692 · Family 125. Nemonychidae

Superfamily CURCULIONOIDEA

125. NEMONYCHIDAE Bedel 1882

by Robert S. Anderson

Family common name: The pine flower snout beetles

mong the weevils, these rarely collected beetles are easily recognized by their straight antennae, and elongate rostrum combined with the presence of a distinct labrum. Adults are found in association with the male pollen- Abearing flowers of Pinus species. Description (based on ing four pairs of setae. Antenna of a single membranous article Lawrence 1982). Shape elon- bearing an accessory appendage. Mandible with two apical teeth, gate, slightly convex; length an obtuse protuberance on cutting edge, a distinctly produced 3.0-5.5 mm; color pale brown molar area with a flattened grinding surface, and one pair of setae. to black; vestiture of fine short Hypopharyngeal bracon present. Maxillary palp with three ar- to moderately long appressed ticles, palpiger present or absent. Labial palp of two articles. or suberect pubescence. Ros- Premental sclerite present, may be divided medially. Thorax with trum moderately to very long pronotal sclerite transverse, lightly pigmented or unpigmented, and mostly narrow. Antennae sparsely covered with setae. Legs very small, subconical, of two or straight, ending in a weak, three segments, with or without a terminal claw. Abdomen with loose club of three articles; an- first eight segments with two dorsal folds and bearing annular or tennal insertions lateral at the bicameral spiracles. Anal opening terminal. middle or near the apex of the Pupae are undescribed. rostrum. Labrum distinct, not Habits and habitats. These beetles are rarely collected, likely fused with clypeus. Mandibles because of their specialized habits and life history. In North with a small but distinct mola. America, adults are found on male flowers of several pine species Maxillae with separate galea and very early in the season, often while snow is still on the ground. lacinia and maxillary palps flex- Adults feed on pollen. After mating, females lay eggs in the ible. Labial palps attached ven- flowers where the larvae also feed on pollen. Mature larvae drop FIGURE 1.125. Cimberis compta trally near the base of the from the flowers to the ground and pupate in the soil. Pupation (LeConte) (from Bright 1993, prementum. Gular sutures can take from a few months to two years. In some instances reproduced with the permission of well-developed and separate. larvae have been observed feeding on plant parts other than the Minister of Public Works and Proventriculus lacks sclerotized pollen (Thomas and Herdy 1961). In other areas of the world Government Services, 2001) plates. Procoxae contiguous gymnosperms such as Araucariaceae and Podocarpaceae, and some and the procoxal cavities nar- primitive angiosperms (Fagaceae and Ranunculacaeae) also serve rowly closed posteriorly. Mesocoxal cavities either not closed lat- as host plants. Whereas pines are the only documented hosts in erally or narrowly so. Elytra without an inner subcostal flange. North America, other genera of conifers may serve as hosts. Hind wing mostly with four anal veins or fewer. Tarsal claws of Status of the classification. This family is bipolar in distri- some cleft. Visible sternites of the abdomen are free; pygydium bution with approximately similar numbers of taxa found in the concealed by the elytra. Tegmen simple or bilobed apically and the southern temperate zones of South America, New Zealand and median lobe with a distinct dorsal plate. Australia, and in the northern Holarctic Region. Whereas the Eggs are undescribed. family was once considered to be absent in tropical areas, a few Larvae (based on Anderson 1991) when mature about 4.0- species have recently been collected in Panama and Venezuela on 4.8 mm in length, of moderate thickness throughout length, Podocarpus and appear to represent an undescribed genus. The strongly “C”-shaped. Body white, covered with mixture of long North American fauna has recently been revised (Kuschel 1989) and short setae. Minute legs present on thorax. Head hypogna- and is well-known. A catalog of the North American species was thous, rounded at sides, pigmented, with few to many setae on prepared by Hamilton (1994). frons and epicranium. Frontal sutures complete, reaching articu- Distribution. There are 5 genera and 15 species in North lating membrane of mandible. Clypeus not distinguishable from America. Two additional species in the genus Atopomacer are known frons and incompletely separated from labrum. One pair of ante- from Pinus at high elevations in far northern Mexico. North rior stemmata. Labrum short, the anterior margin rounded, bear- American nemonychid species are generally distributed in the Family 125. Nemonychidae · 693

5 23 4 6

78 91011

FIGURES 2.125-11.125. 2. Atopomacer orites Kuschel 1989, tarsus; 3. Cimberis elongata (LeConte 1876), tarsus; 4. Atopomacer orites Kuschel 1989, tarsal claw; 5. Cimberis turbans Kuschel 1989, tarsal claw; 6. Lecontellus pinicola Kuschel 1989, rostrum apex; 7. Pityomacer carmelites Kuschel 1989, rostrum apex; 8. Cimberis decipiens Kuschel 1989, rostrum apex; 9. Acromacer bombifrons (LeConte 1876), rostrum apex; 10. Pityomacer pix Kuschel 1989, head; 11. Acromacer bombifrons (LeConte 1876), head. western montane and boreal regions and likely occur anywhere 4(3). Labrum trapezoidal or broadly rounded at apex, with where pines are present. six peg-like apical setae in addition to three pairs of dorsal setae (Fig. 7); base of rostrum moder- ately saddled against obliquely rising frons (Fig. KEY TO THE NEARCTIC GENERA 10) in dorsal view, with 1-3 low carinae flanked with rows of confluent punctures; females with 1. Second tarsomere truncate at middle, not projected setiferous patches mostly present on one or two over base of third (Fig. 2); hind tibia with single ventrites ...... Pityomacer apical spur; tarsal claw with broad basal flange — Labrum triangular, pointed at apex, without any peg- (Fig. 4); elytra with punctures arranged into indis- like setae in addition to three pairs of dorsal se- tinct striae (Rhinorhynchinae)...... Atopomacer tae (Fig. 9); base of rostrum very deeply saddled — Second tarsomere lobed at middle, projected over against vertically rising frons (Fig. 11) in dorsal base of third (Fig. 3); hind tibia with two apical view, smooth or sparsely punctate, not carinate; spurs; tarsal claw simple, lacking broad basal females without setiferous patches on ventrites flange (Fig. 5); elytra with punctures not arranged ...... Acromacer into striae (Cimberidinae) ...... 2

2(1). Mandibles inserted dorsally, the sockets fully ex- CLASSIFICATION OF THE NEARCTIC GENERA posed in dorsal view (Fig. 6); in lateral view, man- dibles directed obliquely downwards in relation to rostral plane; antennae situated at middle or Nemonychidae Bedel 1882 after middle of rostrum, as distant from mandibu- lar sockets as combined length of first three ar- Cimberidinae Gozis 1882 ticles (Doydirhynchini) ...... Lecontellus — Mandibles inserted laterally, the sockets only par- tially exposed in dorsal view (Figs. 7-9); in lateral Cimberidini Gozis 1882 view, mandibles continuous with rostral plane; antennae situated at middle or before middle of Acromacer Kuschel 1989, 1 sp., A. bombifrons (LeConte 1876), Brit- rostrum, distinctly closer to mandibular sockets than combined length of first three articles ish Columbia, Alberta, Washington, Oregon, California, Idaho (Cimberidini) ...... 3 and Nevada. Adults collected on Pinus species.

3(2). Mandibles evenly curved on outer margin, each Cimberis Gozis 1881, 7 spp., generally distributed. Adults col- armed with well-developed tooth on inner margin (Fig. 8) ...... Cimberis lected on Pinus species. — Mandibles angulate on outer margin, unarmed on inner margin (Figs. 7, 9) ...... 4 694 · Family 125. Nemonychidae

Pityomacer Kuschel 1989, 3 spp., British Columbia, Alberta, Wash- BIBLIOGRAPHY ington, Oregon, California and Montana. Adults collected on Pinus species. ANDERSON, D. M. 1991. Nemonychidae (Curculionoidea). Pp. 585-586. In: F. W. Stehr, ed. Immature Insects. Volume 2. Doydirhynchini Pierce 1916 Kendall/Hunt. Dubuque, Iowa. BRIGHT, D. E. 1993. The Insects and Arachnids of Canada, Part. Lecontellus Kuschel 1989, 3 spp., Washington, Oregon, California 21. The Weevils of Canada and Alaska: Volume 1. Centre for and Nevada. Adults collected on Pinus species. Land and Biological Research. Ottawa, 217 pp. HAMILTON, R. W. 1994. A catalog of the Coleoptera of America Rhinorhynchinae Voss 1922 north of Mexico. Family: Nemonychidae. USDA Agriculture Handbook 529-134, x + 8 pp. Rhinorhynchini Voss 1922 KUSCHEL, G. 1989. The Nearctic Nemonychidae (Coleoptera: Curculionoidea). Entomologica Scandinavica, 20: 121-171. Atopomacer Kuschel 1989, 1 sp., A. ites Kuschel 1989, Colorado LAWRENCE, J. L. 1982. Coleoptera. Pp. 482-553. In: S. P. Parker, and Arizona. Adults collected on Pinus species apparently at high ed. Synopsis and Classification of Living Organisms. Volume elevations. 2. McGraw Hill. New York. THOMAS, J. B. and H. HERDY. 1961. A note on the life history of Cimberis elongatus (LeC.) (Coleoptera: Anthribidae). Cana- dian Entomologist, 93: 406-408. Family 126. Anthribidae · 695

126. ANTHRIBIDAE Billberg 1820

by Barry D. Valentine Family common name: The fungus weevils

Family synonyms: Anthotribidae Gemminger and Harold 1872; Choragidae Kirby 1819; Platyrrhinidae Everts 1903; Platystomidae Pierce 1916; Platystomoidea Pierce 1916.

hese primitive weevils greatly resemble the snout beetles but the beak is broad, the antennae are not geniculate, the pygidium is exposed, only the third tarsomere is spongy-pubescent beneath, the pronotal pubescence is Tdirected anteriad, and the elytra usually have an abbreviated scutellar stria. Description: Shape un- mostly trapezoidal, apex obliquely truncate, rarely somewhat pro- usually diverse; elongate and duced over the head, base truncate with an antebasal transverse depressed, to oval, convex, and ridge, this mostly turned forward at the sides forming a short or mite-like, mostly more or less more rarely complete side margin (called the lateral carina), the elongate and convex above; transverse ridge is always antebasal but it is referred to as basal in length 0.4 to 40 mm, in U.S. those species in which the surface behind it is more or less vertical; 0.4 to 16 mm; vestiture rarely surface smooth, punctate, reticulate, rugose, or tuberculate; pleu- absent, of hair-like scales ral region broad, supra-coxal sutures well developed; prosternum mostly in mixed shades of short to long in front of coxae, the intercoxal process narrow white, gray, straw, brown or (rarely broader than a coxa); the coxal cavities closed behind. Me- black, in the tropics some with sosternum short, the process separating the mesocoxae ex- bright pink, red, yellow, or tremely variable in shape and proportions. Metasternum mostly green. In collections, species are long, in some so short that meso- and metacoxae are barely sepa- FIGURE 1.126. Toxonotus cornutus often confused with ceram- rate, a transverse groove or suture near posterior margin, and in (Say) bycids, chrysomelids, ciids, many a midventral longitudinal depression or groove. Legs ro- scolytids, and mites. bust to slender; trochantins not exposed; procoxae globular, Head large, rarely retractile into prothorax; if rostrum present mostly protruding, not touching or only barely so, and grooved mostly broad and flattened; surface mostly smooth on vertex, on their inner faces for reception of the intercoxal process of sculpture often progressively heavier on frons and rostral area. prosternum; mesocoxae globular, separate; metacoxae elongate Antennae with eleven antennomeres (rarely nine or ten), not gen- and transverse (except Sicanthus Valentine and Cisanthribus iculate, most with an apical club of three antennomeres, club Zimmerman where they are globular or broadly tear-drop shaped), faint or absent in species with elongate antennae, and involving separate, not reaching elytral margin; trochanters moderate with from two to eight antennomeres in some tropical species; in- the apices oblique; femora swollen postmedially, mostly without serted either on the lateral or ventral surface of the rostrum, or on teeth or spines; tibiae slender, without movable spurs, but with the face between or below the eyes. Labrum distinct, mostly semi- an apical hook or tooth in males of some genera; tarsal formula circular, setose; mandibles unusually large for a weevil, stout, 5-5-5, apparently 4-4-4, the second apically emarginate, the third curved, upper surface flattened and with a setose groove, the deeply bilobed and tomentose beneath, partially embraced by the apices acute, mostly with a postmedian tooth; maxillary palpi second, and rarely with the lobes fused, the fourth very small and slender, with four palpomeres, the first very short, the fourth invisible in dorsal view, the fifth elongate, with claws mostly long; gular sclerite and sutures invisible; postmentum (fused toothed or cleft. Scutellum small, varied in shape, hidden in some mentum and submentum) large, deeply emarginate in front, flightless species. Elytral apices always locking into a longitudinal heavily sclerotized; ligula large, corneous, mostly notched to deeply groove in pygidial base, pygidial disc always partially exposed in emarginate in front, rarely entire; labial palpi with three palpomeres, posterior view but may be invisible from above; normally 10 the second shortest, both pairs of palpi elongate, cylindrical, flex- elytral striae or groups of punctures plus an abbreviated scutellar ible, and with acute (very rarely truncate) apices. Eyes dorsal, dorso- row, rarely 12, 14 or 19-21 rows of punctures; epipleural fold lateral, or lateral, mostly large, ranging from entire to deeply emar- present, narrowed or obsolete apically. Wing venation and fold- ginate, the facets very variable in size and number, and without ing pattern approach that of the Chrysomelidae. intermixed setae. Abdomen with five visible sterna, the four sutures entire, Pronotum apex narrower than the maximum head width fused, and immovable (rarely the fourth suture flexible); surface (except those choragine genera with retractile heads); some exotic without coarse sculpturing. Male genitalia with the penis mostly genera with the eyes out on broad stalks; base mostly nearly as slender, curved, with a movable dorsal plate which when raised wide as the elytra, constricted or produced laterad in some; shape gives an open “bird’s beak” appearance, with paired slender basal 696 · Family 126. Anthribidae struts; parameres and pars basalis fused, the latter with apex mostly and T. lepidus Valentine on fungus-infested sagebrush. The hosts hooded, rarely flattened, the ventral struts fused into a single of T. ornatus (Schaeffer), T. strigosus (Jordan), and T. nigromaculatus median structure. Female genitalia sclerotized, the valvifers with (Schaeffer) are not known. Sweeping bushy, overgrown areas will long apodemes, the coxites apically toothed or ridged, the styli sometimes reveal species of Ormiscus or Eusphyrus, especially in lateral, the spermatheca C-shaped. early spring, but beating is better for these and all other members Larvae are crescent-shaped, subcylindrical, fleshy, widest in of our fauna except species of Euparius, which occur on polypore the mid-abdominal region; size 4 to 12 mm in length (the major- fungi. Larvae of all our species feed on vegetation; they can be ity still undescribed); body with lateral fleshy protuberances and a found in the twigs and branches of trees, in hard or polypore few to many short or long setae scattered over various segments fungi, or under bark of dead or dying trees. One species, the and the head; color near white. Head exserted, hypognathous, coffee bean weevil, Araecerus fasciculatus (DeGeer 1775), lives in setiferous, or rarely retracted into the prothorax, narrower than seeds and all sorts of dried plant materials from banana flour to prothorax, with epicranial suture present. Antennae reduced, one- strychnine. A related species in Australia and New Zealand occurs segmented or absent. Clypeus transverse; labrum setiferous; on dried fruit, especially apples. Our introduced species of the mandibles robust, with or without molar areas, bi- or tri-dentate; European genus Anthribus (formerly Brachytarsus) have larvae maxillae with cardo, stipes, three-segmented palpi, the galea that feed on scale insects of the subfamily Lecaniinae; the female setiferous, lacinia acute and inconspicuous; labium with lays each of her eggs under a female scale that has laid her eggs, submentum, mentum, ligula, and one-segmented palpi. Ocelli the weevil larvae then develop exclusively at the expense of the absent. Thorax with legs absent or if present, l-, 2-, or 3-seg- eggs of the scale insect. This is a startling deviation from the mented, without tarsunguli. Abdomen nine-segmented, with phytophagous habits of the Rhynchophora. two or three plicae per segment, the ninth segment smaller than Status of the classification. This family has been subject the eighth. Spiracles annular or annular-biforous. Urogomphi to considerable nomenclatorial confusion and lack of studies of absent. Anderson (1947) provides a partial key. higher taxa beyond all proportions consistent with the rest of the Habits and habitats. Distributions and biologies of Nearc- order. The genera need considerable study. The world catalogues tic species are summarized by B. D. Valentine (1999). In general, of Wolfrum (1929, 1953) need many additions and revisions. adults may be found on the larval food plants. Those whose The Nearctic fauna is summarized by Valentine (1999) who pro- larvae feed in the stems or receptacles of various weeds appear to vides a synonymic checklist and keys to most genera. Other basic feed upon the pollen of the same plants; species with fungivorous references are LeConte (1876), Blatchley and Leng (1916), Pierce larvae mostly are found feeding on the surface of the same fungi; (1930), Ting (1936), and Valentine (1960, 1971, 1972, 1991). those with wood-boring larvae occur on dead or dying tree trunks Distribution. There are about 360 genera and 4,000 species or branches, and in some cases at least feed upon bark. Adults of known from all regions; 88 described and 32 undescribed species the majority of anthribid species in the United States are rare in occur in North America. Of 30 North American genera, Araecerus museum collections. The best all-around collecting technique is is cosmopolitan; Choragus, Eurymycter, Gonotropis, Allandrus, and beating dead or diseased branches, clumps of dead twigs or leaves, Trigonorhinus are holarctic (although the latter extends south to or tangles of dead vines. The beating cloth should be as close Argentina); Sicanthus is endemic but probably neotropical; under the plants as possible and must be examined at once, for Euxenulus, Phoenicobiella, and Araeoderes appear to be North Ameri- anthribids recover and fly off much more rapidly than other can but are obviously related to Latin American groups; and all weevils. Some of the choragine genera complicate things by being remaining genera are Neotropical, forming the northern fringe of active jumpers, and are as a result exasperatingly difficult to catch. an extensive Antillean, Central, or South American fauna. Another technique is to range the woods examining the trunks of trees, especially the thin-barked ones, for signs of infirmity, and also all dead branches both above and on the ground. Look KEY TO THE NEARCTIC TRIBES AND GENERA in particular for smooth gray or black patches flush with the wood surface, or small, protruding, black, brown or reddish 1. Antennae inserted on anterior surface of rostrum lumps. These are pyrenomycete fungi, often in the genera or head (Fig. 2); antennal club never with four antennomeres (Choraginae) ...... 2 Hypoxylon and Biscogniauxia, which are ignored by most collec- — Antennae inserted on lateral or ventral surface of tors; they are very hard and when cut with a knife have a dry rostrum (Fig. 3); or antennal club sometimes with crumbly or charcoal-like consistency. All our species of Eurymycter, four antennomeres. (Anthribinae) ...... 10 Piesocorynus, Choragus, and Euxenus are associated with these 2(1). Eyes rounded, upper edges not closer together than growths. Sweeping weedy fields usually produces only lower ...... 3 Trigonorhinus spp., the species T. tomentosus (Say) on common — Eyes elongate-oval, upper ends closer to each other ragweed, T. limbatus (Say) and T. griseus (LeConte) on sneeze- or than lower (Choragini) ...... 7 bitter-weed and other composites, T. alternatus (Say) on 3(2). Hind coxae elongate-transverse, almost reaching fungus-infested morning glory, T. sticticus (Boheman) on smutty elytral edge (Araecerini) ...... 4 grasses and corn, T. rotundatus (LeConte) on smutty grass — Hind coxae globular or short tear-drop shaped (Andropogon sp.) and ferns, and the western T. annulatus (Carr) (Cisanthribini) ...... Sicanthus Family 126. Anthribidae · 697

4(3). Head not retractile into prothorax, the eyes too wide; elytra with a scutellar plus 10 striae or rows of punctures ...... 5 — Head capable of being retracted into prothorax past the eyes (Fig. 2); elytra with 12 or more rows of punctures ...... 6 234 5(4). Lateral prothoracic carina present; transverse pronotal carina basal ...... Araecerus FIGURES 2.126-4.126. 2. Euxenus sp., head, anterior view; 3. — Lateral prothoracic carina absent; transverse Euparius marmoreus (Olivier), head, dorsal view; 4. Gymnognathus sp., pronotal carina antebasal ...... Neoxenus scutellum, dorsal view.

6(4). Lateral prothoracic carina double, not upturned at — White pubescence, if present, confined to spots, if apex; elytra with 12-14 rows of punctures ...... conspicuous, pronotum will have the disc con- ...... Habroxenus cave with a central swelling, and elytra will have — Lateral prothoracic carina single, mostly with apex multiple tufts of erect pubescence ...... 17 upturned; elytra with 19-21 rows of punctures ...... Acaromimus 16(15). Pronotum with a smooth, longitudinal, shallow groove from central pit to antebasal carina; elytral 7(2). Head not retractile into prothorax, the eyes too wide; pale patch antemedian and not reaching side pronotum with a raised reticulum forming a honey- margins ...... Gonotropis comb pattern ...... 8 — Pronotum with a sinuous, transverse, median — Head retractile into prothorax past the eyes; groove; elytral pale band postmedian and reach- pronotum punctate, not reticulate ...... 9 ing the side margins ...... Eurymycter

8(7). Antennae with 11 antennomeres ...... Choragus 17(15). Antennal club with 4 antennomeres; rostrum thick, — Antennae with 9 or 10 antennomeres ...... width at apex less than twice depth; scrobes ...... Pseudochoragus dorso-lateral, interscrobal distance mostly less than interoccular distance (Discotenini) ...... 9(7). Elytra with 10 rows or fields of punctures, plus a ...... Discotenes short scutellar row ...... Euxenus — Antennal club with 1 or 3 antennomeres; rostrum — Elytra with 11 to 15 rows of punctures, one or more depressed apically, width at apex more than mostly incomplete ...... Euxenulus twice depth; interscrobal distance more than interoccular distance ...... 18 10(1). Each elytron partially enclosing anterior part of scutellum, resulting scutellar notch shaped like 18(17). Eyes finely faceted, more than 26 rows across maxi- an inverted omega (Fig. 4); eyes large, entire, very mum width ...... 19 finely faceted (Gymnognathini) ...... — Eyes coarsely faceted, 26 or fewer rows across ...... Gymnognathus maximum width (Piesocorynini) ...... 23 — Scutellar notch forming a simple V or U; eyes as above, or small, or notched, or coarsely faceted 19(18). Sides of rostral apex abruptly widened to cover ...... 11 the laterally protruding mandibular bases; our species with a small post-ocular tooth on apex of 11(10). Rostrum with a dorsal, median carina interrupted at prothorax (Ischnocerini) ...... Ischnocerus base by an abrupt, small, very deep pit or trans- — Sides of rostral apex with mandibular sheaths verse groove (Platystomini) ...... 12 weakly or not wider than rostral dorsum; no small — Rostrum carinate or not, at most with a basal punc- post-ocular teeth on prothoracic apex ...... 20 ture ...... 13 20(19). Merger of rostrum with venter of head capsule form- 12(11). Lobes of all third tarsomeres separate .. Toxonotus ing a broad curve; rostrum long, flattened, and — Lobes of third tarsomeres fused..... Phoenicobiella apically flared...... 21 — Merger of rostrum with venter of head capsule indi- 13(11). Mandibles with strongly toothed ventral cutting cated by a transverse groove or abrupt angle; edge (Fig. 3), as well as normal dorsal edge rostrum shorter, not apically flared (Platyrhinini) (Cratoparini) ...... Euparius ...... 22 — Mandibles without a toothed ventral cutting edge ...... 14 21(20). Antennae with whorls of long, erect setae; length (head excluded) more than 5 mm (Stenocerini) . 14(13). Eye entire, or truncate, or faintly sinuate on ante- ...... Stenocerus rior face ...... 15 — Antennae without whorls of long erect setae; length — Eye strongly notched or emarginate on anterior face (head excluded) less than 4 mm (Allandrini) ...... (nearest insertions of antennae) ...... 24 ...... Allandrus

15(14). Entire face with conspicuous white pubescence; 22(20). Face with a pair of abrupt depressions between the center of pronotum with a pit or groove; our spe- upper ends of the eyes; lateral prothoracic ca- cies with a conspicuous patch or band of white rina with an apical tubercle ...... Trachytropis crossing the suture (Tropiderini) ...... 16 — Face with at most a weak concave area at rostral base; lateral prothoracic carina not toothed at apex...... Goniocloeus 698 · Family 126. Anthribidae

23(18). Eyes with 14 or more rows of facets across maxi- Araeosarus Walker 1859 mum width ...... Piesocorynus Doticus Pascoe 1882 — Eyes with 12 or fewer rows of facets across maxi- mum width ...... Brachycorynus Metadoticus Olliff 1890

24(14). All third tarsomeres with the lobes fused down the Neoxenus Valentine 1999, 1 sp., N. versicolor Valentine, 1999, Texas midline (Anthribini) ...... Anthribus to Panama; four others in Central America and Antilles. — All third tarsomeres with the lobes separate ..... 25

25(24). Rostrum quadrate, or with rounded apical angles . Habroxenus Valentine 1999, 1 sp., H. politus Valentine, 1999, Mary- ...... 26 land, Texas; four others in Central America and Antilles. — Rostrum, excluding mandibles, narrowed from base to apex; rostral apex with central third longer than sides (Trigonorhinini) ...... Trigonorhinus Acaromimus Jordan 1907, 1 sp., A. americanus (Motschoulsky 1873), Florida, Alabama, and Texas; six others in Central America and 26(25). Intercoxal process of mesosternum laterally Antilles. angulate, or swollen, or bent; scrobes sulciform, Acaropsis Jordan 1907 continued across the rostral sides and ending below the eyes (Basitropidini)...... 27 Xenorchestes Motschulsky 1873, not Wollaston 1854 — Intercoxal process of mesosternum simple; scrobes Euxenus Blatchley 1920, not LeConte 1876 foveiform, more or less rounded and not contin- ued transversly or ventrally around and under Cisanthribini Zimmerman 1994 the rostrum (Zygaenodini) ...... 28

27(26). Lateral prothoracic carina extending to apex where Sicanthus Valentine 1999, 1 sp., S. rhizophorae Valentine 1999, it is toothed; antennal scrobes strongly produced Florida keys. toward ventral midline ...... Eugonus — Lateral prothoracic carina not reaching anterior mar- gin, nor apically toothed, antennal scrobes widely Choragini Kirby 1819 separated on venter...... Phaenithon Choragus Kirby 1819, 6 spp., eastern United States; 45 others 28(26). Lateral prothoracic carina absent or if present not almost worldwide. reaching anterior margin ...... 29 — Lateral prothoracic carina extending to apex ...... Alticopus Villa and Villa 1833 ...... Araeoderes Pseudochoragus Petri 1912, 1 sp., P. nitens (LeConte 1884), Massa- 29(28). Transverse pronotal carina clearly antebasal, inca- chusetts and Oklahoma; another species in Europe. pable of contacting the elytral base; pronotal hind angles not projecting laterad of the humerae ... Choragus, of American authors, in part, not Kirby 1819 ...... Ormiscus Holostilpna, of American authors, not Jordan 1907 — Transverse pronotal carina subbasal or basal, ca- pable of contacting elytral base at some point; Euxenus LeConte 1876, 3 spp., eastern United States; at least six pronotal hind angles often projecting laterad of the humerae ...... Eusphyrus more in Central America and Antilles. Holostilpna Jordan 1907

CLASSIFICATION OF THE NEARCTIC GENERA Euxenulus Valentine 1960, 1 sp., E. piceus (LeConte 1878), south Florida; also three undescribed in Antilles. Suprageneric taxa are diagnosed in the generic key. NOTE: Many undescribed species are included in the comments on distribu- Anthribinae Billberg 1820 tion. As used below, the designation Neotropical includes Antillean species; Central and South America do not. Discotenini Lacordaire 1866

Anthribidae Billberg 1820 Discotenes Labram and Imhoff 1841, 2 spp., Texas; Arizona about 10 more south to Brazil. Choraginae Kirby 1819 Phanosolena Schaeffer 1904

Araecerini Lacordaire 1876 Ischnocerini Lacordaire 1866

Araecerus Schoenherr 1823, 2 introduced spp., including the cof- Ischnocerus Schoenherr 1839, 3 spp., Maryland to Texas; Arizona; fee bean weevil, A. fasciculatus (DeGeer 1775): Worldwide. Also 10 more in Neotropics. about 75 Indopacific species. Meconemus Labram and Imhoff 1839 Araeocerus Schoenherr 1839 Araeocorynus Jekel 1855 Family 126. Anthribidae · 699

Allandrini Pierce 1930 Toxonotus Lacordaire 1866, 6 spp., eastern United States to New Mexico and Arizona; about 35 others throughout the Neotropics. Allandrus LeConte 1876, 3 spp., United States except the desert Anthribus, of authors, in part, not Geoffroy 1762 southwest; also several in Palaearctic. Neanthribus Jordan 1906 Tropiderinus Reitter 1916 Pseudanthribus Pierce 1930

Stenocerini Kolbe 1897 Basitropidini Lacordaire 1866

Stenocerus Schoenherr 1826, 1 sp., S. longulus Jekel 1855, south Eugonus Schoenherr 1833, 1 sp., E. bicolor Valentine 1972, south- Texas to Brazil; seven more Mexico to Argentina. eastern Arizona; about 30 more in Central and South America. Schimatocheilus Fahraeus 1839 Gymnognathini Valentine 1960 Phaenisor Motschoulsky 1874

Gymnognathus Schoenherr 1826, 2 spp., south Texas; south Ari- Phaenithon Schoenherr 1823, 1 sp., P. platanum (Schaeffer 1906), zona; about 90 described and many undescribed Neotropical spe- southeastern Arizona; about 65 others in Central and South cies. America. Analotes Schoenherr 1839 Camaroderes Jekel 1855 Griburiosoma Schaeffer 1906 Tropiderini Lacordaire 1866 Zygaenodini Lacordaire 1866 Gonotropis Leconte 1876, 1 sp., G. gibbosus LeConte 1876, Canada south to Pennsylvania and Colorado; plus three Holarctic. Ormiscus G. R. Waterhouse 1845, 14 described and about 30 Tropideres, of European authors and Valentine 1960, in part, undescribed spp., eastern United States and desert southwest; not Schoenherr 1823 probably 200 more throughout the Neotropics, mostly undescribed. Eurymycter LeConte 1876, 3 spp., United States except the desert Entomops Lacordaire 1866 southwest. Hormiscus Gemminger and Harold 1872 Tropideres, of Valentine 1960, not Schoenherr 1923 Toxotropis LeConte 1876 Gonops LeConte 1876 Piesocorynini Valentine 1960 Eusphyrus LeConte 1876, 6 spp., eastern United States and desert Piesocorynus Dejean 1834, 5 spp., eastern United States; Arizona; southwest; 30 more in Central and South America. about 30 others in Neotropics and Antilles. Opisthotropis Hoffmann and Tempere 1954 Piezocorynus Schoenherr 1839 Camptotropis Jekel 1855 Araeoderes Schaeffer, 1906, 1 sp., A. texanus Schaeffer, 1906, southern Texas and southern Alabama. Brachycorynus Valentine 1999, 3 spp., eastern United States to Texas; a few others in Central America and Antilles. Trigonorhinini Valentine 1999 Tropideres, of LeConte 1876 and others, not Schoenherr1823 Brevibarra, of Valentine 1960, not Jordan 1906 Trigonorhinus Wollaston 1861, 11 spp., southern Canada, entire United States; seven others south to Argentina, and in the Pale- Platyrhinini Imhoff 1856 arctic Region. Brachytarsus, of many authors, not Schoenherr 1823 Goniocloeus Jordan 1904, 1 sp., G. bimaculatus (Olivier 1795), east- Trigonorrhinus Gemminger and Harold 1872 ern United States to Texas; 36 others south to Brazil. Anthribulus LeConte 1876 Tropideres, of various authors, in part, not Schoenherr 1823 Brachytarsoides Pierce 1930 Strabus Jekel 1860, not Gerstaecker 1860 [Curculionidae] Cratoparini LeConte 1876 Trachytropis Jordan 1904, 1 sp., T. arizonicus (Sleeper 1954), south- eastern Arizona; two others in Central America. Euparius Schoenherr 1823, 5 spp., eastern United States to Mon- tana and Arizona; at least 70 others in Central and South America, Platystomini Pierce 1916 and Japan to Australia. Cratoparis Dejean 1834 Phoenicobiella Cockerell 1906, 2 spp., South Carolina to Lousiana, Caccorhinus Sharp 1891 Florida and Bahamas; south Texas; a third species in Cuba. Phoenicobius LeConte 1876, not Moerch 1852 [Mollusca] 700 · Family 126. Anthribidae

Anthribini Billberg 1820 TING, P. C. 1936.The mouthparts of the coleopterous group Rhynchophora. Microentomology, 1: 93-114. Anthribus Geoffroy 1762, 2 adventive spp. from Europe, Atlantic VALENTINE, B. D. 1960. The genera of the weevil family coast and ?California. Anthribidae north of Mexico (Coleoptera). Transactions of Brachytarsus Schoenherr 1823 the American Entomological Society, 86: 41-85. Anthotribus Gemminger and Harold 1872 VALENTINE, B. D. 1971. Family Anthribidae. Pp. 243-248. In: Pseudobrachytarsus Pierce 1930 M. H. Hatch. Beetles of the Pacific Northwest, Part V. Univer- sity of Washington Publications in Biology, 16: i-xvi + 1-662. BIBLIOGRAPHY pl. I-LV. VALENTINE, B. D. 1972. Notes on anthribid weevils. III. New ANDERSON, W. H. 1947. Larvae of some genera of Anthribidae species and records primarily from Arizona. Coleopterists (Coleoptera). Annals of the Entomological Society of America, Bulletin, 26: 1-11, fig.1-4. 40: 489-517, pl. I-IV. VALENTINE, B. D. 1991. The Choragus-Holostilpna problem BLATCHLEY, W. S. and C. W. LENG. 1916. Rhynchophora or (Coleoptera: Anthribidae). Coleopterists Bulletin, 45: 301-307. Weevils of North Eastern America. The Nature Company. VALENTINE, B. D. 1998(1999). A review of Nearctic and some Indianapolis, 682 pp., fig. 1-155. related Anthribidae (Coleoptera). Insecta Mundi, 12: 251-296. LECONTE, J. L. 1876. The Rhynchophora of America, North of WOLFRUM, P. 1929. Anthribidae. Coleopterorum Catalogus, Mexico. Proceedings of the American Philosophical Society, 26(102): 1-145. XV (96): i-xvi + 1-455. WOLFRUM, P. 1953. Anthribidae. Coleopterorum Catalogus, PIERCE, W. D. 1930. Studies of the North American weevils of supplementum, 26(102): 1-63. the superfamily Platystomoidea. Proceedings of the United States National Museum, 77(no. 2840): 1-34, pls. 1-5. Family 127. Belidae · 701

127. BELIDAE Schönherr 1826

by Robert S. Anderson

Family common name: The cycad weevils

he only North American members of this family are odd weevils, recognized by their straight antennae, some- what truncated elytra mostly exposing the last one or two tergites, and the short stout legs with femora which Tare expanded in all legs, especially so in males. These weevils are only found in southern Florida where they are associated with native and adventive Zamia cycads. Description. (based on of Zamia (Cycadaceae). The biology has been well-studied by Lawrence 1982). Shape elon- Norstog and Fawcett (1989). Adult weevils swarm on male cones gate, slightly convex; length 3- of the cycads, where mating, feeding and oviposition occur. Lar- 6 mm; color pale to dark brown vae feed within the male cones. Weevils also visit female cones or black; vestiture of very fine but do not feed. Pollen transport to the female cones occurs short appressed pubescence. during these visits and these beetles (and their relatives elsewhere) Rostrum moderately to very may be obligate pollinators of the cycads. long and mostly narrow. An- Status of the classification. The family-level classification tennae straight, ending in a of these beetles is somewhat controversial. Crowson (1986) con- weak, loose club of three ar- sidered them as a distinct family, the Allocorynidae. Kuschel (1995) ticles; articles 9 and 10 with places them as Belidae, subfamily Oxycoryninae. The most recent deep apical pockets; antennal catalog (Alonso-Zarazaga and Lyal 1999), probably following insertions ventral at or near the Lawrence (1982) and Thompson (1992), places them as the sub- base of the rostrum. Labrum family Allocoryninae of the Oxycorynidae. Here they are placed as fused with clypeus. Labial palps Belidae, subfamily Allocoryninae. with two articles and attached Distribution. There is one genus with two species in North dorsally near the apex of the America. Both species occur only in Florida; one is adventive and prementum. Gular sutures one native. FIGURE 1.127. Rhopalotria slossonae (Schaeffer) (from Bright fused. Proventriculus lacks scle- 1993, reproduced with the rotized plates. Pronotum with CLASSIFICATION OF THE NEARCTIC GENERA permission of the Minister of complete lateral edges. Public Works and Government Procoxae contiguous and the Belidae Schönherr 1826 Services, 2001) procoxal cavities narrowly closed posteriorly. Elytra with- Allocoryninae Sharp 1890 out an inner subcostal flange. Hindwing with fewer than five anal veins. Tarsal article 2 broadly lobed as article 3; tarsal claws simple. Rhopalotria Chevrolat 1878, 2 species, R. mollis (Sharp 1890), Florida Visible sternites of the abdomen free; pygydium exposed by the (adventive on Zamia furfuracea L.) and R. slossonae (Schaeffer 1905), elytra. Tegmen truncate or slightly emarginate apically and the Florida (native on Zamia integrifolia L., coontie). median lobe with a distinct dorsal plate. Allocorynus Sharp 1890 Eggs are undescribed. Larvae (based in part on van Emden 1938, Lawrence 1982, BIBLIOGRAPHY Crowson 1986) short, broad, ventrally curved and subglabrous. Body lightly sclerotized. Legs absent. Head strongly retracted, ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL. 1999. A world endocarina indistinct, with very short antennae of one article and catalogue of families and genera of Curculionoidea (Insecta: no epipharyngeal rods. Frontal sutures incomplete, not reaching Coleoptera) (Excepting Scolytidae and Platypodidae). articulating membrane of mandible. Maxilla with palp with two Entomopraxis. Barcelona, Spain. articles, palpiger absent. Thoracic spiracles on mesothorax. Ab- CROWSON, R. A. 1986. On the systematic position of domen with segments with two dorsal folds. Anal opening ven- Allocoryninae. Coleopterists Bulletin, 40: 243-244. tral. EMDEN, F. VAN. 1938. On the taxonomy of Rhynchophora larvae Pupae are undescribed. (Coleoptera). Transactions of the Royal Entomological Soci- Habits and habitats. Within this family, only the genus ety of London, 87: 1-37. Rhopalotria occurs in North America. It is restricted to south Florida where two species are associated with native and adventive species 702 · Family 127. Belidae

KUSCHEL, G. 1995. A phylogenetic classification of NORSTOG, K. J. and P. K. S. FAWCETT. 1989. Insect-cycad Curculionoidea to families and subfamilies. Memoirs of the symbiosis and its relation to the pollination of Zamia furfuracea Entomological Society of Washington, 14: 5-33. (Zamiaceae) by Rhopalotria mollis (Curculionidae). American LAWRENCE, J. L. 1982. Coleoptera. Pp. 482-553. In: S. P. Parker, Journal of Botany, 76: 1380-1394. ed. Synopsis and Classification of Living Organisms. Volume THOMPSON, R. T. 1992. Observations on the morphology and 2. McGraw Hill. New York. classification of weevils (Coleoptera, Curculionoidea) with a key to major groups. Journal of Natural History, 26: 835-891. Family 128. Attelabidae · 703

128. ATTELABIDAE Billberg 1820 by Robert W. Hamilton

Family common names: The leaf rolling weevils, tooth-nose snout beetles, and thief weevils.

Family synonyms: Rhynchitidae Gistel 1856

he members of this family are considered to be primitive weevils based on the straight antennae (non-genicu- late), mouthpart structure, wing venation, etc. The family in the inclusive sense contains three subfamilies in TAmerica north of Mexico, Attelabinae, Rhynchitinae and Pterocolinae. In the Attelabinae (leaf rolling weevils), the body is more or less stout, the rostrum short and apically widened, the mandibles robust and toothed only on the inner margin, the front legs enlarged and the tarsal claws connate. In the Rhynchitinae (tooth nose snout beetles), the body is more or less elongate, the rostrum mostly narrow and elongate, the mandibles flat and toothed on the inner and outer margins, the legs subequal in size and the tarsal claws appendiculate. In the Pterocolinae (thief weevils), the body is short and compact, the short rostrum apically depressed and parallel-sided, the mandibles flat and toothed on the inner and outer margins (as in the Rhynchitinae), the middle and hind femora larger than the front femora, and the tarsal claws have a broad basal tooth. The Pterocolinae are a unique New World group that are clearly related to the Rhynchitinae although the rhynchitine link has not yet been determined.

Description: The follow- postmentum in some males with pair of acuminate ventral pro- ing adult characters have been jections; antennae inserted dorsolaterally, club 3-segmented, more given by Thompson (1992) or less compact, scrobe short, distinct, more or less vertical; man- and Kuschel (1995) to support dibles robust, toothed on inner apical margin, outer margin more the single family status of or less rounded; maxillary palps 4-segmented; labial palps indis- Attelabidae: maxillary palps 3 tinctly 1 or 2 segmented, located ventrally on prementum; elytra or 4 segmented (if 3 seg- more or less quadrate, not much longer than wide, humeri simple mented, elytral punctuation to strongly protuberant, scutellary striole present or absent, striae confused); tergite 8 concealed becoming less distinct posteriorly in many; ventral abdominal in both sexes; elytra separately sutures 1-3 more or less rigid; procoxae prominent, conical; pro- and broadly rounded at apices, thoracic legs enlarged, profemora distinctly swollen, armed or more or less exposing a py- unarmed; protibia longer, narrower and more arcuate in males; gidium (tergites 6 and 7); body tibial apices uni-uncinate in males and bi-uncinate in females; setose, without broad scales; tarsal claws connate. cuticle metallic or otherwise Larva with thoracic spiracle located in mesothorax or in in- brightly colored in many; elytra tersegmental fold between prothorax and mesothorax; with scutellary striole in many; prementum and mentum fused, equally sclerotized; maxillary abdominal ventrites decreasing palps 2-segmented; prodorsal fold more convex than postdorsal in length, first 3 or 4 fused; fold, postdorsal fold sloping posteriorly; abdominal tergum 9 at pygidium visible in both sexes; least twice as long as sternum 9; anus subterminal or ventral; male genitalia consisting of a cuticular asperities more or less inconspicuous. free pedon and tectum and a The subfamily Rhynchitinae is characterized by the follow- FIGURE 1.128. Merhynchites bicolor large tegmen (as in all groups ing: Adult with body more or less elongate, mostly with distinct (Fabricius) (from Bright 1993, with non-geniculate antennae); semi-erect to erect setae above, color variable; head mostly trian- reproduced with the permission of intersegmental membrane be- gular, widest at the base; eyes small to medium in size, rounded, the Minister of Public Works and tween sternites 8 and 9 partly longest diameter more or less horizonal; rostrum mostly slen- Government Services, 2001) or entirely sclerotized. der, parallel-sided and longer than the head, more or less weakly The subfamily Attelabinae is characterized by the following: arcuate in lateral view; antennae inserted laterally, club 3-segmented, Adult with body more or less short, stout, glabrous or with more or less loosely united, scrobes shallow, longitudinally ob- appressed pubescence above, color variable; head rectangular to lique, not well defined; mandibles flat, toothed on inner and weakly triangular, elongate in males of some; eyes large, mostly outer margins; maxillary palps 4 segmented; labial palps 2 or 3 reniform, longest diameter more or less vertical; rostrum short to segmented, inserted laterally or apically on prementum; elytra moderate in length, distinctly widened apically, weakly arcuate in elongate, mostly distinctly longer than wide, humeri simple, lateral view; lateral apical angle in some with tooth-like projection; scutellary striole present or absent, striae mostly distinct through- 704 · Family 128. Attelabidae out; ventral abdominal sutures distinct, suture between 1 and 2 ied. Comments on the biology of three North American species rigid; all legs more or less equally developed; tibial apices mostly are available. Packard (1890), Frost (1908), Blatchley and Leng with one or two small straight mucros; tarsal claws with long (1916), and Lutz (1935) commented on the biology of clawlike inner processes; in some inner processes shorter, wider Himatolabus pubescens (Say). Frost (1908) recorded a eulophid para- and more toothlike. site bred from Attelabus rhois Boheman (=H. pubescens). Loding Larva with thoracic spiracle located on postero-lateral por- (1945) commented on the biology of Homoeolabus analis (Illiger). tion of prothorax; prementum and mentum separate, mostly Murtfeldt (1872), Packard (1890), Girault (1904), and Edwards unequally sclerotized; maxillary palp 2 or 3 segmented; prodorsal (1949) treated aspects of the biology of Attelabus bipustulatus and postdorsal folds (segments 6-8) more or less subequal in Fabricius. Most of the information in these papers deals with convexity; tergum and sternum of abdominal segment 9 subequal taxonomy, host plants, nidus formation and distributions. in length; anus terminal or subventral, transverse x-shaped or Hopkins (1905) reported a trichogrammid egg parasite of A. simple transverse cleft; cuticular asperities more or less conspicu- bipustulatus. Van Emden (1938) included the larvae of H. pubescens, ous. Homoeolabus analis and Attelabus nigripes LeConte in a key to the The subfamily Pterocolinae is characterized by the following: genera of Attelabini. Also, he associated the niduses of these Adult with body robust and dorsally convex, with appressed to species, and A. bipustulatus as well, with specific host plants. Vogt semi-erect inconspicuous fine setae above, coloration metallic blu- (1992) discussed the leaf rolling behavior, host plants and asso- ish-green to bluish-black; head weakly triangular; eyes large, oval, ciated rhynchitid weevils (Pterocolinae) of attelabine weevils oc- posteriorly more protuberant, anteriorly emarginate; rostrum curring from Canada to the Republic of Panama. Hamilton(1998), short, as long as or shorter than the head, straight, in dorsal view in a revision of the New World Pterocolinae, described 15 new more or less parallel-sided throughout, apically depressed, with species from Central and South America and clarified many beard like ventral setosity (more setose in males); antennae short, attelabid-pterocoline-host plant associations initiated by Vogt. subequal to combined length of head and rostrum, inserted near Hamilton (1994) provided a summary of known North basal 1/4 under basirostral ridge; scrobe fossa like; club large, American rhynchitine biology. The larvae of these weevils de- strongly abrupt, compact; mandibles flat, toothed on inner and velop in living or dead leaves as leaf miners, in fruits, in cut outer margins; maxillary palps 4-segmented, labial palps 3-seg- flower heads, in cut terminal shoots, in terminal buds, in flower mented, inserted apically on prementum; pronotum with dis- buds, and in cut leaf primordia. The cuts are made by the adult tinct lateral carinae; elytra short, apices individually rounded; py- female with her mandibles after oviposition in the specific plant gidium and two complete abdominal terga visible; humeri simple; part. The flower heads, buds or leaf primordia are either com- scutellary striole present; striae mostly distinct throughout; inter- pletely cut from the plant or are partially cut and eventually drop vals more or less convex; all ventral abdominal sutures entire; to the ground. Information has been published on the life stages mesepimeral “side pieces” strongly developed, visible in dorsal or life cycles of only eight North American rhynchitine weevils - view between pronotum and elytra; front coxae small, globose, Haplorhynchites aeneus (Boheman), Merhynchites bicolor (Fabricius), distinctly separated; middle and hind coxae widely separated; Merhynchites wickhami (Cockerell), Eugnamptus angustatus (Herbst), middle and hind femora larger than front femora; tibial apices Auletobius cassandrae (LeConte), Temnocerus perplexus (Blatchley), with two blunt-tipped mucros; tarsal claws with broad basal tooth. Temnocerus naso (Casey) and Deporaus glastinus (LeConte). Boving Larva is undescribed. and Craighead (1931) provided the first basic illustrations of H. Attelabid eggs that have been examined are oval, creamy aeneus larvae. Hamilton (1973, 1981) provided detailed illustra- white to yellowish-white, shiny and without surface sculpture tions of life stages, host plants, and behavior for H. aeneus. (Balduf 1959; Hamilton 1980, 1983, 1994). Chittenden (1901), Dickerson (1910), Blatchley and Leng (1916), Only a few North American pupae in the subfamily Ewing (1915), and Essig (1958) comment on the biology of the Rhynchitinae have been described (Hamilton 1980, 1983; eastern rose curculio, M. bicolor. Balduf (1959) published the most Hamilton and Kuritsky 1981). They possess taxonomically sig- comprehensive study on the biology of M. bicolor to date. nificant tuberculate setae and posterior processes associated with Hamilton and Kuritsky (1981) commented on the life cycle of M. abdominal segment 9. bicolor and provided detailed descriptions of the larva and pupa. Habits and habitats. All known members of the subfamily Cooley (1903), Lovett (1915), Robertson (1923), and Hoerner Attelabinae are leaf rollers and apparently cause no serious dam- (1936) comment on the life cycle of the western rose curculio, M. age to their hosts although one Central American species of wickhami. Boving and Craighead (1931) illustrated what appears Hybolabus has been reported as a serious defoliator of its host to be the larva of E. angustatus but they did not rear it and they tree, Cariniana pyriformis Miers (Lecythidaceae). All known females determined it as Orsodacne sp. by the “method of elimination and lay their egg(s) on leaves that they prepare by biting and cutting locality.” Hamilton (1980) provided notes on the biology of E. with their mandibles. The leaves are then rolled into a more or angustatus (Herbst) and described and illustrated the larva and less barrel-shaped structure (nidus) that nourishes and protects pupa. Hamilton (1983) discussed the life cycles of A. cassandrae the developing larvae. The enlarged front legs and the uncinate and T. perplexus and provided detailed descriptions and illustra- tibial apices manipulate the leaf tissue during nidus formation. tions of their immature stages. Hamilton (1994) provided new Pupation occurs in the ground in the species that have been stud- Family 128. Attelabidae · 705 life cycle data for T. naso and D. glastinus on Quercus wislizenii A.D.&C. in the family Attelabidae but suggested a possible alternative in southern California. would be to establish a family Rhynchitidae for the Rhynchitinae Published biological information on other rhynchitid species is and Pterocolinae. Lawrence (1982) also placed these weevils in the lacking except for brief comments by Kissinger (1964), given him by family Attelabidae. O’Brien and Wibmer (1982), in an annotated Vogt, stating that Eugnamptus spp. mine the dead leaves of various checklist of North American weevils, recognized the families hardwoods and that females of Temnocerus aeratus (Say) lay eggs in Rhynchitidae and Attelabidae but Wibmer and O’Brien (1986), and cut terminal oak twigs in which the larvae subsequently develop. in their annotated checklist of South American weevils, regrouped Kissinger also indicated that, again according to Vogt, Involvulus hirtus these weevils into the single family Attelabidae. Thompson (1992) (Fabricius) has habits similar to T. aeratus.Van Emden (1938) in- using abdominal characters, etc. lumped the three subfamilies in cluded three North American species in a paper on the taxonomy of the family Attelabidae. More recently Kuschel (1995), in a phylo- Rhynchophora larvae. He provided key characters for the larvae of H. genetic approach, recognized only the family Attelabidae and Farrell aeneus, M. bicolor and Rhynchites velatus LeConte and included them in (1998), in a molecular approach, also grouped the rhynchitines a key to select species of Rhynchitini. He also associated these three and attelabines in the family Attelabidae. Hamilton (1969 to 1998) species with host plants and localities. Based on his host plant data, has consistently used the family Rhynchitidae for the Rhynchitinae he examined the larva of H. aeneus not Rhynchites (Involvulus) hirtus and Pterocolinae. Most recently, Alonso-Zarazaga and Lyal (1999) (Fabricius) as listed. have recognized the family Rhynchitidae in their world catalogue Haplorhynchites aeneus, M. bicolor and M. wickhami are the only of families and genera of Curculionoidea. North American rhynchitine species of known economic impor- The family name Attelabidae is credited to Billberg (1820). The tance. Haplorhynchites aeneus attacks commercially grown sunflow- genus Attelabus, on which the family name is based, was originally ers while M. bicolor and M. wickhami attack cultivated roses and described by Linnaeus (1758). Linnaeus included only A. coryli sometimes damage blackberry and raspberry. Comments on the Linnaeus in the genus and it was therefore the type by monotypy. economic importance of M. bicolor have been made by Harris Since the time of Linnaeus, many species from all over the world (1862), Chittenden (1901), Gates (1909), Dickerson (1910), have been added to the genus Attelabus. Olivier (1807) decided that Blatchley and Leng (1916) and Essig (1958). Cooley (1903), Lovett A. coryli was generically different from the other species that had been (1915), Robertson (1923), and Hoerner (1936), give information placed in Attelabus and described the new genus Apoderus with 13 on the economic importance and control of M. wickhami. Schulz species including A. coryli. Attelabus coryli is technically the type of and Lipp (1969) and McBride and Oseto (1978) commented on Attelabus but was not recognized by Olivier, or anyone else, until the status of damage to sunflowers by H. aeneus. Bedel (1888) pointed out that Attelabus should be used for those Vogt (1992) discussed the biology of the Pterocolinae and species placed by Olivier in Apoderus. Bedel proposed the name Cyphus their attelabid hosts and coined the term “thief weevils” for their for the species that were left in Attelabus by Olivier as well as many habit of taking over the niduses of their attelabid hosts. Al- other species placed there by subsequent authors. Bedel’s proprosal though not specifically stated, his comments on the biology of followed the International Code but subsequent workers, unaware these weevils were probably based mainly on observations of P. of the change or unwilling to change the names of numerous spe- ovatus in Maryland. According to Vogt, female pterocolines force cies, have not followed Bedel. Silfverberg (1977) successfully appealed their way into freshly made attelabid leaf rolls where they eat or to the International Commission on Zoological Nomenclature to destroy the host egg and oviposit their own. The pterocoline accept common usage of Attelabus in the interest of stability and to larva develops rapidly feeding on the decaying leaf tissue and confirm the designation by Schoenherr (1823) of Attelabus leaves the hollowed out leaf roll to pupate in the ground. curculionoides Linnaeus 1767 (= Curculio nitens Scopoli 1763) as type- Status of the classification. The subfamilies Rhynchitinae species. Two major groups of attelabids are now officially recognized and Pterocolinae, here considered in the family Attelabidae, have - Apoderus with A. coryli as its type and Attelabus with Attelabus nitens been placed in the family Rhynchitidae by other workers. The Scopoli as its type. Jekel (1860) divided the genus Attelabus into 16 family name Rhynchitidae is credited to Gistel 1856 (Alonso- subgenera and Voss (1925) elevated most of them to the generic Zarazaga and Lyal 1999). John L. LeConte (1876) was the first level. The genus Attelabus is applicable to only two species of attelabids American entomologist to use the family name Rhynchitidae for in America, north of Mexico - A. bipustulatus Fabricius and A. nigripes weevils in the subfamilies Rhynchitinae and Pterocolinae. Subse- LeConte. quent workers (Sharp 1889, Pierce1909, Blatchley and Leng 1916, Distribution. The family includes 1,914 world species in 97 Ting 1936, Anderson 1991) grouped these weevils in the family genera (Kuschel 1995). These weevils occur throughout the world Curculionidae. Pierce (1913) recognized the family Attelabidae but Kuschel points out that they do not occur in New Zealand, including the Rhynchitinae and the new tribes Rhynchitini and New Caledonia and the Pacific Islands. In the New World, 362 Auletini. Voss (1922-1969), in a worldwide taxonomic mono- species have been described including 180 species of leaf rolling graph series on the subfamilies Attelabinae, Rhynchitinae and weevils in four subfamilies (Attelabinae, Euscelinae, Hybolabinae Pterocolinae, consistently listed these groups in the family and Pilolabinae), 162 species of Rhynchitinae and 20 species of Curculionidae. Boving and Craighead (1931), in a synopsis of the Pterocolinae. In America north of Mexico, there are 51 total spe- larval forms of Coleoptera placed the Rhynchitinae and Attelabinae cies in 11 genera including 6 species of Attelabinae, 44 species of in the family Attelabidae. Crowson (1955) grouped these weevils Rhynchitinae and one species of Pterocolinae. 706 · Family 128. Attelabidae

KEY TO NEARCTIC SUBFAMILIES AND GENERA

1. Tarsal claws appendiculate (Figs. 7 and 8); man- dibles depressed, toothed on inner and outer margin; front legs subequal to middle and hind legs or middle and hind femora larger than front femora; tibial apices unarmed or with small spurs or mucros ...... 2 3 — Tarsal claws connate (Fig. 9); mandibles robust, not 2 toothed on outer margin; front legs enlarged, dis- tinctly larger than middle and hind legs; profemora distinctly swollen (Fig. 2); tibial apices uni-unci- nate (male) or bi-uncinate (female) (Figs. 5 and 6) (Attelabinae) ...... 10 6

2(1). Prothorax with distinct lateral carina (Fig. 3); propleura strongly excavated beneath carina; 5 body small, robust; metallic bluish-green to blu- ish-black (Pterocolinae) ...... Pterocolus — Prothorax not laterally carinate (Fig. 4); propleura not strongly excavated; body size, shape and 7 color variable (Rhynchitinae) ...... 3 4 9 3(2). Scutellary striole present ...... 4 — Scutellary striole absent ...... 6 8 FIGURES 2.128-9.128. 2. Himatolabus pubescens (Say), male, lateral 4(3). Pygidium completely or almost completely covered habitus; 3. Pterocolus ovatus (Fabricius), male, lateral habitus; 4. Rhynchites by elytra; elytra with some erect setae; males velatus LeConte, male, lateral habitus; 5. Himatolabus pubescens (Say), with one tooth on outer edge of mandibles and female, protibial apex; 6. Himatolabus pubescens (Say), male, protibial females with two ...... Eugnamptus — Pygidium mainly exposed, not completely or almost apex; 7. Pterocolus ovatus (Fabricius), tarsal claw; 8. Haplorhynchites completely covered by elytra; elytra without aeneus (Boheman), tarsal claw; 9. Himatolabus pubescens (Say), tarsal erect setae; mandibular teeth similar in both sexes claw...... 5 — Elytra not short, covering all abdominal terga and 5(4). Elytral striae distinct, more or less quadrate, moder- part of the pygidium; hind basitarsal segment ately to deeply impressed; intervals narrower shorter or subequal to hind tarsal segments 3 than width of striae, convex, smooth; interval and 4 combined ...... 8 punctures much smaller than striae; pubescence inconspicuous, more or less appressed; body 8(7). Elytral intervals wide, more or less flat, with numer- dark colored, in some feebly metallic bronze or ous punctures, punctures in some as large or blue, less than 4 mm in length ...... Temnocerus nearly as large as striae; males mostly more se- — Elytral striae more or less indistinct (more distinct tose along elytral suture at declivity; antennal in M. bicolor), weakly impressed, especially pos- club symmetrical in both sexes ...... teriorly; discernable intervals wider than width ...... Haplorhynchites of striae, more or less flat, in some minutely rug- — Elytral intervals narrow, more or less convex, mod- ose; interval punctures numerous, as large or erately punctured; punctures much smaller than nearly as large as striae; pubescence conspicu- striae; males not more setose along elytral suture ous, fine, semi-erect; body variable reddish-or- at declivity; males with asymmetrical antennal ange and black to brownish-black with faint blu- club ...... Involvulus ish metallic luster, greater than 4 mm in length ...... Merhynchites 9(6). Body with violaceous to greenish metallic luster; elytral pubescence of appressed whitish setae 6(3). Elytral striae distinctly rowed; intervals distinct, with and widely rowed, dark, erect setae; males with punctures more or less smaller than striae; punc- anterolateral spine on each side of pronotum(Fig. tures not masked by pubescence ...... 7 4); length mostly greater than 4 mm ...... — Elytral striae not distinctly rowed; intervals indis- ...... Rhynchites tinct, punctures as large or nearly as large as — Body without violaceous or greenish metallic lus- striae; punctures in some masked by pubescence ter; elytral pubescence not as above; males with- ...... 9 out anterolateral spines on each side of pronotum; length mostly less than 4 mm ...... Auletobius 7(6). Elytra short, exposing pygidium and two abdominal terga; hind basitarsal segment longer than com- 10(1). Upper surface with appressed pubescence ...... bined length of hind tarsal segments 3 and 4 ...... Himatolabus ...... Deporaus — Upper surface without appressed pubescence, gla- brous except for a few scattered fine erect setae ...... 11 Family 128. Attelabidae · 707

11(10). Profemora unarmed in both sexes; submentum in reddish humeral maculae; (key to New World species Hamilton males with pair of ventrally projecting acuminate 1992). spines; ventral rostral apex without median coni- cal prominence ...... 12 — Profemora in males armed with one or two blunt Xestolabus Jekel 1860, 1 sp., X. constrictipennis (Chittenden); fe- spine like projections; profemora in females un- males are leaf rollers; larvae develop in leaf rolls; occurs in the armed or armed with a single peg like projection; southwestern states of AZ and NM and south into Mexico; submentum in males without pair of ventral pro- jecting accuminate spines; ventral rostral apex associated with Rhus toxicodendron L. and Rhus spp. in the south- in lateral view with median conical prominence western U.S. (more pronounced in females) ...... Attelabus Rhynchitinae Gistel 1856 12(11). Body uniformly shiny black to brownish-black; head in both sexes subequal in size and shape; sur- face of abdominal sternites smooth, without tu- Rhynchitini Gistel 1856 bercles; pronotum with pair of basilateral pitlike depressions ...... Xestolabus Eugnamptus Schoenherr 1839, 9 spp.; known larvae are leaf min- — Body bicolored; pronotum, elytra and abdomen red to reddish-orange; legs, sterna and head black; ers; associated with species of Quercus, Sassafras, Juglans, Carya, head elongate in males; females with abdominal Cornus, Liquidambar, etc.; distributed in the eastern half of the sternites 1-3 with pair of acute tubercles; U.S. and southwestern states of AZ and NM; Eugnamptus pronotum without pair of pitlike pronotal depres- angustatus (Herbst) is the most abundant species in North America sions ...... Homoeolabus and occurs in the eastern half of the U. S. in 4 sympatric color forms (key to Nearctic species, Hamilton 1990). CLASSIFICATION OF NEARCTIC GENERA Haplorhynchites Voss 1938, 6 spp., larvae develop in flower heads Attelabidae Billberg 1820 cut by females; associated with various composite species of Helianthus, Silphium, Coreopsis, Viquiera, etc.; H. aeneus is the most Attelabinae Billberg 1820 common North American species and is widely distributed from south central Canada throughout the middle and eastern U.S.; Attelabus Linnaeus 1758, 2 spp., A. bipustulatus Fabricius and A. the other species occur in the southwestern states (key to Nearctic nigripes LeConte; females are leaf rollers; larvae develop in leaf species, Hamilton 1974). rolls; A. bipustulatus is associated with Quercus spp. and Carpinus sp. and ranges from SE Canada throughout the eastern U.S. and Involvulus Schrank 1798, 1 sp., I. hirtus (Fabricius); larvae develop southwest to OK and TX; A. nigripes is associated with Rhus in cut terminal shoots of Quercus spp.according to Vogt in copallina L. and other Rhus sp. and ranges throughout the eastern Kissinger (1964); ranges throughout northeastern U.S. and south- half of the U.S. and south into Mexico; the two species can be west to AZ. distinguished by color. A. bipustulatus is black with reddish hu- Euvolvulus Reitter 1916 meral maculae while A. nigripes is reddish throughout with vari- able darker areas (western form of A. nigripes is darker overall Merhynchites Sharp 1889, 4 spp.; larvae develop in buds or fruit with pale areas in basal half of elytra). (hips) of Rosa spp.; M. bicolor (Fabricius), the eastern rose curculio, develops in the hips and is distributed across the northern U.S. Homoeolabus Jekel 1860, 1 sp., H. analis (Illiger); females are leaf and southern Canada from coast to coast. The western rose curculio, rollers; larvae develop in leaf rolls; associated with many Quercus Merhynchites wickhami (Cockerell), develops in the buds and ranges sp.; ranges from SE Canada throughout the eastern half of the throughout the western half of the U.S. and southwestern U.S. (Volume 2, Color Fig. 28) Canada; the other 2 species occur locally in the southwestern U.S. (key to Nearctic species, Hamilton 1985). Himatolabus Jekel 1860, 2 spp., H. pubescens (Say), Fig. 2, and H. axillaris (Gyllenhal); females are leaf rollers; larvae develop in leaf Temnocerus Thunberg 1815, 14 spp.; known larvae develop in cut rolls; both species are associated with Quercus spp.; H. pubescens terminal shoots, buds and leaf primordia; associated with a wide also rolls the leaves of Alnus incana (L.) and Corylus americana variety of hosts including species of Quercus, Comptonia, Acacia, Walt. in northeastern North America; H. pubescens ranges from Manzanita, etc.; regionally distributed in U.S. (key to Nearctic spe- SE Canada and the northeastern U.S. southwest into TX, NM cies, Hamilton 1971). and AZ and south through Mexico; H. axillaris is recorded in the Pselaphorhynchites Schilsky 1903 southwestern states of AZ and UT and ranges south into Mexico; The two species can easily be separated by color. H. pubescens is Rhynchites Schneider 1791, 1 sp., R. velatus LeConte (Fig. 4); larvae reddish- brown throughout (with blackish-brown areas and darker develop in fruits of the desert plum, Prunus andersonii Gray; re- extremities in some) and H. axillaris is black to dilute black with corded only from CA and NV. 708 · Family 128. Attelabidae

Auletini Desbrochers 1908 CHITTENDEN, F. H. 1901. Some insects injurious to violet, rose and other ornamental plants. United States Department of Auletobius Desbrochers 1869, 9 spp.; known larvae develop in cut Agriculture Bulletin, 27:1- 114. terminal leaf primordia; associated with a wide variety of hosts COOLEY, R. A. 1903. Two insect pests: the rosebud curculio, including species of Quercus, Comptonia, Potentilla, Eriogonum, etc.; Rhynchites bicolor Fabr.- the poplar leaf-folding sawfly, Pontania regionally distributed in U.S. (key to Nearctic species, Pierce 1909; bozemani Cooley. Montana Experiment Station. Bulletin, 46: key to World species, Voss 1933-37). 107-118. CROWSON, R. A. 1955. The natural classification of the families Deporaini Voss 1929 of Coleoptera. E. W. Classey. Middlesex, England, 214 pp. DICKERSON, E. L. 1910. Notes on Rhynchites bicolor Fabr. Deporaus Samouelle 1819, 1 sp., D. glastinus LeConte; larvae are Journal of Economic Entomology, 3: 316-17. leaf miners; associated with Q. wislizenii in southern CA and EDWARDS, J. C. 1949. Coleoptera or beetles east of the Great other Quercus spp. throughout its range; ranges throughout west- Plains. J. W. Edwards. Ann Arbor, MI, 181 pp. ern U.S., east to CO and southeast to NM and west TX. ESSIG, E. O. 1958. Insects and mites of western North America. Platyrhynchus Thunberg (1815) MacMillan. New York, NY, ix-xiii + 1050 pp. EWING, H. E. 1915. A case of persistent melanism. Biological Pterocolinae Lacordaire 1866 Bulletin, 28: 224-228. FARRELL, B.D. 1998. “Inordinate fondness” explained: Why are Pterocolus Say 1831, 1 sp., P. ovatus (Fabricius) (Fig. 3); leaf roll there so many beetles? Science, 281: 555-559. thief, north of Mexico the larvae develop in leaf rolls prepared by FROST, C.A. 1908. Notes on Attelabus rhois and parasite. Psyche, three species of attelabine weevils (H. pubescens, A. bipustulatus 15: 26-32. and H. analis); distributed throughout eastern half of U.S., south- GATES, B. N. 1909. The rose curculio (Rhynchites bicolor Fabr.) in west to TX and AZ and south into Mexico (key to New World Massachusetts. Journal of Economic Entomology, 2: 465- species, Hamilton 1998). 66. GIRAULT, A. A. 1904. Attelabus bipustulatus Fabr., the theory of oviposition and construction of nidus; misc. notes. Ento- BIBLIOGRAPHY mological News, 15: 189-193. GISTEL, J. 1856. Die Mysterien der Europaischen Insectenwelt. ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL. 1999. A Dannheimer. Kempten, 12 + 532 pp. world catalogue of families and genera of Curculionoidea HAMILTON, R.W. 1969. Studies of the Rhynchophorous fami- (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae). lies Nemonychidae, Attelabidae and Rhynchitidae, with a Entomopraxis. Spain, 350 pp. revision of North American species of Attelabus Linnaeus, ANDERSON, D. M. 1991. Curculionidae (broad sense) (Cur- Rhynchites Schneider and Eugnamptus Schoenherr (Coleoptera: culionoidea). Pp. 594-612. In: F. W. Stehr, ed. Immature Curculionoidea). Unpublished Ph.D. dissertation, The Ohio Insects. volume 2. Kendall/Hunt. Dubuque, IA. State University, Columbus, Ohio, microfilm, 492 pp. BALDUF, W. V. 1959. Obligatory and facultative insects in rose HAMILTON, R.W. 1971. The genus Pselaphorhynchites in America, hips, their recognition and bionomics. University of Illinois north of Mexico (Coleoptera: Rhynchitidae). Annals of the Press. Urbana, IL., 194 pp. Entomological Society of America, 64: 982-996. BEDEL, L. 1882-1888. Faune des Coleopteres du Bassin de la HAMILTON, R.W. 1973. Observations on the biology of Seine. Rhynchophora. Annales del la Société Entomologique Haplorhynchites aeneus (Boheman) (Coleoptera: Rhynchitidae). de France, 6 (hors. Serie), 442 pp. Coleopterists Bulletin, 27: 83-86. BILLBERG, G. J. 1820. Enumeratio Insectorum in Museo G. J. HAMILTON, R.W. 1974. The genus Haplorhynchites in America Billberg. Gadel, Stockholm, 138 pp. north of Mexico (Coleoptera: Rhynchitidae). Annals of the BLATCHLEY, W. S. and C. W. LENG. 1916. Rhynchophora or Entomological Society of America, 67: 787-794. weevils of North Eastern America. The Nature Publishing HAMILTON, R.W. 1979. Taxonomic use of endophallic struc- Company. Indianapolis, IN, 682 pp. tures in some Attelabidae and Rhynchitidae of America north BØVING, A. G. and F. C. CRAIGHEAD. 1931. An illustrated of Mexico with notes on nomenclature. Annals of the Ento- synopsis of the principal larval forms of the order Coleoptera. mological Society of America, 72: 29-34. Entomologica Americana, 11: 1-351. HAMILTON, R.W. 1980. Notes on the biology of Eugnamptus BRIGHT, D. E. 1993. The Insects and Arachnids of Canada. Part collaris (Fabricius) (Coleoptera: Rhynchitidae) with descrip- 21. The weevils of Canada and Alaska: Volume 1. Coleoptera: tions of the larva and pupa. Coleopterists Bulletin, 34: 227- Curculionoidea, excluding Scolytidae and Curculionidae. 236. Publication 1882. Research Branch, Agriculture Canada. Ottawa, Canada. Family 128. Attelabidae · 709

HAMILTON, R. W. 1981. Description of the larva and pupa of LINNAEUS, C. V. 1758. Systema naturae per regna tria naturae Haplorhynchites aeneus (Boheman), (Coleoptera: Cur- secundum classes, ordines, genera, species, cum characteribus, culionoidea: Rhynchitidae). Journal of the Kansas Entomo- differentis, synonymis, locis, ed. 10. Salvius. Holmiae, vol. 1, logical Society, 54: 616-624. 823 pp. HAMILTON, R. W. 1983. Biological data on two North American LOVETT, A. L. 1915. The rose curculio, Rhynchites bicolor Fabr. rhynchitids (Coleoptera: Rhynchitidae) associated with sweet injures blackberry buds. Reports of the Department of En- fern, Comptonia peregrina (Linnaeus): with descriptions, illus- tomology, Oregon Agricultural Experiment Station Bulletin, trations and comparisons of their immature stages. Journal pp. 150-153. of the Kansas Entomological Society, 56: 511-522. LODING, H. P. 1945. Catalogue of the beetles of Alabama. HAMILTON, R. W. 1985. The genus Merhynchites Sharp in Wetumpka Printing. Wetumpka, AL, 172 pp. America north of Mexico. Southwestern Entomologist, 10: LUTZ, F. E. 1935. Field book of insects, 3rd ed. G. P. Putnam and 49-64. Sons. New York, NY, 510 pp. HAMILTON, R. W. 1990. A revision of the weevil genus MCBRIDE, D. K. and C. Y. OSETO. 1978. Sunflower insect Eugnamptus Schoenherr (Coleoptera: Rhynchitidae) in America pests. Cooperative Extension Service. North Dakota State north of Mexico. Transactions of the American Entomologi- University, Fargo, ND, Circular E-623, pp. 1-6. cal Society, 115: 475-502. MURTFELDT, M. E. 1872. Notes on Attelabus bipustulatus Fabr. HAMILTON, R. W. 1992. Revision of the New World Genus Canadian Entomologist, 4: 143-145. Himatolabus Jekel (Coleoptera: Attelabidae) in North America. O’BRIEN, C. W. and G. J. WIBMER. 1982. Annotated checklist Transactions of the American Entomological Society, 118:197- of the weevils (Curculionidae, sensu lato) of North America, 226. Central America and the West Indies (Coleoptera: HAMILTON, R. W. 1994. New life cycle data for two western Curculionoidea). Memoirs of the American Entomological North American weevils (Coleoptera: Rhynchitidae) with a Institute, 34: ii-ix, 1- 382. summary of North American rhynchitid biology. Coleopter- OLIVIER, A. G. 1807. Entomologie, ou histoire naturelle des ists Bulletin, 48: 331-343. insectes, avec leurs caractères génériques et spécifiques, leur HAMILTON, R. W. 1998. Taxonomic revision of the New World description, leur synonymie et leur figure enluminée. Desray. Pterocolinae (Coleoptera: Rhynchitidae. Transactions of the Paris, Coleopteres, vol. 5, 612 pp. American Entomological Society, 124: 203-269. PACKARD, A. S. 1890. Fifth report of the U.S. Entomological HAMILTON, R. W. and S. S. KURITSKY. 1981. Description of Commission on insects injurious to forest and shade trees. the larva and pupa of Merhynchites bicolor (Fabricius). Coleop- Government Printing Office. Washington, DC, 957 pp. terists Bulletin, 35: 189-195. PIERCE, W. D. 1909. Studies of North American weevils. Pro- HARRIS, T. W. 1862. A treatise on some of the insects injurious ceedings of the United States National Museum, 37(1708): to vegetation. Crosby and Nichols. Boston, MA, 640 pp. 325-364. HOERNER, J. L. 1936. Western Rose Curculio, Rhynchites bicolor PIERCE, W. D. 1913. Miscellaneous contributions to the knowl- wickhami Cockerell. Colorado Agricultural Experiment Sta- edge of the weevils of the families Attelabidae and tion Bulletin, 432: 1-19. Brachyrhinidae. Proceedings of the United States National HOPKINS, A. A. 1905. (Secretarial Notes). Proceedings of the Museum, 45 (1988): 365-426. Entomological Society of Washington, 7: 130-133. ROBERTSON, H. A.1923. The rose curculio in Manitoba with JEKEL, H. 1855-1860. Insecta saundersiana: or characters of notes on other insects affecting roses. Annual Reports of the undescribed insects in the collection of William Wilson Entomological Society of Ontario, 54: 12-16. Saunders, Esq. E. Newman. London, 242 pp. SCHOENHERR, C. J. 1823. Tabula synoptica familiae cur- KISSINGER, D. G. 1964. Curculionidae of America North of culionidum. Isis Oken, heft X, columns 1132-1146. Mexico. A key to the genera. Taxonomic Publications. South SCHULZ, T. J. and W. V. LIPP. 1969. The status of the sunflower Lancaster, MA, 143 pp. insect complex in the Red River Valley of North Dakota. KUSCHEL, G. 1995. A phylogenetic classification of Curculionoidea Proceedings of the North Central Branch, Entomological to families and subfamilies. Memoirs of the Entomological Society of America, 24: 99-100. Society of Washington, 14: 5-33. SHARP, D. E. 1889. Biologia Centrali Americana, Insecta, Co- LAWRENCE, J. L. 1982. Coleoptera. Pp. 482-553. In: S. P. Parker, leoptera, Curculionidae: Attelabinae. 4: 1-45. ed. Synopsis and classification of living organisms. McGraw/ SILFVERBERG, H. 1977. Attelabus Linnaeus, 1758 (Insecta, Hill. NY. Coleoptera): Request for confirmation of designation of type- LECONTE, J. L. 1876. In J. L. LeConte and G. H. Horn, The species. Z. N. (S.) 2209. Bulletin of Zoological Nomenclature, Rhynchophora of America, north of Mexico. Proceedings of 34: 189-90. the American Philosophical Society, 15: 1-455. THOMPSON, R. T. 1992. Observations on the morphology and classification of weevils (Coleoptera, Curculionoidea) with a key to major groups. Journal of Natural History, 26: 835-891. 710 · Family 128. Attelabidae

TING, P.C. 1936. The mouthparts of the coleopterous group Stettiner Entomologische Zeitung, 99: 59-117, 302-363 (1938); Rhynchophora. Microentomology, 1: 93-114. 103: 129-155 (1942); 104: 46-63 (1943). VAN EMDEN, F. 1938. On the taxonomy of Rhynchophora VOSS, E. 1938. Monographie der Rhynchitinen-Tribus Rhynchitini. larvae (Coleoptera). Transactions of the Royal Entomological 2. Gattungsgruppe: Rhynchitina. V. 2. Teil der Monographie Society of London, 87: 1-37. der Rhynchitinae-Pterocolinae. (45. Beitrag zur Kenntnis der VOGT, G. 1992. Leaf rolling weevils (Coleoptera: Attelabidae) Curculioniden), Koleopterologische Rundschau, 24: 129- their host plants, and associated rhynchitid weevils in North 171. America (Canada through the Republic of Panama): Sum- VOSS, E. 1941. Monographie der Rhynchitinen-Tribus Rhinocartini mary of a long term field study. Pp. 392-420. In: D. Quainter sowie der Gattungsgruppe Eugnamptina der Tribus and A. Aiello, eds. Insects of Panama and Mesoamerica: Rhynchitini. IV. Teil der Monographie der Rhynchitinae- Selected Studies. Oxford University Press, vxii + 692 pp. Pterocolinae. (32. Beitrag zur Kenntnis der Curculioniden). VOSS, E. 1922. Monographische Bearbeitung der Unterfamilie Deutche Entomologische Zeitschrift, 1941: 113-215. Rhynchitinae (Curc.) I, Teil, Nemonychini-Auletini. Archiv VOSS, E. 1941. Monographie der Rhynchitinen-Tribus Rhynchitini. für Natrurgeschichte, 88: 1-113. 2. Gattungsgruppe: Rhynchitina. V. 2. Teil der Monographie VOSS, E. 1925. Die Unterfamilien Attelabinae und Apoderinae der Rhynchitinae-Pterocolinae. (45. Beitrag zur Kenntnis der (Col. Curc.) (18. Beitrag zur Kenntnis der Curculioniden). Curculioniden). Mittheilungen Muenchener Entomologische Stettiner Entomologische Zeitung, 85: 1-78, 191-304. Gesellschaft, 31: 628-680. VOSS, E. 1932-33. Monographie der Rhynchitinen-Tribus VOSS, E. 1965. Die Unterfamilie Camarotinae, ihre Beziehungen Rhynchitini. 2. Gattungsgruppe: Rhynchitina. V. 1. Teil der zur Familie Attelabidae sowie ein Versuch, die Monographie der Rhynchitinae-Pterocolinae. (41. Beitrag zur phylogenetischen Zusammenhange innerhalb der letzteren Kenntnis der Curculioniden.) Koleopterologische zur Darstellung zu bringen (Coleoptera, Curculionidae) (191. Rundschau, 18: 153-189 (1932); 19: 25-56 (1933). Beitrag zur Kenntnis der Curculioniden). Entomologische VOSS, E. 1933-37. Monographie der Rhynchitinen-Tribus Auletini. Abhandlungen Staatliches Museum fur Tierkunde in Dresden, III. Teil der Monographie der Rhynchitinae-Pterocolinae. (37. 32 : 221-244. Beitrag zur Kenntnis der Curculioniden.) Stettiner VOSS, E. 1969. Monographie der Rhynchitinen-Tribus Rhynchitini. Entomologische Zeitung. 94: 108-136, 273-286 (1933); 95: 2. Gattungsgruppe: Rhynchitina. V. 2. Teil der Monographie 109-135, 330-344 (1934); 96: 91- 105, 229-241 (1935); 97: 279- der Rhynchitinae-Pterocolinae. Entomologische Arbeiten, 289 (1936); 98: 101-108 (1937). aus dem Museum G. Frey, 20: 117-375. VOSS, E. 1938-1943. Monographie der Rhynchitinen-Tribus WIBMER, G. J. and C. W. O’BRIEN. 1986. Annotated checklist Deporaini sowie der Unterfamilien Pterocolinae- Oxycoryninae of the weevils (Curculionidae, sensu lato ) of South America (Allocorynini). VII. Teil der Monographie der Rhynchitinae- (Coleoptera: Curculionoidea). Memoirs of the American Pterocolinae . (73. Beitrag zur Kenntnis der Curculioniden) Entomological Institute, 39: i-xvi, 1- 563. Family 129. Brentidae · 711

129. BRENTIDAE Billberg 1820

by Robert S. Anderson and David G. Kissinger

Family common name: The straight-snouted weevils; pear shaped weevils

Family synonyms: Apionidae Schoenherr 1823; Cycladidae Schoenherr 1823; Eurhynchidae Lacordaire 1863.

s used here, Brentidae is a composite of primitive weevil forms, not recognizable by any one or a few characters. Nearly all of them, with the exception of only the Nanophyinae, have straight, non-geniculate Aantennae. Apioninae and Nanophyinae are small ‘pear-shaped’ weevils immediately recognized by a long cylin- drical trochanter with the femur attached at its apex, a character state that is known in no other beetles. Brentinae, Cyphagoginae and Trachelizinae have elongated, parallel-sided bodies and a long, generally straight rostrum that often exhibits marked sexual dimorphism. Cyladinae are the sweet potato weevils, recognizable by their unique habitus.

Description (based on connate and much longer than following two; pygydium con- Lawrence 1982). Shape variable cealed by the elytra. Tegmen bilobed or notched apically and the elongate, narrow and parallel- median lobe with a “trough-like” ventral plate and a narrow dor- sided (Brentinae, Cyphag- sal plate. oginae, Trachelizinae), or Larvae (based on Lawrence 1982) when mature about 2.0- stouter and more robust with 18.0 mm in length, slender and subcylindrical (Brentinae) or broad expanded oval elytra and a dis- and strongly curved (Apioninae). Small legs rarely present on tinctive “pear-like” shape thorax (Brentinae), usually absent, but with well-developed pedal (Apioninae, Nanophyinae); flat lobes (Apioninae). Frontal sutures complete although rarely in- to convex; length 1.0-40.0 mm; distinct, reaching articulating membrane of mandible. Antenna color pale brown to black, very of a single membranous article. Hypopharyngeal bracon present. rarely red or bicolored or with Abdomen with first eight segments with two, three or four dor- contrasting colors or markings; sal folds and bearing annular or rarely biforous spiracles. vestiture lacking, or of fine Habits and habitats. As noted, Brentidae is a composite of short to moderately long ap- taxa of subfamily rank each of which has a rather different body pressed or suberect pubescence; form. Natural history aspects of Brentidae similarly are correlated rarely with scales. with, and best discussed, at the subfamily level (Thomas 1996). Rostrum moderately to Brentinae, Cyphagoginae and Trachelizinae are the traditional very long and usually narrow brentids. They are small to large, elongate weevils that are usually and cylindrical; sexual dimor- found under bark as both adults and larvae. Where known, fe- phism in rostrum form males lay their eggs in holes they have excavated with their ros- (Brentinae, Cyphagoginae, trum in living, dying or recently felled hardwood trees. Larvae 1.129. Arrenodes minutus (Drury) Trachelizinae) or length tunnel deep into the heartwood and appear to feed on wood and (from Bright 1993, reproduced (Apioninae) generally evident. fungus mycelia. Brentus anchorago Linnaeus is one of North with the permission of the Minister Antennae straight or rarely gen- America’s largest weevils, with males reaching a body length of of Public Works and Government iculate (Nano-phyinae), funicle up to 40 mm. In some species, adult males guard females during Services, 2001) very narrow and slender, rarely egg-laying. filiform or moniliform Cyladinae includes only the adventive Cylas formicarius (Fab- (Brentinae, Cyphagoginae, Trachelizinae), ending in a weak, loose ricius), the sweet potato weevil. Adults and larvae are associated club (Nanophyinae) or a compact club (Apioninae, Cyladinae) of with various Convolvulaceae (especially Ipomoea pescapreae L.) and three articles. Antennal insertions lateral at the middle of the are pests of sweet potatoes, Ipomoea batatas L. Larvae mine the rostrum or rarely dorsal near the base. Maxillary palps of two or stems. three articles. Labial palps small, of one or two articles, attached Little is known about Nearctic Nanophyinae. Most Palearctic dorsally near the apex of the prementum, rarely in cavities. Proven- Nanophyinae are associated with plants of the family Lythraceae, triculus lacks sclerotized plates. Front coxae vary from contiguous especially the genus Lythrum, but also Crassulaceae, Tamaricaceae to widely separated. Middle coxae and hind coxae narrowly to and Ericaceae. Larvae are associated with the fruits, leaves and widely separated. Front tibiae rarely equipped with an antenna- stems and many appear to cause galls. One species of Nanophyes cleaning organ. Hind wing usually with reduced anal venation. Schoenherr has been introduced into North America for biologi- Tarsal claws variable. First two visible sternites of the abdomen cal control of Lythrum salicaria L. (purple loosestrife; Lythraceae) 712 · Family 129. Brentidae

(Harris 2001) and a second is under consideration. Zeugonyx sabinae Notman has been reared from galls on Juniperus ashei Buchh. in Texas, but oth- erwise no information is avail- able on the native North American fauna. Apioninae are associated with a wide variety of plants, 34 mostly Fabaceae and Aster- aceae, but also Malvaceae, Polygonaceae, Caprifioliaceae, and Umbelliferae. Many of the FIGURES 3.129-4.129. 3. Front leg, Brentinae (schematic); 4. Front leg, Apioninae (schematic). newly elevated genera, for- merly subgenera of Apion sensu there are now 5 genera and 5 species recognized in North America. 2.129. Trichapion centrale (Fall) lato, are host specific on certain (from Bright 1993, reproduced This is based on phylogenetic evidence and follows the division families of plants (e.g., with the permission of the Minister of the once larger widespread genus Nanophyes Schoenherr by Alocentron on Malvaceae; of Public Works and Government Alonso-Zarazaga (1989) and followed by Alonso-Zarazaga and Perapion on Polygonaceae). Services, 2001) Lyal (1999). The situation with Apioninae is even more complex Larvae mine stems, feed on with many of the various subgenera (e.g., Kissinger 1968) now developing pericarp of fruit, or develop in the seeds. A few spe- elevated to generic rank, many generic concepts have been nar- cies are adventive. Rhopalapion longirostre (Olivier) is associated rowed with some North American species transferred into previ- with the ornamental hollyhock, Althea rosea Cav. (Tuttle 1954) ously Palearctic genera, and a number of new genera have been and Ischnopterapion virens (Herbst) with clover, Trifolium (Hoebeke proposed (Kissinger 1989, 1990, 1992, 1998, 1999; Alonso- et al. 2000). Two species of Exapion have been introduced into Zarazaga 1990). Alonso-Zarazaga (1990) treats the Palearctic clas- the western United States as biological control agents for gorse, sification of Apioninae in detail and is the basis for the develop- Ulex europaeus L. and scotch broom, Cytisus scoparius (L.) (both ment and understanding of the new system as applied in North Fabaceae) (O’Brien 1995) and Omphalapion hookerorum (Kirby) has America. At present in North America there are approximately been introduced for biological control of scentless chamomile, Matricaria perforata Mérat (Harris and McClay 2001). Podapion gallicola Riley 1883 has been associated with Pinus, larvae occur in galls on twigs (Bright 1993). Status of the classification. At present there is continuing controversy over the classification of the members of this family. Alonso-Zarazaga and Lyal (1999) recognize Brentidae (including Brentinae, Cyphagoginae, Trachelizinae and Cyladinae) as distinct from Apionidae and Nanophyidae and accord each family level status. This system is different from others such as Lawrence (1982) which recognizes Brentidae (including Brentinae, Cyphagoginae and Trachelizinae) and Apionidae (including Apioninae, Nanophyinae and Cyladinae). The family level classi- fication used here follows Lawrence and Newton (1995) and groups all of these taxa within Brentidae; however, at the subfamilial level, within Brentidae, the classification follows 6 Alonso-Zarazaga and Lyal (1999). O’Brien and Wibmer (1982) provide a checklist of the North American species of Nanophyinae and species of Apioninae (all as Apion sensu lato). Brentinae, Cyphagoginae, Trachelizinae and Cyladinae are well- known in North America and their classification remains un- changed (Warner 1960; Kissinger 1968). Brentinae have 3 species in 3 genera, and Cyphagoginae, Trachelizinae and Cyladinae each 5 have only 1 genus and 1 species represented. Apioninae and Nanophyinae on the other hand have undergone substantial changes at the generic level over the last few years. In Nanophyinae, FIGURES 5.129-6.129. 5. Dorsal habitus, Cylas formicarius (Fabricius 1798); 6. Lateral view, pronotum and base of elytra, Cylas formicarius. Family 129. Brentidae · 713

140 species placed in the 19 genera. A small number of North 7 910 American species have not yet been assigned to a genus in this new system. Identifying Apioninae is no easy task and the key to the genera of Apioninae formerly placed in Apion sensu lato (start- ing at couplet 15) is based extensively on male characters only. Also, many of the characters used in establishing the new generic classification are based on male genitalia and require dissections and detailed study. Distribution. Collectively this family is widely distributed in North America, but some of the constituent subfamilies have very restricted distributions. Apioninae are widespread in North America and found in most habitat types from sea level to the alpine zone. Nanophyinae are also widespread but do not extend as far north as do Apioninae, nor do they include the diversity of habitats of the latter group, but then there are also many fewer species. Excluding Cyladinae, which has only the one adventive species, Cylas formicarius Fabricius, Brentinae, Cyphagoginae and Trachelizinae are generally tropical in their distributions and 4 of the 5 species that occur in North America only do so in extreme southern Florida (Brentus anchorago (Linnaeus), Stereodermus exilis Suffrian, and Paratrachelizus uncimanus (Boheman)) or southern Texas (Heterobrenthus texanus Schaeffer). The remaining species, Arrenodes minutus (Drury), is widespread in eastern North America.

KEY TO THE NEARCTIC SUBFAMILIES AND GENERA 11 1. Trochanter short and triangular, femur attached to the side of trochanter (Fig. 3); body more elon- 8 gate and narrow, length greater than 3.0 mm, most greater than 10.0 mm (Figs. 1, 5, 14) ...... 2 — Trochanter long and somewhat cylindrical, femur attached to the apex of trochanter (Fig. 4); body “pear-shaped”, length less than 5.0 mm, most less than 3.0 mm (Figs. 2, 16, 31) ...... 7

2(1). Pronotum with a broad constricted collar (that is 13 almost as long as the rest of the pronotum) be- fore base (Fig. 5); mesothorax with a deep verti- cal sulcus near the posterior margin of the pronotum (Fig. 6); tarsal claws connate at base; elytra elongate-oval in dorsal view, lacking im- pressed striae (Fig. 5) (Cyladinae) ...... Cylas — Pronotum with at most a very short constriction be- fore base (Figs. 1, 7, 9, 10, 14); mesothorax lack- ing a deep lateral pit near the posterior margin of the pronotum; tarsal claws free; elytra elongate and parallel-sided in dorsal view, with distinctly 12 or indistinctly impressed striae ...... 3

3(2). All femora lacking distinct tooth (although slight an- gulation may be present on front femur); rostrum with ventral surface punctate and with dense microsetae or sparse elongate recumbent setae ...... 4 FIGURES 7.129-13.129. 7. Dorsal habitus, Stereodermus exilis Suffrian; — At least front femur with obvious tooth; rostrum with 8. Front leg, Stereodermus exilis; 9. Dorsal habitus, Paratrachelizus ventral surface smooth or coarsely punctate, not uncimanus (Boheman); 10. Dorsal habitus, Brentus anchorago (Linnaeus), setose (Brentinae) ...... 5 male; 11. Dorsal view, head and pronotum, Brentus anchorago, female; 12. Dorsal view, head, Arrenodes minutus (Drury), male; 13. Front leg, 4(3). Elytra with stria 3 bent inwards at middle towards Arrenodes minutus. elytral suture and coalescent with stria 2 (Fig. 7); front tibia with inner edge incised in apical one- 714 · Family 129. Brentidae

and female (long and cylindrical anterior to point of antennal insertion) ...... Heterobrenthus

7(1). Antenna geniculate, scape distinctly longer than combined length of first 3 articles of funicle; fu- nicle apparently with only 5 articles, articles 6 and 7 as wide as apical antennal article and form- ing a loose club (Nanophyinae) ...... 8 — Antenna straight, not geniculate, scape shorter than combined length of first 3 articles of funicle; fu- nicle with 7 articles, articles 6 and 7 narrow, not as wide as apical antennal article and distinct from compact apical club (Apioninae) ...... 12

8(7). Tarsal claws equal in length and form ...... 9 — Inner tarsal claw shorter and finer than outer claw ...... 11

9(8). Elytra with interstria 8 with a row of small, dark tu- bercles (Fig. 15); elytra largely pale, with dark, oblique fascia extended from basal one-third at middle to humeri; vestiture consisting of sparse, erect setae and long, decumbent, moderately dense scales intermixed; length 2.0-2.3 mm ..... FIGURE 14.129. Dorsal habitus, Heterobrenthus texanus Schaeffer, ...... Pseudotychius — Elytra with interstria 8 smooth in basal one-quarter; male. elytral vestiture various; length 1.2-1.9 mm .. 10

half, incision surrounded by very dense setae 10(9). Elytra lacking fascia, setae appressed and uniformly (Fig. 8); rostrum with ventral surface with sparse arranged, each seta short, not or hardly exceed- elongate recumbent setae (Cyphagoginae) ...... ing the base of the following one (Fig. 16); ...... Stereodermus pronotum and tibia lacking specialized erect se- — Elytra with stria 3 evident only as a row of shallow tae, only one subhumeral seta on elytral interstria punctures, straight throughout length, not coa- 9 (Fig. 16) ...... Microon lescent with stria 2 (Fig. 9); front tibia with inner — Elytra with fascia, setae arcuate and disordered, edge straight, not incised; rostrum with ventral each seta long, usually exceeding the base of surface with dense microsetae in basal portion the following one (Fig. 17); pronotum, tibia and (Trachelizinae) ...... Paratrachelizus odd elytral interstriae with specialized erect se- tae (Fig. 17) ...... Nanophyes 5(4). Head longer than broad, not distinctly constricted behind eyes, lateral margins continuous, pro- 11(8). Antenna with basal 5 articles of funicle slender; longed and convergent behind eyes (Figs. 10, elytra largely pale, with basal dark area; vestiture 11); pronotum medially deeply sulcate in basal fine and short, consisting of whitish and dark one-half (Figs. 10, 11); male with pronotum ex- scales ...... Nanodactylus tremely elongated, lateral margins constricted — Antenna with basal 4 articles of funicle slender; near middle (Fig. 10) ...... Brentus elytra uniformly testaceous; vestiture of long — Head short, width more or less equal to length, dis- whitish setae ...... Zeugonyx tinctly constricted behind eyes, lateral margins at most slightly prolonged but interrupted by con- 12(7). Middle coxae contiguous, not separated by junc- striction, not convergent behind eyes (Figs. 1, tion between mesosternal process and intercoxal 12, 14); pronotum uniformly smooth dorsally, not process of metasternum ...... 13 sulcate (Figs. 1, 14); male with pronotum not elon- — Middle coxae separated by junction between gated, and lateral margins not constricted near mesosternal process and intercoxal process of middle...... 6 metasternum...... 15

6(5). Antenna with articles 2 to 11 more or less equal in 13(12). Tarsal claws with acute basal tooth .... Chrysapion length and width towards apex (Fig. 1); head con- — Tarsal claws simple, at most swollen as base, not stricted behind eyes, hind angles not projecting toothed...... 14 (Figs. 1, 12); front tibia with at most a swelling on inner margin, no distinct tooth evident (Fig. 13); 14(13). Rostrum straight, in side view not forming angle at rostrum very dissimilar in male (short and broad junction with frons, apical half of rostrum usually with large mandibles, Fig. 12) and female (narrow tapered to apex (Fig. 18); eyes not or scarcely and cylindrical, Fig. 1) ...... Arrenodes prominent; antennae inserted at basal 0.29-0.47 — Antenna with articles 2 to 11 increasing in length of rostrum; scape at most as long as width of ros- and width towards apex (Fig. 14); head strongly trum at midlength; prescutellar fovea conspicu- constricted behind eyes, hind angles projecting; ous, lanceolate to sulciform; tarsi normal, front front tibia with distinct tooth on inner margin (Fig. tarsus with first article 1.25-1.60 X as long as wide; 14); rostrum dissimilar in male (dilated at apex) internal sac of aedeagus without baculi, other Family 129. Brentidae · 715

15 16 17

18 19

20

21 22 23 24

25 26 27 28 29

35 32 33 34 30 31

FIGURES 15.129-35.129. 15. Lateral habitus, Pseudotychius watsoni Blatchley; 16. Dorsal habitus, Microon canadense (Brown); 17. Lateral habitus, Nanophyes marmoratus (Goeze); 18. Lateral view, head, Perapion; 19. Lateral view, head, Podapion; 20.Front leg, Fallapion, male; 21. Dorsal view, pygydium, Aspidapiini; 22. Lateral view, pygydium, Aspidapiini; 23. Dorsal view, pygydium, Ixapiini; 24. Lateral view, pygydium, Ixapiini; 25. Dorsal view, pronotum, Alocentron (schematic); 26. Dorsal view, pronotum, Rhopalapion (schematic); 27. Dorsal view, pygydium, Oxystomatini; 28. Lateral view, pygydium, Oxystomatini; 29. Dorsal view, tegmen, Betulapion simile (Kirby), male; 30. Dorsal view, tegmen, Mesotrichapion, male; 31. Habitus, Apion, undetermined species, Alberta, Canada; 32. Elytral apex, Apion; 33. Elytral apex, Ischnopterapion; 34. Dorsal view, head, Sayapion; 35. Dorsal view, head, Mesotrichapion, male (Figs. 21-30 redrawn after Alonso-Zarazaga 1990).

structures may be present; associated with — Front femur of male without polished area on ven- Polygonaceae ...... Perapion tral surface ...... 16 — Rostrum strongly curved, in side view forming a conspicuous angle at junction with frons, apical 16(15). Pygydium of male lacking distinct, deep, transverse half of rostrum parallel-sided to slightly widened preapical sulcus, profile of pygydium not inter- to apex (Fig. 19); eyes prominent; antennae in- rupted (Figs. 21, 22); abdominal ventrite 5 of male serted at basal 0.10-0.19 of rostrum; scape longer rounded apically; elytra with specialized seta on than width of rostrum at midlength; prescutellar interstria 7 lacking ...... 17 fovea inconspicuous, similar to pronotal punc- — Pygydium of male with distinct, deep, transverse tures; tarsi robust, front tarsus with first article preapical sulcus, profile of pygydium distinctly 0.85-1.00 X as long as wide; internal sac of interrupted by sulcus (Figs. 23, 24, 27, 28); ab- aedeagus with two baculi and denticles; associ- dominal ventrite 5 of male truncate apically; elytra ated with Pinus (Pinaceae) ...... Podapion with specialized seta on interstria 7 at middle or near apical 1/3 present or lacking ...... 19 15(12).* Front femur of male with polished (sometimes stri- ate) area on ventral surface, the area generally * From this point on the key is complex and relies extensively on limited posteriorly by a prominent longitudinal characters of males. Males can be recognized by the shorter, more carina (Fig. 20); metasternum generally with a pair coarsely sculptured rostrum and (sometimes) by an upward deflection of spicules near the middle of the posterior mar- of the last abdominal ventrite. These are the taxa traditionally treated gin ...... Fallapion as Apion sensu lato and the reader may wish to consult Bright (1993) and Kissinger (1968) for species level identifications. 716 · Family 129. Brentidae

17(16). Antennal club with sutures obsolete or absent; tar- gins of pronotum various; male genitalia with sal claws simple, not toothed; on Matricaria prostegium with at most moderately high basal perforata (Asteraceae) ...... Omphalapion median carina not projected basally; elytral — Antennal club with sutures distinct; tarsal claws interstria 2 only slightly prolonged outward at toothed at base; on Malvaceae ...... 18 apex (Fig. 33); on Fabaceae ...... 24

18(17). Pronotum with vestiture directed away from the mid- 24(23). Elytra blue-green; elytral interstria 7 with one spe- line at the base, more or less parallel to the mid- cialized seta in apical one-third; legs dark; lateral line along the lateral margins, and perpendicular margins of pronotum parallel in basal one-half; to the apical margin along the apical margin (Fig. rostrum not prominently expanded at point of in- 25); pronotum with basal flange well-developed sertion of antennae; eyes prominent ...... (Fig. 25); on Malvaceae ...... Alocentron ...... Ischnopterapion — Pronotum with vestiture directed towards the mid- — Elytra black; elytral interstriae 7 and 9 with one spe- line, in some cases, totally transverse at the basal cialized seta near apex; legs various, may be light or apical margins (Fig. 26); pronotum with basal in color; lateral margins of pronotum various in flange obsolete (Fig. 26); on Althea rosea basal one-half; rostrum variously expanded at (Malvaceae) ...... Rhopalapion point of insertion of antennae; eyes prominent or not ...... 25 19(16). Elytra with transverse pattern of contrasting light and dark scales; pygydium of male with sulcus 25(24). Pronotum without basal flange, with lateral margins incomplete laterally, not reaching side margins nearly evenly convergent from base to apex, of pygydium (Fig. 24); front tarsus with article 2 subconical in dorsal form; rostrum prominently stout, about as wide as long; femora and tibiae expanded at point of insertion of antennae; legs robust; middle coxae widely separated (0.25-0.30 and article 1 of antenna dark ...... Apionion x diameter of middle coxa); aedeagus with inter- — Pronotum with or without basal flange, with lateral nal sac with large apically bifurcate structure; on margins various; rostrum smooth at point of inser- Rutaceae, Caprifoliaceae ...... Neapion tion of antennae, not expanded; legs and article — Elytra with scales more uniformly colored, not con- 1 of antenna dark or light ...... 26 trasting light and dark; pygydium of male with sulcus complete laterally, reaching side margins 26(25). Pronotum with basal flange indistinct, subconical of pygydium (Figs. 27, 28); front tarsus with ar- in dorsal form (Fig. 34); legs and article 1 of an- ticle 2 usually longer than wide; femora and tibiae tenna light; aedeagus with median lobe and teg- slender; middle coxae moderately widely sepa- men narrow, elongate, subcylindrical; internal sac rated (<0.25 x diameter of middle coxa); aegeagus with large teeth (>0.030 mm long) ...... Sayapion with internal sac without large structure; on — Pronotum without basal flange, lateral margins Fabaceae and other plants...... 20 subparallel in basal one-half; legs and article 1 of antenna light or dark; aedeagus with median lobe 20(19). Genitalia of male with tegmen with prostegium ar- slightly depressed, broader at base; internal sac ticulated with free ring (Fig. 29) ...... 21 with largest teeth (<0.020 mm long) .... Kissingeria — Genitalia of male with tegmen with prostegium fused with free ring (Fig. 30) ...... 28 27(21). Head with ventral surface with subcephalic ridges lacking or low, not extended basally on head to 21(20). Tibiae of male simple, not mucronate ...... 22 point equivalent to middle of eye; vestiture of — Middle tibia (at least) of male mucronate ...... 27 interstriae 2-5 sparse, generally inconspicuous, consisting of one row of scales; one specialized 22(21). Scrobe with dorsal margin produced into a long, seta on interstria 9; aedeagus with parameroid slender acute process projected ventrally; elytra lobe with one or more macrochaetae; on Betula with interstria 7 with specialized seta inserted papyrifera (Betulaceae) ...... Betulapion near midlength of stria; metasternum very con- — Head with ventral surface with subcephalic ridges vex in lateral view; meso- and metasternal pro- moderately to well developed, extended to or cesses directed inwardly where meeting between beyond the middle of the eye; vestiture of hind coxae; on Fabaceae ...... Exapion interstriae 2-5 relatively dense, consisting of two — Scrobe with dorsal margin simple, not produced into or more rows of scales; one specialized seta on a long, slender acute process; elytra with interstria each of interstriae 7 and 9; aedeagus with 7 or 9, or 7 and 9, with or without specialized parameroid lobe generally lacking macrochaetae; seta inserted at or behind apical 1/3 of elytron; associated mainly with Fabaceae and Asteraceae metasternum slightly convex in lateral view; meso- ...... Trichapion and metasternal processes either flat or with metasternal process more prominent where meet- 28(20). Elytra blue in color, legs dark; tarsus with claw ing between hind coxae; on various plants .. 23 simple; rostrum of male markedly expanded later- ally at point of antennal insertion (Fig. 35); frons 23(22). Body color red throughout; lateral margins of flat, striate ...... Mesotrichapion pronotum parallel in basal one-half (Fig. 31); male — Elytra not blue in color; rostrum of male smooth, not genitalia with prostegium with high median crista expanded laterally at point of antennal insertion projected basally; elytral interstria 2 prolonged ...... 29 outward at apex (Fig. 32); on Polygonaceae ...... Apion 29(28). Male with metasternum tuberculate on median pos- — Body color black or with metallic sheen, occasion- terior margin and apical area of parameroid lobe ally with light colored appendages; lateral mar- Family 129. Brentidae · 717

bearing four or more macrochaetae more than Apioninae Schoenherr 1823 0.055mm long ...... Eutrichapion — Male metasternum not tuberculate or parameroid lobe lacking macrochaetae more than 0.010mm Apionini Schoenherr 1823 long ...... Coelocephalapion Apion Herbst 1797, 1 sp., identity uncertain, Alberta. Adults have been collected around edges of high elevation snowfields. Asso- CLASSIFICATION OF THE NEARCTIC GENERA ciated with Polygonaceae in Palearctic Region. Apiolum Kirby 1808 Brentidae Billberg 1820 Apionus Rafinesque 1814 Apius Billberg 1820, not Panzer 1806 Brentinae Billberg 1820 Apium Agassiz 1846 Oxystomum Gistel 1856 Brentini Billberg 1820 Oxeostomum Gistel 1856 Erythrapion Schilsky 1906 Brentus Fabricius 1787, 1 sp., B. anchorago (Linnaeus 1758), Florida. Adults are often found under bark of Bursera simaruba (L.) Sarg. Aplemonini Kissinger 1968 Brenthus Illiger 1801 Perapion Wagner 1907, 4 spp., generally distributed. Associated Arrhenodini Lacordaire 1866 with Polygonaceae. Eroosapion Ehret 1994 (valid subgenus) Arrenodes Schoenherr 1823, 1 sp., A. minutus (Drury 1770), gener- Hemiperapion Wagner 1930 (valid subgenus) ally distributed in eastern United States, into extreme southern Rhaphidoplectron Alonso-Zarazaga 1990 (valid subgenus) Canada. Adults are found under bark especially of oaks; larvae bore into the wood (Buchanan 1960). Males are territorial and Podapion Riley 1883, 1 sp., P. gallicola Riley 1883, generally distrib- guard females during egg-laying (Sanborne 1983). uted in eastern United States into extreme southern Canada; also Brentus Panzer 1788, not Fabricius 1787 in California and Oregon. Associated with Pinus spp. Arrhenodes Schoenherr 1826 Eupsalis Lacordaire 1866 Aspidapiini Alonso-Zarazaga 1990 Platysystrophus Kleine 1917 Alocentron Schilsky 1901, 6 spp., generally distributed. Associated Heterobrenthus Sharp 1895, 1 sp., H. texanus Schaeffer 1915, south- with Malvaceae. ern Texas. Bulborhinapion Schatzmayr 1926 (valid subgenus) Nearctalox Alonso-Zarazaga 1990 (valid subgenus) Cyphagoginae Kolbe 1892 Ceratapiini Alonso-Zarazaga 1990 Stereodermini Sharp 1895 Omphalapion Schilsky 1901, 1 sp., O. hookerorum (Kirby 1808), Brit- Stereodermus Lacordaire 1866, 1 sp., S. exilis Suffrian 1870, south- ish Columbia, Alberta, Saskatchewan, Manitoba and Nova Scotia; ern Florida. adventive. This species was first discovered in Nova Scotia (Peschken et al. 1993) but has recently been introduced into west- ern Canada for biological control of Matricaria perforata Mérat Trachelizinae Lacordaire 1866 (scentless chamomile; Asteraceae) (Harris and McClay 2001).

Trachelizini Lacordaire 1866 Exapiini Alonso-Zarazaga 1990

Paratrachelizus Kleine 1921, 1 sp., P. uncimanus (Boheman 1839), Exapion Bedel 1997, 2 spp., California, Oregon and Washington. southern Florida. Adventive; introduced for biological control of gorse, Ulex europaeus L. and scotch broom, Cytisus scoparius (L.) (Fabaceae) Cyladinae Schoenherr 1823 (O’Brien 1995). Ulapion Ehret 1997 (valid subgenus) Cylas Latreille 1802, 1 sp., C. formicarius (Fabricius 1798), generally distributed in southern United States. Adventive on Ixapiini Alonso-Zarazaga 1990 Convolvulaceae including Ipomoea batata (L.) Lam. (sweet potato). Cylanus Rafinesque 1815 Neapion Alonso-Zarazaga 1990, 6 spp., generally distributed in Protocylas Pierce 1941 eastern North America west to southern Texas, north into ex- 718 · Family 129. Brentidae treme southern Canada. Subgenus Neotropion, 1 sp., N. xanthoxyli Incertae sedis (Fall), associated with Rutaceae (genus Zanthoxylum); subgenus Neapion, 5 spp., associated with Caprifoliaceae (genus Viburnum). Apionion Kissinger 1998, 2 spp., A. crassum (Fall 1898), eastern Xixias Kissinger 1990 United States; A. dilatatum (Smith 1884), Arizona. Associated Neotropion Alonso-Zarazaga 1990 (valid subgenus) with Fabaceae.

Malvapiini Alonso-Zarazaga 1990 Coelocephalapion Wagner 1914, 22 spp., generally distributed. As- sociated mostly with Fabaceae and Asteraceae. Rhopalapion Schilsky 1906, 1 sp., R. longirostre (Olivier 1807), gen- erally distributed. Adventive on Althea rosea Cav. (hollyhock; Sayapion Kissinger 1999, 5 spp., eastern United States, Texas and Malvaceae). Arizona. One species has been associated with Fabaceae.

Oxystomatini Alonso-Zarazaga 1990 Nanophyinae Gistel 1856

Oxystomatina Alonso-Zarazaga 1990 Nanophyini Gistel 1856

Eutrichapion Reitter 1916, 3 spp., generally distributed. Associ- Microon Alonso-Zarazaga 1989, 1 sp., M. canadense (Brown 1944), ated with Fabaceae. western United States south to Arizona and western Canada east Cnemapion Bokor to Manitoba. At least one Palearctic species is associated with the Leconteapion Alonso-Zarazaga 1990 (valid subgenus) genus Lythrum (Lythraceae). Phalcrolobus Alonso-Zarazaga 1990 (valid subgenus) Psilocalymma Alonso-Zarazaga 1990 (valid subgenus) Nanodactylus Blatchely 1922, 1 sp., N. obesulus Blatchley 1922, Illi- nois, Indiana, Texas. Mesotrichapion Györffy 1956, 1 sp., M. cyanitinctum (Fall 1927), Alaska to northern Quebec, south to southern Manitoba. Asso- Nanophyes Schoenherr 1838, 1 sp., N. marmoratus (Goeze 1777), ciation with Fabaceae (genus Astragalus). Manitoba, New York; introduced into Canada in 1997 for the biological control of Lythrum salicaria L. (Lythraceae) (Harris 2001). Trichapiina Alonso-Zarazaga 1990 This weevil species is also approved for introduction into the United States and is established in the vicinity of Ithaca, New Betulapion Ehret 1994, 1 sp., B. simile (Kirby 1811), generally dis- York. A second species, N. brevis Boheman 1845, also is under tributed. Adventive on Betula papyrifera Marsh. (paper birch; consideration for introduction. Salicaceae); larvae develop in flowers. Pseudotychius Blatchley 1922, 1 sp., P. watsoni Blatchley 1922, east- Kissingeria Alonso-Zarazaga 1990, 6 spp., generally distributed. ern United States, north into Ontario. Associated with Fabaceae. Zeugonyx Notman 1922, 1 sp., Z. sabinae Notman 1922, Texas. Trichapion Wagner 1912, 45 spp., generally distributed. Associ- This species was reared from galls on twigs of Juniperus ashei ated mostly with Fabaceae and Asteraceae. Buchh. (Cupressaceae).

Synapiina Alonso-Zarazaga 1990 BIBLIOGRAPHY

Ischnopterapion Bokor 1923, 1 sp., I. virens (Herbst 1797), New ALONSO-ZARAZAGA, M. A. 1989. Revision of the supraspecific York, Pennsylvania, Maryland, New Jersey, Connecticut, Dela- taxa in the Palaearctic Apionidae Schoenherr, 1823. 1. Intro- ware and Virginia. Adventive on various species of clover, Trifo- duction and subfamily Nanophyinae Seidlitz, 1891 (Co- lium (Fabaceae) (Hoebeke et al. 2000). leoptera, Curculionoidea). Fragmenta Entomologica, 21: 205- 262. Piezotrachelini Voss 1959 ALONSO-ZARAZAGA, M. A. 1990. Revision of the supraspecific taxa in the Palaearctic Apionidae Schoenherr, 1823 (Co- Chrysapion Kissinger 1968, 2 spp., Arizona, California and Texas. leoptera, Curculionoidea). 2. Subfamily Apioninae Schoenherr, 1823: Introduction, keys and descriptions. Fallapion Kissinger 1968, 30 spp., generally distributed. Associ- Graellsia, 46: 19-156. ated mostly with Asteraceae and Umbelliferae. ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL. 1999. A world catalogue of families and genera of Curculionoidea (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae). Entomopraxis. Barcelona, Spain. Family 129. Brentidae · 719

BRIGHT, D. E. 1993. The Insects and Arachnids of Canada. Part KISSINGER, D. G. 1998. Apionidae from North and Central 21. The weevils of Canada and Alaska: Volume 1. Coleoptera: America. Part 5. Description of the genus Apionion and 4 new Curculionoidea, excluding Scolytidae and Curculionidae. species (Coleoptera). Insecta Mundi, 12: 93-102. Publication 1882. Research Branch, Agriculture Canada. Ot- KISSINGER, D. G. 1999. Description of a new genus, Sayapion, tawa, Canada. from North and Central America (Coleoptera: Apionidae). BUCHANAN, W. D. 1960. Biology of the oak timberworm, Insecta Mundi, 13: 72. Arrhenodes minutus. Journal of Economic Entomology, 53: LAWRENCE, J.L. 1982. Coleoptera, Pp. 482-553. In: S. P. Parker, 510-513. ed. Synopsis and Classification of Living Organisms. Volume HARRIS, P. 2001. Nanophyes marmoratus (Goeze). Flower-feeding 2. McGraw Hill. New York. weevil. http://res2.agr.ca/lethbridge/weedbio/agents/ LAWRENCE. J. F. and A. F. NEWTON, Jr. 1995. Families and ananmar.htm subfamilies of Coleoptera (with selected genera, notes, refer- HARRIS, P. and A. MCCLAY. 2001. Omphalapion hookeri Kirby. ences and data on family-group names). Pp 779-1006. In: J. Seed-head weevil. http://res2.agr.ca/lethbridge/weedbio/ Pakaluk and S. A. Slipinski, eds., Biology, phylogeny and agents/aomphook.htm classification of Coleoptera: Papers celebrating the 80th birth- HOEBEKE, E. R., R. A. BYERS, M. A. ALONSO-ZARAZAGA, day of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, and J. F. STIMMEL. 2000. Ischnopterapion (Chlorapion) virens Warsaw. (Herbst) (Coleoptera: Curculionoidea: Brentidae: Apioninae), O’BRIEN, C. W. 1995. Curculionidae, premiere biological control a Palearctic clover pest new to North America: recognition agents. Memoirs of the Entomological Society of Washing- features, distribution, and bionomics. Proceedings of the ton, 14: 129-136. Entomological Society of Washington, 102: 151-161. O’BRIEN, C. W. and G. J. WIBMER. 1982. Annotated checklist KISSINGER, D. G. 1968. Curculionidae subfamily Apioninae of of the weevils (Curculionidae sensu lato) of North America, North and Central America with reviews of the world genera Central America, and the West Indies (Coleoptera: of Apioninae and world subgenera of Apion Herbst (Co- Curculionoidea). Memoirs of the American Entomological leoptera). Taxonomic Publications. South Lancaster, Massa- Institute, 34: i-ix, 1-382. chusetts. PESCHKEN, D. P., K. C. SAWCHYN and D. E. BRIGHT. 1993. KISSINGER, D. G. 1989. Apionidae from North and Central First record of Apion hookeri Kirby (Coleoptera: Curculionidae) America. Part 1. Notes on the classification of the Apion in North America. Canadian Entomologist, 125: 629-631. subgenus Trichapion Wagner with description of new species SANBORNE, M. 1983. Some observations on the behaviour of from the United States of America (Coleoptera). Insecta Arrhenodes minutus (Drury) (Coleoptera: Brentidae). Coleop- Mundi, 3: 271-227. terists Bulletin, 37: 106-113. KISSINGER, D. G. 1990. Apionidae from North and Central THOMAS, M. C. 1996. The primitive weevils of Florida (Co- America. Part 2. Description of a new subgenus and two new leoptera: Brentidae: Brentinae). Florida Department of Agri- species of Apion from Mexico. (Coleoptera). Insecta Mundi, culture. Division of Plant Industry. Entomology Circular No. 4: 33-40. 375. 3 pp. KISSINGER, D. G. 1992. Apionidae from North and Central TUTTLE, D. M. 1954. Notes on the bionomics of six species of America. Part 4. Generic classification and introduction to the Apion (Curculionidae, Coleoptera). Annals of the Entomo- genus Coelocephalapion Wagner, with new species from Mexico logical Society of America, 47: 301-307. and Venezuela (Coleoptera). Insecta Mundi, 6: 65-77. WARNER, R. E. 1960. The genus Stereodermus new to America north of Mexico with a revised key to the genera of Brentidae. Coleopterists Bulletin, 14: 29. 720 · Family 130. Ithyceridae

130. ITHYCERIDAE Schönherr 1823

by Robert S. Anderson

Family common name: The New York weevil

mong the primitive weevils with straight, non-geniculate antennae, this enigmatic family contains only one species, whose adults can be recognized by their large size (12-18 mm long), stout form, and distinct pubes- Acence. Description (based on palp of two articles; palps widely separated. Premental sclerite Lawrence 1982): Body large, subtriangular. Thorax with pronotal sclerite moderately pig- stout. Pubescence of scale-like mented. Legs very small, widely separated, subconical, two or bristles, distinct. three jointed. Abdomen with first four segments with three dor- Head with single gular su- sal folds, segments 5-8 with two dorsal folds. Spiracles annular. ture; pregular sutures absent. Pupa (based on Sanborne 1981) with distinct labrum present; Rostrum broad, stout, not mandibles with one pair of short setae; setosity extensive. sexually dimorphic; antennal Habits and habitats. Adult I. noveboracensis are associated insertions lateral. Maxilla with- with various species of Fagaceae, Betulacae and Juglandaceae. out lacinia; maxillary palp of Adults appear to prefer white oak (Quercus alba L.) and American three articles, rigid, partially re- beech (Fagus grandifolia Ehrh.). They feed mainly on the bark of tracted into large palpifer; labial shoots, leaf petioles, leaf buds and acorn buds. Eggs are laid in palp of three articles; labrum the ground and larvae feed on the roots of the same host plants. FIGURE 1.130. Ithycerus absent; mandible stout. An- Pupation takes place in the soil. A detailed study of the biology noveboracensis (Forster) (from Bright tennae straight, moniliform; has been published by Sanborne (1981). 1993, reproduced with the permission of the Minister of antennal club compact, of three Status of the classification. Once enigmatic in its place- Public Works and Government articles. Proventriculus without ment there now seems to be a consensus that this species be- Services, 2001) sclerotized plates. longs in a separate and distinct family (Lawrence 1982, Thomp- Notosternal suture com- son 1992, Alonso-Zarazaga and Lyal 1999). Kuschel (1995) how- plete. Procoxae contiguous, mexocoxae narrowly separated and ever, placed it in Curculionidae. Relationships to other metacoxae moderately widely separated. Hind wing with four Curculionoidea are still disputed. anal veins. Legs with trochanter short; triangular; femur attached Distribution. This family contains only the species Ithycerus to side of trochanter. Tarsal claws toothed. Abdomen with noveboracensis (Forster). It is found throughout eastern North ventrites 1 and 2 fused together but separated by distinct suture. America. Cap piece of tegmen strongly bilobed; the median lobe with a broad ventral plate and a narrow dorsal plate. Egg (based on Sanborne 1981) with length 1.2-2.2 mm. CLASSIFICATION OF THE NEARCTIC GENERA Subspherical, longer than wide. External surface of chorion of hexagonal facets with impressed borders. Surface strongly punc- Ithyceridae Schönherr 1823 tate, with numerous aeropyles. Larva (based in part on Sanborne 1981, Lawrence 1991) when Ithycerus Schönherr 1823, 1 sp., I. noveboracensis (Forster 1771), mature about 20-25 mm in length. Body relatively short and eastern United States into extreme southern Canada. Associated broad, subcylindrical, strongly c-shaped. Body yellowish, with with various species of Fagaceae, Betulacae and Juglandaceae. heavily sclerotized mouth frame and mandibles. Vestiture of Pachyrhynchus Kirby 1837 moderately long, scattered setae. Head protracted and hypogna- thous, about as long as wide. Frontal sutures complete, reaching articulating membrane of mandible. Clypeus distinguishable from BIBLIOGRAPHY frons and completely separated from labrum. Three pairs of st- emmata on each side. Labrum free, bearing one sensillum and ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL. 1999. A four pairs of setae. Antenna of a single dome-like article bearing world catalogue of families and genera of Curculionoidea a sensorium, a bifurcate appendage and several setae. Mandible (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae). with two apical teeth, and three blunt teeth on inner edge, acces- Entomopraxis. Barcelona, Spain. sory ventral process and mola absent. Hypopharyngeal bracon present. Maxilla with palp with 2 articles, palpiger present. Labial Family 130. Ithyceridae · 721

KUSCHEL, G. 1995. A phylogenetic classification of SANBORNE, M. 1981. Biology of Ithycerus noveboracensis (Forster) Curculionoidea to families and subfamilies. Memoirs of (Coleoptera) and weevil phylogeny. Evolutionary Mono- the Entomological Society of Washington, 14: 5-33. graphs, 4: 1-80. LAWRENCE, J.L. 1982. Coleoptera. Pp. 482-553. In: S. P. Parker, THOMPSON, R.T. 1992. Observations on the morphology and ed. Synopsis and Classification of Living Organisms. Volume classification of weevils (Coleoptera, Curculionoidea) with a 2. McGraw Hill. New York. key to major groups. Journal of Natural History, 26: 835-891. LAWRENCE, J. L. 1991. Ithyceridae (Curculionoidea). Pp. 590- 591. In: F. W. Stehr, ed. Immature Insects. Volume 2. Kendall/Hunt. Dubuque, Iowa.