups associated with the closely related nesia in the west to Turkmenistan and Uzbekistan genera Scrophularia [R. bipustulata (Rossi) group] in the east (Meregalli & Borovec 2011). The adults and Verbascum [ R. tetra (Fabricius) group], whereas are associated with Amaranthaceae (Chenopodio- the other two clades encompass the species occur- ideae, Salicornioideae, and Salsoloideae) and repre- ring on Plantaginaceae of the tribe Antirrhineae. sent the dominant weevil component of halophytic These results indicate that host associations in this coenoses on fl at, warm, or hot plains with saline genus are narrow and conservative as well as partly soil or at the margins of brackish marshes. The wee- congruent with the phylogenetic pattern among vils crawl along the stems of their apparent hosts their hosts, the two plant families being closely during the day, feeding on the succulent stems and related. leaves or hiding in debris at the base of the shrubs. The larvae of Mecinini develop in roots, shoots, It is very likely that goosefoots are the hosts also of leaves, and fl owers, many causing their host organs Curculioninae Latreille, 1802 327 to swell or develop into galls. The latter habit in the position of their fore coxae, whereas occurs in several genera and often in unrelated spe- Morimoto (1962 a) proposed a separate subfamily, cies, and some species of Rhinusa are inquilines in Galloisiinae, for it but evidently did not know galls produced by other species of the same genus the relevant African taxa in detail. Kuschel (1987) (Hoffmann 1958; Arzanov 2000; Caldara 2001, included Metatygini and Galloisiini in Molytinae 2003, 2005, 2007, 2010; Korotyaev et al. 2005). but gave no specifi c reasons for this placement, and Mecinini are predominantly associated with the Alonso-Zarazaga & Lyal (1999) followed it in their families Scrophulariaceae (tribe Scrophularieae), generic catalogue. None of these genera have been Plantaginaceae (tribes Antirrhineae and Plan- studied in detail, however, and several of them tagineae), and Campanulaceae. Mecinus species share characters that contradict Voss’ superfi cial live on Plantagineae and Rhinusa species on both group differences and classifi cation as well as that Scrophulariaceae and Plantaginaceae as presently of Alonso-Zarazaga & Lyal (1999). The distinction circumscribed (Olmstead et al. 2001; Albach et al. between Microstylus and Zantes is obscure, with their 2005), as association supporting the close rela- species agreeing in a number of characters (e.g., tionship between these families as indicated by Zantes also possesses separate fore coxae (Pascoe recent molecular studies. Several species of Mecinus 1888), contra Voss ’ classifi cation, and several having and Rhinusa have recently become the subject of been moved between these two genera in the litera- detailed ecological studies (Tos ˇ evski & Gassmann ture. Microstylini , and Nerthopini are here treated 2004) investigating their potential use as addi- in Curculioninae, whereas the tribe Metatygini is tional biological control agents of Palaearctic retained in Molytinae (q.v. ). species of toadfl ax (Linaria spp., Plantaginaceae) The tribe Microstylini is here taken to include that have become invasive in North America. The the genera Microstylus (ca. 5 species), Zantes (ca. 22), Palaearctic species of Gymnetron live on Veronica Microgalloisia (1), Menesinus Faust (1), and Galloisia (Caldara 2008), currently included in Plantag- (2), the fi rst three African, Menesinus Indian and inaceae (Olmstead et al. 2001; Albach et al. 2005), Galloisia Oriental (one species each in Japan and whereas those in the Afrotropical region (Caldara China). Microstylus species are peculiar subglobu- 2003; Caldara et al. 2010), where Plantaginaceae lar weevils with a short, broad rostrum; bulging are poorly represented, appear to live on various eyes and a defi nite neck; the head with a pair of genera of Scrophulariaceae distributed mainly in deep, posteriorly convergent furrows between the the southern hemisphere, i.e., Diascia, Hemimeris , eyes; the mandibles fl at, adentate, with broadly and Nemesia (Hemimerideae), Hebenstreitia, Selago, rounded apices; the prementum large with labial Sutera , and Tetraselago (Selagineae), and Buddleja palps absent; the postmentum with a median (Buddlejaeae), all closely related to Scrophularieae, tooth or ridge in the male; the fore coxae nar- as well as on Stilbaceae, i.e., Anastrebe, a family very rowly separated; the fore femora with two small closely related to Scrophulariaceae (Olmstead et al. teeth; the tibiae shallowly grooved on the inside 2001). The Palaearctic species of Miarus, and Cleo- and apically with an inner praemucro and a large, pomiarus are associated with genera of Campanula- sharp outer uncus; the claws strongly appen- ceae in the subfamily Campanuloideae ( Adenophora, diculate; the scutellum triangular to pentagonal, Campanula, Jasone, Phyteuma), whereas species of centrally depressed to sometimes forming lateral Cleopomiarus in South Africa and Mexico live on carinae; the elytra with ten striae and interstria genera of the subfamilies Campanuloideae ( Light- 10 strongly costate, without stridulatory fi le; footia, Roella, Wahlenbergia ) and Lobelioideae (Lobe- ventrite 1 posteriorly broadly emarginate in the lia ) (Caldara 2005, 2007). middle; the pygidium exposed; the aedeagus fl at, Microstylini (Fig. 3.7.10.7 H and I). The genera with very short temones; and the tegmen without included in this tribe, proposed by Lacordaire apodeme. In fresh specimens, the body is covered (1865) for Microstylus Schoenherr, were generally by a reddish waxy secretion. The very similar Men- placed in several others, namely Galloisiini, esinus differs in not having the rostrum separated Metatygini, and Nerthopini, but the classifi ca- from the head by a pair of oblique grooves, the tion of these groups has been controversial and fore femora with a single tooth, the tibial apices confusing and remains somewhat unclear. Voss with only an uncus, and ventrite 1 posteriorly (1962) grouped all of them in a subfamily Ner- straight (after Faust 1898). Legalov (2006) discov- thopinae, distinguishing four tribes on the differ- ered that the African Archolabus Voss, described ent combinations of separate or contiguous fore and long treated as a representative of an ances- coxae and free or connate tarsal claws: Nerthopini tral attelabine lineage, is very similar to Menesi- with Nerthops Schoenherr, Microstylus and Teluropus nus; it is in fact congeneric with Microstylus , and its Marshall; Metatygini (as Omophorini ) with Omoph- type species, A. methneri Voss, may be the same as orus Schoenherr, Physarchus Pascoe, Sternechosomus M. hypocritus Hustache (R. Oberprieler, personal Voss, and Zantes Pascoe; Galloisiini with Galloisia observation). Zantes also agrees with Microstylus in Hustache and Microgalloisia Voss; Acallopistini most characters, in particular, the two-pronged with only Acallopistus Schoenherr. Marshall (1935), tibiae, appendiculate claws, male postmental however, had placed the Oriental Galloisia in tooth, and similar scutellum, but it has a lon- Nerthopinae near Microstylus despite the difference ger rostrum that in the female may be ventrally 328 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler infl ated in front of the almost vertical scrobes. personal observation), and evidently also A. malvae Microgalloisia was considered by its author (Voss Boheman in South Africa – and in India – an unde- 1962) to be close to both Microstylus and Zantes scribed species on Abutilon indicum (Pajni & Nanda but differs most conspicuously by having con- 1992). Nerthops guttatus Schoenherr (Fig. 3.7.10.7 J) nate claws. Galloisia is similar to Microstylus develops in the seed capsules of Hibiscus aethiopicus and Zantes in habitus and vestiture (very fi ne, and H. microcarpus , and adults have also been found sparse setae and reddish waxy secretion) and the feeding in fl owers of cotton (Gossypium ) (R. Stals, absence of labial palps (see Morimoto 1962 a), personal communication). but it differs in having the femora with a single Ochyromerini (Fig. 3.7.10.7 L). Often treated as large tooth, the tibiae internally carinate and with a subtribe of Tychiini (Clark et al. 1977; Oberprieler only an uncus, and the claws connate. The ros- 1993), this group currently contains approximately trum in the female is ventrally similar but more 28 genera of fl ower weevils generally characterized spectacularly infl ated than in Zantes . by the robust fore femora carrying a large median Nothing has been reported about the biology of anterior tooth, although this does not occur in any genus of Microstylini . Microstylus appears asso- all the genera. The majority of the genera occurs ciated with fabaceous trees, M. rufus Schoenherr in Asia and was well studied by H. Kojima and (Fig. 3.7.10.7 I) living on Acacia nigrescens and Dal- coauthors in a series of papers (Kojima & Morim- bergia obovata and M. hypocritus on Pterocarpus rotun- oto 1995 a – e, 1996 b – c; Kojima 1997; Kojima & difolius in South Africa (R. Oberprieler, personal Matoba 2002). One genus, Neochyromera Heller, has observation). The stout, subglobular body shape been described from New Guinea (Setliff 2007), of this genus and also its similarity in some other but further undescribed taxa occur there as well characters (dentate fore femora, double-pronged as in northeastern Australia, some of the latter tibial apex, toothed claws, laterally costate elytra) related to Hexeria Pascoe, which was uncertainly with the American genus Camarotus suggest that placed in Erirhininae by Alonso-Zarazaga & Lyal its larvae may also be leaf miners or leaf gallers, and (1999). Five genera are known from Africa, which the same probably applies to Menesinus in India. were last treated by Oberprieler (1993). Two gen- Zantes armatipes has been collected feeding on era listed in the tribe by Alonso-Zarazaga & Lyal leaves of Lecaniodiscus cupanioides (Sapindaceae) and (1999), Sphinxis and Thamnobius , were transferred fogged from Terminalia ivorensis (Combretaceae) in to Anoplini (q.v. ) by Kojima & Morimoto (2000, West Africa (records in Natural History Museum, 2002). A defi nition of Ochyromerini was provided London), whereas Galloisia appears to be associated by Kojima & Morimoto (1995 d); salient features with Actinidia arguta (Actinidiaceae) (H. Kojima, are the uncinate tibiae (uncus often reduced to lost personal communication). in the hind tibiae), femora generally with large Nerthopini (Fig. 3.7.10.7 J and K ). In its cur- fl at tooth fl anked on outside by a row of long erect rent composition (Alonso-Zarazaga & Lyal 1999), setae, appendiculate tarsal claws, elytro-tergal this tribe comprises eight African genera and one stridulation structures in both sexes, endophallus Indian genus, but there is no evidence that they with a pair of elongate sclerites, and tegmen with- together and exclusively form a natural (mono- out parameres. Katsurazo Kojima has spectacularly phyletic) group. Nerthops (1 species in South Africa) modifi ed and infl ated antennal funicles. However, agrees closely with Acallopistus (ca. 11 species in the exact concept and composition of the group Africa, also in India) in the structure of the anten- remains unclear. nae (seventh funicular segment closely fi tting to Biological information for the Ochyromerini club), the dentate fore femora, the uncinate tibiae was summarized by Oberprieler (1993) for the with long, ascending talus, the strongly bifi d claws, African fauna and is included for the Oriental and the exposed pygidium, although Nerthops dif- genera in the papers by Kojima & Morimoto (1995 fers in having a shorter, tapering rostrum, large, a– e, 1996 b, c). The adult weevils are associated ventrally contiguous eyes, and adelognathous with many plant families (e.g., Anacardiaceae, mouthparts. The two genera appear closely related, Celastraceae, Clusiaceae, Combretaceae, Ebena- but further study of both is required. The relation- ceae, Geraniaceae, Hamamelidaceae, Fabaceae, ships of the Madagascan genera Acallopistellus Hus- Lauraceae, Moraceae, Oleaceae), but the habits of tache, Achynius Fairmaire, Helaeniella Hustache, the larvae are much less well known. The larvae of Paranerthops Hustache, and Pseudodesmidophorus the Oriental Drepanoscelus Marshall and Ochyromera Hustache to Acallopistus and Nerthops are in need of Pascoe and the African Daeneus Oberprieler have verifi cation, whereas Microstylus, and Menesinus rep- the spiracles of segment VIII located on elongated resent a different group ( Microstylini). dorsal protuberances (Gardner 1938; Cuda et al . All recorded hosts of Nerthopini belong to the 2008; Oberprieler 1993, respectively), an appar- family Malvaceae. Acallopistus develops in fruits ent adaptation for breathing when developing in of Abutilon , both in Africa – A. abutilonis in Sudan the mostly fl eshy fruits of Machilus (or Persea, Lau- and A. maculithorax Hustache in Kenya and Uganda raceae), Ligustrum (Ebenaceae), and Bequaertioden- (Marshall 1950), A. fallax Boheman of Abutilon dron (Sapotaceae), respectively. This modifi cation angulatum in Namibia (S. Neser & R. Oberprieler, also occurs in undescribed Australian species and Curculioninae Latreille, 1802 329

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Fig. 3.7.10.8 Habitus, adults. A, Myrmex scutellaris (Sharp) (Otidocephalini); B, Piazorhinus scutellaris (Say) ( Piazorhinini); C, Pyropus cyaneus (Herbst) (Pyropini); D, Tachyerges salicis (Linnaeus) (Rhamphini); E, Orthochaetes setiger (Beck) ( Styphlini); F, Smicronyx discoidus LeConte ( Smicronychini); G, Storeus maximus Lea ( Storeini); H, Tychius uralensis Pic ( Tychiini); I, Lignyodes enucleator (Panzer) (Tychiini); J, Ulomascus parallelus Marshall ( Ulomascini); K, Tivicis sp. (Viticiini); L, Pachytychius simillimus Desbrochers des Loges (Curculioninae incertae sedis). Scale bars 1 mm. 330 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler probably more widely in the tribe. The larvae of A key to the species of Prionobrachium was given by the South African Teinomphasus robustus Oberpri- Hustache (1938). eler (Fig. 3.7.10.7 L) develop in the fl eshy fruits Prionobrachiine larvae appear to be leaf min- of Hexalobus monopetalus (Annonaceae), pupating ers as well, with one species of Chelotonyx living on in fallen fruits on the ground (R. Oberprieler, Malvaceae (Anderson 1993). personal observation). Ochyromera ligustri Warner, Pyropini (Fig. 3.7.10.8 C ). This very small tribe the ligustrum weevil, is a pest as well as a possible is characterized by a thin, angled lateral carina natural control agent of Chinese privet (Ligustrum separating the pronotum from the pleuron of the sinense , Oleaceae) in North America (Cuda et al . prothorax (Marvaldi & Lanteri 2005) and composed 2008), the larva feeding in the seeds, and Ochy- of two genera from central and southern America, romera miwai Kô no is considered a pest of Diospyros Craspedotus Schoenherr (5 species) and Pyropus kaki (Oleaceae) in Taiwan, the larvae mining in the Schoenherr (3). leaves (Kojima & Morimoto 1996 b, c). The lar- Adults of Craspedotus and Pyropus have been col- vae of Endaenidius and Endaeus appear to develop lected on Brassicaceae and Capparaceae, respec- in fl owers of various magnoliids, with the adults tively (Silva et al. 1968; Anderson 1993; Pakaluk shown or indicated to be the main pollinators of 1996), the larvae of the former genus forming galls several basal angiosperm families (see Biology and on leaves and shoots of Rubiaceae or developing in Ecology above). seeds pods of Lamiaceae and those of the latter tun- Otidocephalini (Fig. 3.7.10.8 A ). This American neling in stems and leaf petioles of Brassica (Ander- tribe, comprising only the fi ve genera Laemomerus son 1993; Pakaluk 1996). Kirsch (2 species), Micromyrmex Sleeper (10), Myr- Rhamphini (Fig. 3.7.10.8 D). This large, almost mex Sturm ( > 100), Oopterinus Casey (8), and Ptinopsis cosmopolitan tribe contains about 20 genera and Champion (2), appears closely related to Erodiscini is presently divided into three subtribes, Ixalmina , (Vanin 1986; Marvaldi & Lanteri 2005) and is some- Rhamphina , and Tachygonina . All the Holarctic times combined with the latter group (e.g., Ander- genera, among them Orchestes Illiger (ca. 80 spe- son 1993). The species are similarly characterized cies), Pseudorchestes Bedel (ca. 35), Rhamphus (ca. by a narrow, pedunculate prothorax and often 15), Rhynchaenus (ca. 10), and Tachyerges Schoen- mimic ants. Ivie & Sikes (1995) commented on the herr (ca. 12), are included in the fi rst subtribe. In taxonomic challenges regarding the large genus the Afrotropical region, in addition to these, there Myrmex and presented a key to the genera. are two monobasic, apparently endemic genera, Larvae and adults of Myrmex are associated Macrorhynchaenus Hustache from Madagascar with numerous plant families including Araceae, and Rhamphonyx Voss from Sudan. Orchestes and Asteraceae, Fagaceae, Platanaceae, Santalaceae, Rhamphus also occur in Australia – the species of Sapotaceae, Smilacaceae, and Ulmaceae, with the the former placed in Rhynchaenus by Zimmerman larvae mining in stems and in several instances (1992), but their characters agreeing with those associated with galls of cynipid wasps (Burke et al. of Orchestes as differentiated by Kojima & Morim- 1975; Anderson 1994). The larvae of Micromyrmex oto (1996 a) – and several further unidentifi ed or asclepia Ivie & Sikes develop in the stems of Ascle- undescribed genera are known from the region piadaceae (Ivie & Sikes 1995). (Zimmerman 1992). The tribe is not represented in New Zealand, however; the sole genus listed Piazorhinini (Fig. 3.7.10.8 B ). This tribe includes there by Leschen et al. (2003), Geochus, displays fea- only the American genus Piazorhinus (ca. 30 species). tures of phrynixine Molytinae (q.v. ) rather than of Often included in Prionomerini (thus Camarotini, Rhamphini . Its larva mines in dead leaves of Wein- q.v .), e.g., by O’ Brien & Wibmer (1982), Anderson mannia (Cunoniaceae) and is similarly adapted to (1993), and Lyal & King (1996), its fore femora, this lifestyle (May 1992, 1993), but leaf mining however, carry only a small tooth. A key to most of has evolved numerous times in Curculionidae and the species of Piazorhinus was provided by Hustache does not represent direct evidence of relationship. (1939) and Voss (1940, 1944). The subtribe Ixalmina comprises only one genus, Piazorhinus is associated with Quercus (Fagaceae) the Oriental Ixalma (ca. 15 species), whereas the and Coccoloba (Polygonaceae) (Anderson 1993, three genera of Tachygonina, Laemorchestes Cham- 2002), the larvae mining in leaves (Marvaldi & pion (1), Tachygonidius Champion (4), and Tachygo- Lanteri 2005). nus Gu é rin-M é neville (ca. 40), occur only in the Prionobrachiini . Another American tribe of New World. Morimoto (1984) studied the phy- uncertain status, the Prionobrachiini comprise logenetic relationships among the Japanese gen- only two South American genera, Chelotonyx Water- era of Rhamphini and Kojima & Morimoto (1996 house (5 species) and Prionobrachium Faust (ca. 10). a) those of the Oriental fauna, establishing the Anderson (1993) included the group in Priono- monophyly of the tribe on the basis of two synapo- merini and regarded also Macrorhoptus LeConte as morphies (maxillary palps one- to two-segmented belonging in it, sharing the host family (Malvaceae) and hind femora with an apodeme, modifi ed for with Chelotonyx . The adults are again characterized jumping). The sister group of the tribe is, how- by the fore femora carrying a large, serrate tooth. ever, not established. Anderson (1993) sketched Curculioninae Latreille, 1802 331 the evolution of the North American genera, and subgenus of Smicronyx (Anderson & Cox 1997). The Hespenheide (1992) reviewed the North American nominate subgenus Smicronyx extends its distri- species of Tachygonus . bution into northern Australia, with two species Larvae of Rhamphini generally mine in young described but several more present in collections, leaves of broad-leaved trees belonging to many although it is unclear whether any of them is truly families, e.g., Anacardiaceae, Betulaceae, Capri- native there. foliaceae, Fagaceae, Myricaceae, Rosaceae, Salica- The larvae of Smicronyx develop in seeds or ceae, and Ulmaceae, feeding on the parenchyma in stems of herbaceous plants in numerous families, winding tunnels and often causing serious dam- including Asteraceae, Convolvulaceae (Cuscuta ), age by defoliation, but those of Pseudorchestes live Fabaceae, Malvaceae, and Scrophulariaceae. In on herbaceous Asteraceae (Hoffmann 1958; Scherf some cases, they incite galls in the stems or seeds 1964; Anderson 1989; Hespenheide 1992; Kojima of their hosts and pupation occurs in infested & Morimoto 1996 a; Lee & Morimoto 1996). Pupa- plant tissues or in the soil (Hoffmann 1958; tion generally occurs in the mines, but some spe- Anderson 1962; Scherf 1964; Anderson & Cox cies drop to the ground in pieces of leaf, in which 1997; Anderson et al. 2006). Several species have they pupate. The larvae of Orchestes hustachei Klima been used as biological control agents of weeds, develop in aphid galls, feeding on the aphids as e.g., the seed-feeding Mexican S. lutulentus Dietz the primary source of food (Tomisawa & Akimoto of Parthenium hysterophorus (Asteraceae) in Austra- 2004). In Australia, Orchestes australiae Lea has been lia (Dhileepan & Strathie 2009) and the galling reared from seedpods of Brachychiton populneus S . (Smicronyx) guineanus Voss, S. ( Afrosmicronyx ) (Malvaceae) and an undescribed Orchestes species dorsomaculatus Cox, and S. (Afrosmicronyx) umbrinus from leaf mines on Argyrodendron trifoliolatum (Mal- Hustache of parasitic witchweeds, Striga hermonth- vaceae), whereas Rhamphus lives on Acacia (Faba- ica and Alectra vogelii (Scrophulariaceae), in Africa ceae), where the larvae presumably mine in leaves (Anderson & Cox 1997), all with only moderate (although some adults have been reared from success. Adults of Smicronyx squalidus Casey appear galls). A number of Rhamphus species also occur to the only weevil recorded to feed on nectar, from on Acacia in southern Africa, which is remarkable extra-fl oral nectaries of Desmanthus (Fabaceae) because the African and Australian acacias are not (Davis 2007). closely related and do not normally share any wee- Sphaeriopoeini . This monobasic tribe was vil taxa. The larvae of Ixalma feed in blotch-like recently proposed for a single Chilean genus and mines along the edge of leaves of Euptelea (Eupt- species, Sphaeriopoeus faber Kuschel, with a most eleaceae) and Kadsura japonica (Schisandraceae) in unusual biology (Kuschel 2003). It is a small Japan, hibernating and, in spring, pupating in the (2 – 3 mm long), broad, compact species with mines (Morimoto 1984; Lee & Morimoto 1996). In appressed pubescence and erect sensory dorsal North America, larvae of Tachygonus mine in leaves hairs; a short, stout rostrum with large, broad, of Quercus (Fagaceae), and adults occur on Fabaceae tooth-like ventral process locking behind the fore ( Coursetia, Robinia), Rhamnaceae ( Berchemia ), and coxae; coarsely facetted eyes; seven-segmented Ulmaceae (Ulmus ) (Hespenheide 1992), whereas funicles; three-segmented maxillary and labial Melo da Silva et al . (2007) recorded a Tachygonus palps, the latter contiguous; anteriorly toothed species mining in leaves of Erythroxylum (Eryth- fore coxae; toothed femora; uncinate tibiae, the roxylaceae) in Brazil. Also in Brazil, larvae of fore and middle ones in female with long prae- Tachygonidius mine in leaves of Ochnaceae (Vanin & mucro; thin, basally connate tarsal claws; ten-stri- Mermudes 2007). ate elytra without stridulatory fi les, infrolateral Smicronychini (Fig. 3.7.10.8 F). This tribe of fl ange extending to middle of length; rudimen- fi ve or six genera is clearly differentiated from tary proventriculus; ring-like rectal valve; sternite other Curculioninae by a basally constricted ros- IX of male asymmetrical and tegmen with com- trum and connate claws. Most species belong to the plete parameres; sternite VIII of female well scler- widespread, mainly Holarctic genus Smicronyx (ca. otized but gonocoxites weakly so, without styli. 140 species), whereas the smaller Sharpia Tournier Kuschel (2003) placed the tribe Sphaeriopoeini (ca. 15) occurs in the Palaearctic region, Promeco- near Molyt inae in his larger concept of a subfam- tarsus Casey (4) in North America, Afrosmicronyx ily Curculion inae (Kuschel 1995), but Elgueta & Hustache (5), and Topelatus Hustache (6) in Africa, Marvaldi (2006) included it in Curculioninae in and Hedychrous Marshall (1) in India. Anderson’ s the narrower sense as adopted here. (1962) monograph of North American Smicronyx is Sphaeriopoeus faber lives in the canopy of the the latest comprehensive treatment of the group, deciduous Nothofagus obliqua (Nothofagaceae), whereas the taxonomy of the numerous Palaearc- where the female rolls and folds whole new leaves tic species of this genus as well as that of the closely into tight balls, without incising the leaf lamina related Sharpia is still confused. Smicronyx is gener- as attelabids do. A single egg is laid into each roll, ally divided into fi ve subgenera, three restricted apparently before the rolling commences, and on to North America and one to western Europe, completion of the rolling, the leaf ball is cut off and and Afrosmicronyx is sometimes also treated as a falls to the ground, where the larva consumes the 332 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler contents. This leaf-rolling method is apparently Notionomus Erichson, and Symbothynus Blackburn unique in weevils (Kuschel 2003). have strongly imbricate to agglutinated scales and Storeini (Fig. 3.7.10.8 G ). The concept of this appear to all live on sedges in semi-aquatic habitats, tribe is unclear and contentious. In a narrow sense, suggesting that they may not be closely related to it includes those genera with a pectoral canal in the other genera. All of these Australian genera are front of the fore coxae and the walls of the canal in need of study to assess both their proper generic being incised to form a narrow cleft to large “ win- delineations and their taxonomic affi nities. Both dow” or “ peep-hole ” (Fig. 3.7.10.2), which may be in the narrow and the wide sense, Storeini appears lined with dense setae and in some species come to be a purely Australo-Pacifi c group, in the nar- to lie over the eyes when the head and rostrum row one extending from Australia, New Caledo- are folded down and through which the weevil nia, and Fiji north to the Philippines, and Taiwan may then be able to see (Lea 1927). On this rather and in the wider sense also including New Zealand unique structure, Storeini sensu stricto include and probably other Pacifi c islands as far north as the large but composite genera Storeus Schoen- the Ryukyus. The four African genera listed in herr (ca. 50 species) and Emplesis Pascoe (ca. 100) Storeini by Alonso-Zarazaga & Lyal (1999) most and also Leucomelacis Lea (2), Microberosiris Lea (2), probably do not belong in this tribe; Anchonocra- Placorrhinus Marshall (1), and Pseudostoreus Lea (1) nus Marshall appears related to the Phacellopterus in Australia as well as Aporotaxus Perroud (1) and Schoenherr group of Anthonomini (Oberprieler Griphosternus Heller (1) in New Caledonia and Dio- 1993) and Pansmicrus to Anoplini ( q.v.), whereas mia Pascoe (1) in New Guinea (the last misplaced the identities and affi nities of Australafer Alonso- in Molytinae: Ithyporini by Alonso-Zarazaga & Zarazaga & Lyal and Merocarterus Hustache require Lyal 1999). Zimmerman (1992, 1994) used an investigation. Also, the placement of the only two expanded concept of Storeini by including also the Palaearctic genera in Storeini, Aubeonymus Jacque- Australian genera Arthriticosoma Lea, Cycloporop- lin du Val and Pachytychius (previously in Erirhin- terus Lea, Euthebus Pascoe, Hybomorphus Saunders & inae), is highly doubtful (Gonz á lez 1968; Caldara Jekel, Hybophorus Waterhouse, Lybaeba Pascoe, 1978), and they are here treated as incertae sedis . Melanterius Erichson, Moechius Pascoe, Neomelan- Biological information on Storeini is mostly terius Lea, Rhinidotasia Lea, and Teutheria Pascoe, limited to host-plant records, although not all of which do not have the “ peep-hole ” type of pectoral these necessarily refl ect true (larval) hosts. Among canal and also differ in several other features from Storeini sensu stricto, Aporotaxus develops in fi gs Storeus and allies, most importantly in possess- (Kuschel 1990 a) and also some Storeus species ing uncinate tibiae and sclerolepidia and appear in Australia occur on Ficus (Moraceae), whereas to be Molytinae of a cleogonine affi nity (except Storeus albosignatus (Blackburn) lives on various Euthebus , Hybomorphus , and Rhinidotasia ). Kuschel Acacia species (Fabaceae) both in Australia and in (1990 a) tacitly used yet another broader concept of New Zealand, where it is adventive, and Storeus bae- Storeini, loosely characterized by the possession of odontus Lea has been collected on Dodonea cuneata divaricate claws and, in the male, of elytral stridu- (Sapindaceae) (Rheinheimer 1989). It is, however, latory fi les and a broadly angulate, asymmetrical uncertain whether these species are congeneric. sternite IX, and he included in it genera such as Emplesis Pascoe is generally found on Eucalyptus , Palontus from New Caledonia, Aneuma Pascoe from and Emplesis bifoveata Lea – misidentifi ed as Storeus New Zealand and Elleschodes and Rhinidotasia from majusculus (Blackburn) by Rheimheimer (1989) Australia, which either have a pectoral canal with- and redescribed as Storeus acmenae by Rheinheimer out an incision in its walls or no canal at all. This (1996) – occurs on Syzygium smithii (Myrtaceae) also concept includes another 14 genera of New Zea- in New Zealand where it is adventive. Leucomelacis land “ fl ower weevils ” (Leschen et al. 2003) as well quadrinotatus Lea has been reared from Syzygium as approximately 47 Australian genera classifi ed fruits and Placorrhinus placitus Lea collected on in Tychiini by Zimmerman (1992) but later placed Doryphora (Monimiaceae). without tribal assignment in Curculioninae (Zim- Biological records for several New Zealand gen- merman 1994; Alonso-Zarazaga & Lyal 1999). Of era of Storeini sensu lato ( Alloprocas Broun, Aneuma , the latter, Imathia was transferred to Storeini by Hypotagaea Pascoe, Neomycta Pascoe, Oropterus, Peri- Kojima & Morimoto (1998). These Australian gen- storeus Kirsch, Phorostichus Broun, Praolepra Broun, era are a diverse group, however, including besides Simachus Broun) are given by May (1987, 1993) and many genera with divaricate claws also others Kuschel (1990 b); the larvae often are leaf miners but with simple, divergent claws (such as Abethas Zim- also develop in stems, fl ower buds, young fruits, merman, Lexithia Pascoe, Olbiodorus Blackburn, and seeds of families spanning conifers (Podocar- Paryzeta Pascoe, Plaesiorhinus Blackburn) or with paceae) to monocotyledons (Asteliaceae) and vari- strongly toothed to bifi d ones (Agestra Pascoe, ous dicotyledons (Asteraceae, Ericaceae, Fabaceae, “Ellescus ” , Hibberticola Lea) as well as several with- Nothofagaceae, Malvaceae, Myrtaceae, Onagraceae, out onychia (Anarciarthrum Blackburn, Cenchrena Pittosporaceae, Polygonaceae, Rubiaceae, Scrophu- Pascoe, Micraonychus Lea, Misophrice Pascoe, Thechia lariaceae). Of the Australian genera, Cydmaea Pascoe, Pascoe). Cenchrena, Ethadomorpha Blackburn, Dicomada Pascoe, and Erytenna Pascoe breed in stems Curculioninae Latreille, 1802 333 or fruits of Hakea and Grevillea (Proteaceae), the lar- unrelated taxa such as Allium (Amaryllidaceae), vae feeding on the seeds to the extent that several Aquilegia and Clematis (Ranunculaceae), Cycla- species have been used as biological control agents men (Primulaceae), Myosotis (Boraginaceae), Plan- against the invasive Hakea sericea in South Africa tago (Plantaginaceae), and Viola (Violaceae), but (Gordon 1999; Kluge & Gordon 2004). The clawless appears to prefer Asteraceae, on which the larvae Micraonychus , Misophrice , and Thechia live on Alloca- mine in leaves before pupating in the soil (Scherf suarina (Casuarinaceae) and Misophricoides Rhein- 1964; Dieckmann 1986; Osella & Zuppa 1994). heimer on Callitris (Cupressaceae), whereas Agestra , Styphlus and Pseudostyphlus also live on Asteraceae and Plaesiorhinus develop in seedpods of Bossiaea and (the genera Barkhausia , Hedypnois, and Matricaria), Daviesia (Fabaceae) and Hibberticola in galls on Hib- where the larvae develop in fl ower heads, seeds, or bertia (Dilleniaceae). Some Elleschodes species, such as roots (Hoffmann 1958; Scherf 1964; Dieckmann E. hamiltoni Blackburn, pollinate Eupomatia (Eupo- 1986; Bland & Nelson 1997; Ostoja-Starzewski matiaceae) (Armstrong & Irvine 1990), whereas E. 2002; Knutelski 2005). Trachystyphlus appears to compactus Lea and some other species live on Pulta- live on Caryophyllaceae ( Cerastium, Heliospermum ) nea (Fabaceae) but are not evidently congeneric with (Franz 1974). Interestingly and seemingly unique the former. Eniopea Pascoe, Epacticus Blackburn, in Curculioninae, some species, e.g., Orthochaetes Eristinus Lea, Myllorhinus Boisduval, and Paryzeta are setiger (Beck) and O. insignis (Aubé ), are partheno- associated with Myrtaceae (Eucalyptus, Leptospermum, genetic at least in part of their distribution range Melaleuca), Myllorhinus boring in shoots of Eucalyptus (Gonzá lez 1967; Osella & Zuppa 1994). (Jones & Potts 2000). Symbothynus lives on Gahnia Thecesternini . Both the composition and the sedges (Cyperaceae), and Cenchrena , Ethadomorpha , classifi cation of this small tribe have been con- and Notionomus occur in similar habitats and are fused in the literature. In the catalogue of Alonso- likely to have similar hosts. Zarazaga & Lyal (1999), it was placed in Entiminae Styphlini (Fig. 3.7.10.8. E ). This Palaearctic tribe, and included four genera; however, the Palaearctic generally included in Erirhininae until 15 years genus Herpes Bedel belongs in Hyperini (Alonso- ago, is mainly characterized by having a stout ros- Zarazaga & Lyal 2002) and the New Caledonian trum, the scrobes directed toward the anterior Cyphomastax Marshall in Storeini (G. Kuschel, per- margin of the eyes, the antennae inserted near the sonal communication). Thus, Thecesternini com- apex of the rostrum, small eyes, the humeral area prise only the two New World genera Thecesternus of the elytra constricted, all or the odd-numbered Say (ca. 6 species) and Arodenius Heller (2). They are interstriae usually distinctly convex, and the legs characterized by having the prosternum anteri- distinctly stout with short tarsi. It currently com- orly deeply and sharply excavate (the canal closed prises ten extant genera: Orthochaetes Germar (ca. off before the fore coxae), the prementum deeply 15 species), Paraphilernus Desbrochers (1), Paroryx excavate, the tibiae with two apical spines (in Thece- Reitter (1), Philernus Schoenherr (1), Pseudostyph- sternus representing a mucro and an apparently lus Tournier (5), Salsolia Bajtenov (1), Styphlidius fused spur, in Adorenius a praemucro and uncus), Penecke (1), Styphlus Schoenherr (20), Trachystyph- the elytra fused in Thecesternus but separate, with lus Alonso-Zarazaga & Lyal (3), and Turanostyphlus unequal fl anges, in Adorenius , the aedeagus dorsally Davidian & Savitsky (1). Of the two other genera solid at its base and with loosely attached temones, listed in the tribe by Alonso-Zarazaga & Lyal (1999), the ovipositor with the distal gonocoxites short, Geranorhinus was recently transferred to the closely cylindrical, with a large apical stylus, the tergite related tribe Itini ( q.v. ) by Meregalli & Borovec above it with the apex produced into two fl at, (2011), whereas the Madagascan Madecastyphlus upcurved processes (for pushing into soil), and the Richard, as the only African genus, is most unlikely proventriculus with eight long, internal processes to belong in it. Several genera were partially taxo- ending in a fl at scoop with serrated margins (Kiss- nomically revised (Gonzá lez 1967; Osella 1981; inger 1963), similar but not identical to those of Osella & Zuppa 1994; Davidian & Savitsky 2000), Gonipterini. Marvaldi (1997) related Thecesternus but none of these papers treated the genera phylo- to various tribes of Entiminae and Cyclominae genetically. Most species have a limited distribu- because the mandibular thecae of its pupa also tion, and some of them [ Orthochaetes alpicola Daniel, carry setae, but it does not belong in Entiminae due Styphlidius corcyreus Reitter, and Trachystyphlus alpi- to its phanerognathous mouthparts and absence nus (Penecke)] were divided in one or more subspe- of deciduous mandibular cusps in the adult and cies (Osella 1981; Osella & Zuppa 1994). conical antenna in the larva, and it also does not fi t The biology of Styphlini is poorly known. into the current concept of Cyclominae (see Ober- Atypically for Curculioninae, the adults are usu- prieler 2010) due to the epicranial position of setae ally found under stones or by sifting mosses, des3 in the larva (Marvaldi 1997; Oberprieler 2010). dead leaves, and marcescent roots in bare patches For the moment, it is therefore here dealt with in or at the margins of woods both in plains and at Curculioninae. high altitudes (some Orthochaetes , Trachystyphlus ) The larva of the Mexican Thecesternus hirsu- or in sandy and dry soil (some Styphlus ). Ortho- tus Pierce feeds semi-ectophytically in gall-like chaetes lives on a wide range of plants, including growths on the roots of Parthenium (Asteraceae) 334 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler

(McClay & Anderson 1985) and the species has been families of Caryophyllaceae. The Afrotropical spe- considered for biological control of P. hysterophorus cies, divided into eight groups of which seven are in Australia (McFadyen 1992), where this plant is considered endemic to the region (Caldara 1989 b, a severe weed of crops and pastures, but it was not 1993), appear to be closely related to both Sibinia released (Dhileepan & Strathie 2009). sensu stricto and Dichotychius but seem to live on dif- Tychiini (Fig. 3.7.10.8 H and I ). This concept ferent plant families, especially Aizoaceae, which is of this tribe is complex and contentious. In its tra- closely related to Caryophyllaceae, and their rela- ditional status as a subfamily, Clark et al. (1977) tionships are still unclear (Caldara et al. 2008). In divided it into four tribes, Ellescini , Lignyodini, Tychius , most species (180) occurs in the Palaearctic Ochyromerini (as Endaeini), and Tychiini, and region, with the remainder in the Nearctic (10) and defi ned it by two apparent synapomorphies, later- Afrotropical (ca. 45, mainly in South Africa) regions ally angled ventrites and mesadly directed pronotal and on the Indian subcontinent (3). The Palaearc- scales or setae. Morimoto (1962 b) and Oberprieler tic species are arranged in two subgenera, Apeltar- (1993) pointed out, however, that both these char- ius Desbrochers and Tychius sensu stricto, the latter acters occur in several other curculionid groups divided into 22 species groups (Caldara 1990). The and cannot be used to identify a group as com- North American species possibly belong to four of posed by Clark et al . (1977) as monophyletic. In the Palaearctic groups (Clark 1971), whereas the a series of subsequent papers on the Asian ochy- southern African species fall into six groups, four romerines, H. Kojima and coauthors (Kojima & apparently endemic but the other two includ- Morimoto 1995 a – e, 1996 b – c; Kojima 1997; ing also several Palaearctic species (Caldara 1989 Kojima & Matoba 2002) treated these as a distinct a, 1996). However, the four endemic groups also tribe, and Ochyromerini and Ellescini are therefore appear closely related to some Palaearctic ones, also here regarded as tribes separate from Tychiini, from which they can be distinguished only by a the latter comprising Demimaeina , Lignyodina , few subtle characters (Caldara et al. 2008). In India, and Tychiina as subtribes (as in Alonso-Zarazaga three groups are represented, only one of them & Lyal 1999). Oberprieler (1993) discussed the dif- endemic, the T. eremita Caldara group (Caldara Q: “or” was fi culties of properly delineating these groups as 1989 a, c). Phylogenetic analysis of all the groups of changed to well as the tribe Tychiini, and a larger phylogenetic Tychius was so far unsuccessful due to the extreme “of ”. Correct? study of all curculionine genera with angled ven- uniformity of the species and the absence of clear trites is required to elucidate natural groups in this synapomorphies. No Tychiini appear to occur in complex. the Australo-Pacifi c region, the various genera All three subtribes of Tychiini comprise only listed in this tribe by Zimmerman (1994) belong- two genera. In Demimaeina, Demimaea Pascoe (ca. ing in Storeini sensu lato (q.v. ) and the New-Guinean 25 species) is largely Oriental but also occurs in Sibiniella Voss probably in that group as well. A Africa (Voss 1962; Oberprieler 1993), whereas the species of Sibinia ( S. fastigiata Clark) has been intro- monobasic Pseudocionus Marshall is only known duced for the biological control of the weed Mimosa from South Africa. The Lignyodina, comprising pigra (Fabaceae) in northern Australia but evidently the largely Holarctic genus Lignyodes Dejean (ca. failed to establish (Ostermeyer & Grace 2007). 25 species) and the American Plocetes LeConte (ca. Host associations are relatively narrow in Tychi- 35), were studied by Clark (1980, 1982), whose ini. The larva of Demimaea tunnels in green stems phylogenetic analyses divided the former genus of Ficus (Moraceae) (Gardner 1934), but no biologi- into three groups (subgenera). Kojima & Morim- cal information is available for Pseudocionus . Lig- oto (2003) recently recorded Lignyodes also from nyodes lives on Oleaceae (Clark 1980) and Plocetes Japan. Both genera of Tychiina, Sibinia Germar on Rubiaceae (Clark 1982), the larvae feeding in and Tychius Germar, are very large (ca. 240 species the seeds and pupating in the soil. The Palaearctic each) and widespread. About 50 species of Sibinia species of Sibinia develop in seeds of mainly Caryo- occur in the Palaearctic region (Caldara 1979, 1984 phyllaceae and Plumbaginaceae (Hoffmann 1958; b), 130 in the New World, mainly South America Scherf 1964; Caldara 1979, 1984), and the Afro- (Clark 1978), and 60 in subsaharan Africa, mainly tropical ones are at least partly associated with the South Africa (Caldara 1989 b, 1993). The genus is related family Aizoaceae (Caldara 1989 b), whereas tentatively divided into three subgenera, Dichoty- the American species (subgenus Microtychius ) breed chius Bedel, Microtychius Casey, and Sibinia sensu in seeds of Fabaceae [Clark 1978; see also Anderson stricto . The almost entirely American Microtychius , (1993) for a discussion of host-mediated speciation distinguishable by the cup-shaped, sclerotized in Sibinia ]. Tychius has an even narrower host range, base of its spermathecal gland and its host-plants all known hosts belonging to Fabaceae (Scherf (Fabaceae: Mimosoideae), was subjected to a pre- 1964; Clark 1971, 1977, 1978; Clark & Burke 1977; liminary phylogenetic analysis by Clark (1978). Clark et al. 1978; Caldara 1989 a, 1990; Anderson No clear apomorphies have been identifi ed for the 2002; Oberprieler et al . 2007) and certain lineages Palaearctic Dichotychius so far (Caldara 1979), but appearing to have phylogenetically highly con- the taxon may be characterized by its different host servative host associations. Most species are seed plants (Plumbaginaceae: Staticoideae), whereas predators, but a few form galls on leaves, fl owers, or species of Sibinia sensu stricto live on several sub- pods (Scherf 1964; Clark & Burke 1977; Korotyaev Curculioninae Latreille, 1802 335 et al. 2005). When mature, seed-feeding larvae typi- Morimoto (2007) added another genus to the cally leave the pods and enter the soil to pupate tribe, Aviticis , which retains very small onychia (Clark & Burke 1977). but without claws. Oberprieler (2010) recently Ulomascini (Fig. 3.7.10.8 J). This small tribe also included the Samoan Samoacis Zimmerman comprises three genera of most bizarrely shaped and the Australian Eupholocis Lea, Peliocis Lea, and weevils, Buttikoferia Roelofs (1 species), Cratopechus Platynotocis Lea as well as Zimmerman ’ s (1992, Marshall (3), and Ulomascus Fairmaire (9). They Plate 561) undescribed genus Austrocis (a nomen are only known to occur in central Africa, from nudum ) in the tribe, based in part on a study of the Sierre Leone in the west to Kenya in the east. The group commenced by E. C. Zimmerman before his weevils are fl at and long, with a broad, spatulate death in 2004. In this composition, the Viticiini rostrum; fl at, calliper-like, non-overlapping man- comprise two groups, one including Viticis, Tivicis, dibles; seven- or six-segmented funicles and often Aviticis, and Eupholocis, characterized by having greatly elongate antennal clubs; large coxae and six-segmented funicles, separated fore coxae, and trochanters; strongly infl ated, fl attened, dentate dentate femora, the other including the closely femora often densely adorned with long hairs; related Austrocis , Platynotocis , and Samoacis as well short stout tibiae folding against or under the fl at, as Peliocis , which have seven-segmented funicles, broad femoral tooth; fl at tarsi with large, deeply contiguous fore coxae, and unarmed femora. A lobed third tarsomeres and strong, widely divari- number of other similar but undescribed gen- cate but simple claws; elytra with eight or nine era occur in Australia and New Guinea, and the punctate striae. In Ulomascus insolitus Hoffmann, precise composition and delineation of the tribe the onychium of the hind tarsi is in the female require further study. Morimoto ’ s (1983) defi - extremely elongate (as long as a tibia) and car- nition of Viticiini therefore only applies to the ries a single, equally long claw folded back onto former group. Viticis (6 species), redescribed by the onychium, a most extraordinary feature of Kuschel (2008), occurs from Guam south to New unclear function. The females of Ulomascus and Caledonia and from Amboina east to the Marque- Cratopechus also have a large, tapering tergal pouch sas Islands. Tivicis (2) is presently known only from situated under a very large, tergite-like apodeme the Ryukyu Islands of Japan, but Eupholocis macula- that is extended by a large membrane with a bun- tus Lea from New Britain evidently also belongs to dle of tendon-like fi bers between apodeme and it as well as at least one undescribed species from pouch, and the pouch itself ends in similar hard- southern New Guinea and northern Australia. ened fi bers from which powerful muscles extend Aviticis (1) is only known from Taiwan and Euphol- to the extraordinary long genitalia, which, when ocis dentipes Lea from northern Queensland. Aus- unfolded, are nearly twice as long as the body of trocis, Platynotocis , and Peliocis (1 species each) are the weevil (Kuschel 1987). The taxonomic affi ni- recorded from northeastern Australia, but a large ties of the group are unclear. species of or near Peliocis occurs in southwestern Nothing specifi c is known about the biology of Western Australia. Ulomascini , except that the adults are apparently The available biological information of Vitici- nocturnal (usually collected at lights) and the ini was summarized by Oberprieler (2010). Viticis Kenyan Cratopechus arundinarum Marshall and appears associated with Ficus (Moraceae), whereas C. gedyei Marshall have been found on reeds. Because Austrocis bicaudatus Zimmerman (undescribed, a of the long, golden hairs on the femora and sides of nomen nudum ) mines in leaves of Elaeocarpus (Elaeo- the prothorax and elytra of some species of Ulomas- carpaceae) and Platynotocis bicaudatus Zimmerman cus , a myrmecophilous or termitophilous habit has (the species also undescribed, a nomen nudum ) in been surmised for the group (Marshall 1928; Ghes- leaves of Polyosma cunninghamii (Escalloniaceae) and quiè re 1932), but the fl at, long, strong body shape Peliocis lives on Allocasuarina (Casuarinaceae). Other and the occurrence on reeds of Cratopechus suggest than probably Peliocis , the larvae of all genera may that all genera probably live on large grasses or per- be leaf miners. haps palms, where they may hide or push into tight Genera of Uncertain Placement ( Incertae Sedis ) spaces between leaves and stems. The very long and (Fig. 3.7.10.8 L ). A number of genera cannot complex ovipositor also indicates that the eggs may be currently placed in any of the above tribes, be laid into deep, narrow spaces, perhaps covered although the list is not as long as that given by with a secretion from the tergal pouch. Alonso-Zarazaga & Lyal (1999). The largest group Viticiini (Fig. 3.7.10.8 K). This group is based of such genera of uncertain placement occurs in on Viticis Lea, small weevils lacking onychia and Madagascar, owing both to their generally inad- distributed throughout the Pacifi c islands. The equate descriptions and to the fact that the Mada- genus was associated with the entimine tribe gascan weevil fauna has not been studied in recent Ottistirini by Lea (1930) and subsequent authors, times. Although these genera are mostly placed until Morimoto (1983) recognized that its mouth- into tribes in the catalogue of Alonso-Zarazaga parts are phanerognathous and proposed a sepa- & Lyal (1999) and tentatively left there in this rate tribe for it and a closely related genus, Tivicis work, their placements need scrutiny in nearly Morimoto, which has short onychia and basally all cases. Some genera in continental Africa are connate but apically divergent claws. Kojima & also currently unplaced, e.g., the closely related 336 Roberto Caldara, Nico M. Franz, and Rolf G. Oberprieler

South African genera Hypsomus Schoenherr and Stals (Plant Protection Research Institute, Preto- Pachytrichus Schoenherr (Alonso-Zarazaga & Lyal ria, South Africa), Frank W. Pelsue, Jr. (Corona, 2002; Oberprieler 2010), the former apparently CA, USA), Andrew Walker (CSIRO Ecosystem Sci- associated with grasses (one species introduced ences, Canberra, Australia). Natalie Banks and in Western Australia, often found there in wheat Debbie Jennings (CSIRO Ecosystem Sciences) and fi elds) and the latter with Moraea (Iridaceae). Valter Fogato (Milan) are thanked for ably pre- Also of unclear affi nity are the South African paring the illustrations and Adriana Marvaldi for genera Sidomenia , and Stenotypus Marshall, both reviewing the manuscript. described in Erirhininae but transferred to Cur- culioninae incertae sedis by Alonso-Zarazaga & Lyal (2002), the former an apparent inquiline in cecidomyid galls on Aspalathus (Fabaceae) (S. Literature Neser, personal observation) and the latter breed- Ahmad, M. & Burke, H. R. (1972): Larvae of the weevil ing in stems of Arctotheca and Senecio (Asteraceae) tribe Anthonomini (Coleoptera: Curculionidae). – (R. Oberprieler, personal observation). The clas- Miscellaneous Publications of the Entomological Society of sifi catory position of the Indian Teluropus ballardi America 8: 33 – 81. Marshall, reported from leaves of jackfruit (Arto- Albach, D. C., Meudi, H. M. & Oxelman, B. (2005): carpus, Moraceae) (Marshall 1917; Jha & Sen-Sarma Piecing together the “ new ” Plantaginaceae. – Amer- 2008) but intercepted in the United Kingdom in ican Journal of Botany 92: 297 – 315. association with fruits of Syzygium jambos (Lyal Alonso-Zarazaga, M. A. (2002): Lista preliminar de los 1996), is also unclear; it certainly does not belong Coleoptera Curculionoidea del á rea Ibero-Balear, in Molytinae: Metatygini . In the New World, the con descripti ó n de Melicius gen. nov. y nuevas citas. important genera of uncertain affi nity are Mac- – Bolet í n de la Sociedad Entomol ó gica Aragonesa 31: rorhoptus, which appears to be oligophagous on 9 – 33. Malvaceae (Burke 1973; Anderson 1993) and was – ( 2005): Diagnoses preliminares de nuevos t á xones associated with the prionobrachiine genus Che- de Curculionidae (Coleoptera). – Bolet í n de la Socie- lotonyx by Anderson (1993), and a group of Chilean dad Entomol ó gica Aragonesa 37: 89 – 93. genera mostly associated with Nothofagus (Noth- – (2007): Acalyptini C. G. Thomson 1859, correct ofagaceae) – Epaetius, Epembates, Nothofaginoides, name for a tribe in Curculioninae (Coleoptera: Nothofaginus , and Wittmerius, all described by Curculionidae) and a comment on nomenclatural Kuschel (1952) – as well as Malaiserhinus Kuschel, terminology. – Coleopterists Bulletin 61 (4): 559. Alonso-Zarazaga, M. A. & Lyal, C. H. C. (1999): A World associated with Kageneckia (Rosaceae). Regarded Catalogue of Families and Genera of Curculionoidea (Insecta: as a “ special group of Curculioninae ” by Kus- Coleoptera) (Excepting Scolytidae and Platypod idae) . 315 chel (1987), these genera may represent a distinct pp. Entomopraxis S.C.P. Edition, Barcelona. tribe. The classifi cation of the South American – ( 2002): Addenda and corrigenda to “ A World Cata- fauna of Curculioninae generally needs attention, logue of Families and Genera of Curculionoidea as nearly all of its typical tribes are poorly delin- (Insecta: Coleoptera) ” . – Zootaxa 63: 1 – 7. eated. In the Palaearctic region, the main genera Anderson, D. M. (1962): The weevil genus Smicronyx in need of placement are Aubeonymus and Pachyty- in America north of Mexico (Coleoptera: Curcu- chius (Fig. 3.7.10.8 L). Aubeonymus mariaefranciscae lionidae). – Proceedings of the United States National Roudier is considered a pest of Beta vulgaris (Ama- Museum 113 (3456): 185 – 372. ranthaceae) in Spain, adults and larvae feeding on – ( 1972): Dodder weevils in simultaneous associa- beetroot and sugar beet (Marco et al. 1998), while tion with parasitic plants and their hosts. – Science Pachytychius lives on Fabaceae (such as Astragalus 168: 132 – 133. and Sarothamnus ) and Poaceae (Caldara 1978), – (1973): Keys to larvae and pupae of the Gymnetr- with P. avulsus Faust able to cause serious damage inae of America north of Mexico (Coleoptera: Cur- to wheat in the eastern Mediterranean region culionidae). – Proceedings of the Entomological Society (Martelli et al. 1960). of Washington 75: 133 – 140. Anderson, D. M. & Cox, M. L. (1997): Smicronyx species (Coleoptera: Curculionidae), economically impor- tant seed predators of witchweeds (Striga spp.) Acknowledgments (Scrophulariaceae) in sub-Saharan Africa. – Bulletin of Entomological Research 87: 3 – 17. We are very grateful to the following colleagues Anderson, D. M., Korotyaev, B. A. & Lingafelter, S. for various taxonomic and biological information W. (2006): Discovery of a new species of Smicronyx and many insightful comments on this diffi cult Schoenherr (Coleoptera: Curculionidae). – Coleo- group of weevils: Horace Burke (Texas A&M Uni- pterists Bulletin 60: 243 – 251. versity, College Station, TX, USA), Hiroaki Kojima Anderson, R. S. (1989): Revision of the subfamily (Tokyo University of Agriculture, Tokyo, Japan), Rhynchaeninae in North America (Coleoptera: Willy Kuschel (Auckland, New Zealand), Chris Curculionidae). – Transactions of the American Ento- Lyal (The Natural History Museum, London, mological Society 115: 207 – 312. United Kingdom), Adriana Marvaldi (CONICET, – ( 1993): Weevils and plants: phylogenetic versus Mendoza, Argentina), Stefan Neser and Riaan ecological mediation of evolution of host plant Curculioninae Latreille, 1802 337

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