Among the weevils, these rarely collected beetles are easily recognized by their straight antennae, and elongate

Transcription

1 692 Family 125. Nemonychidae Superfamily CURCULIONOIDEA 125. NEMONYCHIDAE Bedel 1882 by Robert S. Anderson Family common name: The pine flower snout beetles Among 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 pollenbearing flowers of Pinus species. FIGURE Cimberis compta (LeConte) (from Bright 1993, reproduced with the permission of the Minister of Public Works and Government Services, 2001) Description (based on Lawrence 1982). Shape elongate, slightly convex; length mm; color pale brown to black; vestiture of fine short to moderately long appressed or suberect pubescence. Rostrum moderately to very long and mostly narrow. Antennae straight, ending in a weak, loose club of three articles; antennal insertions lateral at the middle or near the apex of the rostrum. Labrum distinct, not fused with clypeus. Mandibles with a small but distinct mola. Maxillae with separate galea and lacinia and maxillary palps flexible. Labial palps attached ventrally near the base of the prementum. Gular sutures well-developed and separate. Proventriculus lacks sclerotized plates. Procoxae contiguous and the procoxal cavities narrowly closed posteriorly. Mesocoxal cavities either not closed laterally or narrowly so. Elytra without an inner subcostal flange. Hind wing mostly with four anal veins or fewer. Tarsal claws of some cleft. Visible sternites of the abdomen are free; pygydium concealed by the elytra. Tegmen simple or bilobed apically and the median lobe with a distinct dorsal plate. Eggs are undescribed. Larvae (based on Anderson 1991) when mature about mm in length, of moderate thickness throughout length, strongly C -shaped. Body white, covered with mixture of long and short setae. Minute legs present on thorax. Head hypognathous, rounded at sides, pigmented, with few to many setae on frons and epicranium. Frontal sutures complete, reaching articulating membrane of mandible. Clypeus not distinguishable from frons and incompletely separated from labrum. One pair of anterior stemmata. Labrum short, the anterior margin rounded, bearing four pairs of setae. Antenna of a single membranous article bearing an accessory appendage. Mandible with two apical teeth, an obtuse protuberance on cutting edge, a distinctly produced molar area with a flattened grinding surface, and one pair of setae. Hypopharyngeal bracon present. Maxillary palp with three articles, palpiger present or absent. Labial palp of two articles. Premental sclerite present, may be divided medially. Thorax with pronotal sclerite transverse, lightly pigmented or unpigmented, sparsely covered with setae. Legs very small, subconical, of two or three segments, with or without a terminal claw. Abdomen with first eight segments with two dorsal folds and bearing annular or bicameral spiracles. Anal opening terminal. Pupae are undescribed. Habits and habitats. These beetles are rarely collected, likely because of their specialized habits and life history. In North America, adults are found on male flowers of several pine species very early in the season, often while snow is still on the ground. Adults feed on pollen. After mating, females lay eggs in the flowers where the larvae also feed on pollen. Mature larvae drop from the flowers to the ground and pupate in the soil. Pupation can take from a few months to two years. In some instances larvae have been observed feeding on plant parts other than pollen (Thomas and Herdy 1961). In other areas of the world gymnosperms such as Araucariaceae and Podocarpaceae, and some primitive angiosperms (Fagaceae and Ranunculacaeae) also serve as host plants. Whereas pines are the only documented hosts in North America, other genera of conifers may serve as hosts. Status of the classification. This family is bipolar in distribution with approximately similar numbers of taxa found in the southern temperate zones of South America, New Zealand and Australia, and in the northern Holarctic Region. Whereas the family was once considered to be absent in tropical areas, a few species have recently been collected in Panama and Venezuela on Podocarpus and appear to represent an undescribed genus. The North American fauna has recently been revised (Kuschel 1989) and is well-known. A catalog of the North American species was prepared by Hamilton (1994). Distribution. There are 5 genera and 15 species in North America. Two additional species in the genus Atopomacer are known from Pinus at high elevations in far northern Mexico. North American nemonychid species are generally distributed in the

2 Family 125. Nemonychidae FIGURES 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 where pines are present. KEY TO THE NEARCTIC GENERA 1. Second tarsomere truncate at middle, not projected over base of third (Fig. 2); hind tibia with single apical spur; tarsal claw with broad basal flange (Fig. 4); elytra with punctures arranged into indistinct striae (Rhinorhynchinae)... Atopomacer Second tarsomere lobed at middle, projected over base of third (Fig. 3); hind tibia with two apical spurs; tarsal claw simple, lacking broad basal flange (Fig. 5); elytra with punctures not arranged into striae (Cimberidinae) (1). Mandibles inserted dorsally, the sockets fully exposed in dorsal view (Fig. 6); in lateral view, mandibles directed obliquely downwards in relation to rostral plane; antennae situated at middle or after middle of rostrum, as distant from mandibular sockets as combined length of first three articles (Doydirhynchini)... Lecontellus Mandibles inserted laterally, the sockets only partially exposed in dorsal view (Figs. 7-9); in lateral view, mandibles continuous with rostral plane; antennae situated at middle or before middle of rostrum, distinctly closer to mandibular sockets than combined length of first three articles (Cimberidini) (2). Mandibles evenly curved on outer margin, each armed with well-developed tooth on inner margin (Fig. 8)... Cimberis Mandibles angulate on outer margin, unarmed on inner margin (Figs. 7, 9) (3). Labrum trapezoidal or broadly rounded at apex, with six peg-like apical setae in addition to three pairs of dorsal setae (Fig. 7); base of rostrum moderately saddled against obliquely rising frons (Fig. 10) in dorsal view, with 1-3 low carinae flanked with rows of confluent punctures; females with setiferous patches mostly present on one or two ventrites... Pityomacer Labrum triangular, pointed at apex, without any peglike setae in addition to three pairs of dorsal setae (Fig. 9); base of rostrum very deeply saddled against vertically rising frons (Fig. 11) in dorsal view, smooth or sparsely punctate, not carinate; females without setiferous patches on ventrites... Acromacer CLASSIFICATION OF THE NEARCTIC GENERA Nemonychidae Bedel 1882 Cimberidinae Gozis 1882 Cimberidini Gozis 1882 Acromacer Kuschel 1989, 1 sp., A. bombifrons (LeConte 1876), British Columbia, Alberta, Washington, Oregon, California, Idaho and Nevada. Adults collected on Pinus species. Cimberis Gozis 1881, 7 spp., generally distributed. Adults collected on Pinus species.

3 694 Family 125. Nemonychidae Pityomacer Kuschel 1989, 3 spp., British Columbia, Alberta, Washington, Oregon, California and Montana. Adults collected on Pinus species. Doydirhynchini Pierce 1916 Lecontellus Kuschel 1989, 3 spp., Washington, Oregon, California and Nevada. Adults collected on Pinus species. Rhinorhynchinae Voss 1922 Rhinorhynchini Voss 1922 Atopomacer Kuschel 1989, 1 sp., A. ites Kuschel 1989, Colorado and Arizona. Adults collected on Pinus species apparently at high elevations. BIBLIOGRAPHY ANDERSON, D. M Nemonychidae (Curculionoidea). Pp In: F. W. Stehr, ed. Immature Insects. Volume 2. Kendall/Hunt. Dubuque, Iowa. BRIGHT, D. E The Insects and Arachnids of Canada, Part. 21. The Weevils of Canada and Alaska: Volume 1. Centre for Land and Biological Research. Ottawa, 217 pp. HAMILTON, R. W A catalog of the Coleoptera of America north of Mexico. Family: Nemonychidae. USDA Agriculture Handbook , x + 8 pp. KUSCHEL, G The Nearctic Nemonychidae (Coleoptera: Curculionoidea). Entomologica Scandinavica, 20: LAWRENCE, J. L Coleoptera. Pp In: S. P. Parker, ed. Synopsis and Classification of Living Organisms. Volume 2. McGraw Hill. New York. THOMAS, J. B. and H. HERDY A note on the life history of Cimberis elongatus (LeC.) (Coleoptera: Anthribidae). Canadian Entomologist, 93:

4 Family 126. Anthribidae ANTHRIBIDAE Billberg 1820 Family common name: The fungus weevils by Barry D. Valentine Family synonyms: Anthotribidae Gemminger and Harold 1872; Choragidae Kirby 1819; Platyrrhinidae Everts 1903; Platystomidae Pierce 1916; Platystomoidea Pierce These 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 directed anteriad, and the elytra usually have an abbreviated scutellar stria. Description: Shape unusually diverse; elongate and depressed, to oval, convex, and mite-like, mostly more or less elongate and convex above; length 0.4 to 40 mm, in U.S. 0.4 to 16 mm; vestiture rarely absent, of hair-like scales mostly in mixed shades of white, gray, straw, brown or black, in the tropics some with bright pink, red, yellow, or green. In collections, species are FIGURE Toxonotus cornutus often confused with cerambycids, chrysomelids, ciids, (Say) scolytids, and mites. Head large, rarely retractile into prothorax; if rostrum present mostly broad and flattened; surface mostly smooth on vertex, sculpture often progressively heavier on frons and rostral area. Antennae with eleven antennomeres (rarely nine or ten), not geniculate, most with an apical club of three antennomeres, club faint or absent in species with elongate antennae, and involving from two to eight antennomeres in some tropical species; inserted either on the lateral or ventral surface of the rostrum, or on the face between or below the eyes. Labrum distinct, mostly semicircular, setose; mandibles unusually large for a weevil, stout, curved, upper surface flattened and with a setose groove, the apices acute, mostly with a postmedian tooth; maxillary palpi slender, with four palpomeres, the first very short, the fourth long; gular sclerite and sutures invisible; postmentum (fused mentum and submentum) large, deeply emarginate in front, heavily sclerotized; ligula large, corneous, mostly notched to deeply emarginate in front, rarely entire; labial palpi with three palpomeres, the second shortest, both pairs of palpi elongate, cylindrical, flexible, and with acute (very rarely truncate) apices. Eyes dorsal, dorsolateral, or lateral, mostly large, ranging from entire to deeply emarginate, the facets very variable in size and number, and without intermixed setae. Pronotum apex narrower than the maximum head width (except those choragine genera with retractile heads); some exotic genera with the eyes out on broad stalks; base mostly nearly as wide as the elytra, constricted or produced laterad in some; shape mostly trapezoidal, apex obliquely truncate, rarely somewhat produced over the head, base truncate with an antebasal transverse ridge, this mostly turned forward at the sides forming a short or more rarely complete side margin (called the lateral carina), the transverse ridge is always antebasal but it is referred to as basal in those species in which the surface behind it is more or less vertical; surface smooth, punctate, reticulate, rugose, or tuberculate; pleural region broad, supra-coxal sutures well developed; prosternum short to long in front of coxae, the intercoxal process narrow (rarely broader than a coxa); the coxal cavities closed behind. Mesosternum short, the process separating the mesocoxae extremely variable in shape and proportions. Metasternum mostly long, in some so short that meso- and metacoxae are barely separate, a transverse groove or suture near posterior margin, and in many a midventral longitudinal depression or groove. Legs robust to slender; trochantins not exposed; procoxae globular, mostly protruding, not touching or only barely so, and grooved on their inner faces for reception of the intercoxal process of prosternum; mesocoxae globular, separate; metacoxae elongate and transverse (except Sicanthus Valentine and Cisanthribus Zimmerman where they are globular or broadly tear-drop shaped), separate, not reaching elytral margin; trochanters moderate with the apices oblique; femora swollen postmedially, mostly without teeth or spines; tibiae slender, without movable spurs, but with an apical hook or tooth in males of some genera; tarsal formula 5-5-5, apparently 4-4-4, the second apically emarginate, the third deeply bilobed and tomentose beneath, partially embraced by the second, and rarely with the lobes fused, the fourth very small and invisible in dorsal view, the fifth elongate, with claws mostly toothed or cleft. Scutellum small, varied in shape, hidden in some flightless species. Elytral apices always locking into a longitudinal groove in pygidial base, pygidial disc always partially exposed in posterior view but may be invisible from above; normally 10 elytral striae or groups of punctures plus an abbreviated scutellar row, rarely 12, 14 or rows of punctures; epipleural fold present, narrowed or obsolete apically. Wing venation and folding pattern approach that of the Chrysomelidae. Abdomen with five visible sterna, the four sutures entire, fused, and immovable (rarely the fourth suture flexible); surface without coarse sculpturing. Male genitalia with the penis mostly slender, curved, with a movable dorsal plate which when raised gives an open bird s beak appearance, with paired slender basal

5 696 Family 126. Anthribidae struts; parameres and pars basalis fused, the latter with apex mostly hooded, rarely flattened, the ventral struts fused into a single median structure. Female genitalia sclerotized, the valvifers with long apodemes, the coxites apically toothed or ridged, the styli lateral, the spermatheca C-shaped. Larvae are crescent-shaped, subcylindrical, fleshy, widest in the mid-abdominal region; size 4 to 12 mm in length (the majority still undescribed); body with lateral fleshy protuberances and a few to many short or long setae scattered over various segments and the head; color near white. Head exserted, hypognathous, setiferous, or rarely retracted into the prothorax, narrower than prothorax, with epicranial suture present. Antennae reduced, onesegmented or absent. Clypeus transverse; labrum setiferous; mandibles robust, with or without molar areas, bi- or tri-dentate; maxillae with cardo, stipes, three-segmented palpi, the galea setiferous, lacinia acute and inconspicuous; labium with submentum, mentum, ligula, and one-segmented palpi. Ocelli absent. Thorax with legs absent or if present, l-, 2-, or 3-segmented, without tarsunguli. Abdomen nine-segmented, with two or three plicae per segment, the ninth segment smaller than the eighth. Spiracles annular or annular-biforous. Urogomphi absent. Anderson (1947) provides a partial key. Habits and habitats. Distributions and biologies of Nearctic species are summarized by B. D. Valentine (1999). In general, adults may be found on the larval food plants. Those whose larvae feed in the stems or receptacles of various weeds appear to feed upon the pollen of the same plants; species with fungivorous larvae mostly are found feeding on the surface of the same fungi; those with wood-boring larvae occur on dead or dying tree trunks or branches, and in some cases at least feed upon bark. Adults of the majority of anthribid species in the United States are rare in museum collections. The best all-around collecting technique is beating dead or diseased branches, clumps of dead twigs or leaves, or tangles of dead vines. The beating cloth should be as close under the plants as possible and must be examined at once, for anthribids recover and fly off much more rapidly than other weevils. Some of the choragine genera complicate things by being active jumpers, and are as a result exasperatingly difficult to catch. 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 in particular for smooth gray or black patches flush with the wood surface, or small, protruding, black, brown or reddish lumps. These are pyrenomycete fungi, often in the genera Hypoxylon and Biscogniauxia, which are ignored by most collectors; they are very hard and when cut with a knife have a dry crumbly or charcoal-like consistency. All our species of Eurymycter, Piesocorynus, Choragus, and Euxenus are associated with these growths. Sweeping weedy fields usually produces only Trigonorhinus spp., the species T. tomentosus (Say) on common ragweed, T. limbatus (Say) and T. griseus (LeConte) on sneeze- or bitter-weed and other composites, T. alternatus (Say) on fungus-infested morning glory, T. sticticus (Boheman) on smutty grasses and corn, T. rotundatus (LeConte) on smutty grass (Andropogon sp.) and ferns, and the western T. annulatus (Carr) and T. lepidus Valentine on fungus-infested sagebrush. The hosts of T. ornatus (Schaeffer), T. strigosus (Jordan), and T. nigromaculatus (Schaeffer) are not known. Sweeping bushy, overgrown areas will sometimes reveal species of Ormiscus or Eusphyrus, especially in early spring, but beating is better for these and all other members of our fauna except species of Euparius, which occur on polypore fungi. Larvae of all our species feed on vegetation; they can be found in the twigs and branches of trees, in hard or polypore fungi, or under bark of dead or dying trees. One species, the coffee bean weevil, Araecerus fasciculatus (DeGeer 1775), lives in seeds and all sorts of dried plant materials from banana flour to strychnine. A related species in Australia and New Zealand occurs on dried fruit, especially apples. Our introduced species of the European genus Anthribus (formerly Brachytarsus) have larvae that feed on scale insects of the subfamily Lecaniinae; the female lays each of her eggs under a female scale that has laid her eggs, the weevil larvae then develop exclusively at the expense of the eggs of the scale insect. This is a startling deviation from the phytophagous habits of the Rhynchophora. Status of the classification. This family has been subject to considerable nomenclatorial confusion and lack of studies of higher taxa beyond all proportions consistent with the rest of the order. The genera need considerable study. The world catalogues of Wolfrum (1929, 1953) need many additions and revisions. The Nearctic fauna is summarized by Valentine (1999) who provides a synonymic checklist and keys to most genera. Other basic references are LeConte (1876), Blatchley and Leng (1916), Pierce (1930), Ting (1936), and Valentine (1960, 1971, 1972, 1991). Distribution. There are about 360 genera and 4,000 species known from all regions; 88 described and 32 undescribed species occur in North America. Of 30 North American genera, Araecerus is cosmopolitan; Choragus, Eurymycter, Gonotropis, Allandrus, and Trigonorhinus are holarctic (although the latter extends south to Argentina); Sicanthus is endemic but probably neotropical; Euxenulus, Phoenicobiella, and Araeoderes appear to be North American but are obviously related to Latin American groups; and all remaining genera are Neotropical, forming the northern fringe of an extensive Antillean, Central, or South American fauna. KEY TO THE NEARCTIC TRIBES AND GENERA 1. Antennae inserted on anterior surface of rostrum or head (Fig. 2); antennal club never with four antennomeres (Choraginae)... 2 Antennae inserted on lateral or ventral surface of rostrum (Fig. 3); or antennal club sometimes with four antennomeres. (Anthribinae) (1). Eyes rounded, upper edges not closer together than lower... 3 Eyes elongate-oval, upper ends closer to each other than lower (Choragini) (2). Hind coxae elongate-transverse, almost reaching elytral edge (Araecerini)... 4 Hind coxae globular or short tear-drop shaped (Cisanthribini)... Sicanthus

6 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 (4). Lateral prothoracic carina present; transverse pronotal carina basal... Araecerus Lateral prothoracic carina absent; transverse pronotal carina antebasal...neoxenus 6(4). Lateral prothoracic carina double, not upturned at apex; elytra with rows of punctures Habroxenus Lateral prothoracic carina single, mostly with apex upturned; elytra with rows of punctures.... Acaromimus 7(2). Head not retractile into prothorax, the eyes too wide; pronotum with a raised reticulum forming a honeycomb pattern... 8 Head retractile into prothorax past the eyes; pronotum punctate, not reticulate (7). Antennae with 11 antennomeres... Choragus Antennae with 9 or 10 antennomeres Pseudochoragus 9(7). Elytra with 10 rows or fields of punctures, plus a short scutellar row... Euxenus Elytra with 11 to 15 rows of punctures, one or more mostly incomplete... Euxenulus 10(1). Each elytron partially enclosing anterior part of scutellum, resulting scutellar notch shaped like an inverted omega (Fig. 4); eyes large, entire, very finely faceted (Gymnognathini) Gymnognathus Scutellar notch forming a simple V or U; eyes as above, or small, or notched, or coarsely faceted (10). Rostrum with a dorsal, median carina interrupted at base by an abrupt, small, very deep pit or transverse groove (Platystomini) Rostrum carinate or not, at most with a basal puncture (11). Lobes of all third tarsomeres separate.. Toxonotus Lobes of third tarsomeres fused... Phoenicobiella 13(11). Mandibles with strongly toothed ventral cutting edge (Fig. 3), as well as normal dorsal edge (Cratoparini)... Euparius Mandibles without a toothed ventral cutting edge (13). Eye entire, or truncate, or faintly sinuate on anterior face Eye strongly notched or emarginate on anterior face (nearest insertions of antennae) (14). Entire face with conspicuous white pubescence; center of pronotum with a pit or groove; our species with a conspicuous patch or band of white crossing the suture (Tropiderini) FIGURES Euxenus sp., head, anterior view; 3. Euparius marmoreus (Olivier), head, dorsal view; 4. Gymnognathus sp., scutellum, dorsal view. White pubescence, if present, confined to spots, if conspicuous, pronotum will have the disc concave with a central swelling, and elytra will have multiple tufts of erect pubescence (15). Pronotum with a smooth, longitudinal, shallow groove from central pit to antebasal carina; elytral pale patch antemedian and not reaching side margins... Gonotropis Pronotum with a sinuous, transverse, median groove; elytral pale band postmedian and reaching the side margins... Eurymycter 17(15). Antennal club with 4 antennomeres; rostrum thick, width at apex less than twice depth; scrobes dorso-lateral, interscrobal distance mostly less than interoccular distance (Discotenini) Discotenes Antennal club with 1 or 3 antennomeres; rostrum depressed apically, width at apex more than twice depth; interscrobal distance more than interoccular distance (17). Eyes finely faceted, more than 26 rows across maximum width Eyes coarsely faceted, 26 or fewer rows across maximum width (Piesocorynini) (18). Sides of rostral apex abruptly widened to cover the laterally protruding mandibular bases; our species with a small post-ocular tooth on apex of prothorax (Ischnocerini)... Ischnocerus Sides of rostral apex with mandibular sheaths weakly or not wider than rostral dorsum; no small post-ocular teeth on prothoracic apex (19). Merger of rostrum with venter of head capsule forming a broad curve; rostrum long, flattened, and apically flared Merger of rostrum with venter of head capsule indicated by a transverse groove or abrupt angle; rostrum shorter, not apically flared (Platyrhinini) (20). Antennae with whorls of long, erect setae; length (head excluded) more than 5 mm (Stenocerini).... Stenocerus Antennae without whorls of long erect setae; length (head excluded) less than 4 mm (Allandrini) Allandrus 22(20). Face with a pair of abrupt depressions between the upper ends of the eyes; lateral prothoracic carina with an apical tubercle... Trachytropis Face with at most a weak concave area at rostral base; lateral prothoracic carina not toothed at apex... Goniocloeus

7 698 Family 126. Anthribidae 23(18). Eyes with 14 or more rows of facets across maximum width... Piesocorynus Eyes with 12 or fewer rows of facets across maximum width... Brachycorynus 24(14). All third tarsomeres with the lobes fused down the midline (Anthribini)... Anthribus All third tarsomeres with the lobes separate (24). Rostrum quadrate, or with rounded apical angles Rostrum, excluding mandibles, narrowed from base to apex; rostral apex with central third longer than sides (Trigonorhinini)... Trigonorhinus 26(25). Intercoxal process of mesosternum laterally angulate, or swollen, or bent; scrobes sulciform, continued across the rostral sides and ending below the eyes (Basitropidini) Intercoxal process of mesosternum simple; scrobes foveiform, more or less rounded and not continued transversly or ventrally around and under the rostrum (Zygaenodini) (26). Lateral prothoracic carina extending to apex where it is toothed; antennal scrobes strongly produced toward ventral midline... Eugonus Lateral prothoracic carina not reaching anterior margin, nor apically toothed, antennal scrobes widely separated on venter... Phaenithon 28(26). Lateral prothoracic carina absent or if present not reaching anterior margin Lateral prothoracic carina extending to apex Araeoderes 29(28). Transverse pronotal carina clearly antebasal, incapable of contacting the elytral base; pronotal hind angles not projecting laterad of the humerae Ormiscus Transverse pronotal carina subbasal or basal, capable of contacting elytral base at some point; pronotal hind angles often projecting laterad of the humerae... Eusphyrus CLASSIFICATION OF THE NEARCTIC GENERA Suprageneric taxa are diagnosed in the generic key. NOTE: Many undescribed species are included in the comments on distribution. As used below, the designation Neotropical includes Antillean species; Central and South America do not. Anthribidae Billberg 1820 Choraginae Kirby 1819 Araecerini Lacordaire 1876 Araecerus Schoenherr 1823, 2 introduced spp., including the coffee bean weevil, A. fasciculatus (DeGeer 1775): Worldwide. Also about 75 Indopacific species. Araeocerus Schoenherr 1839 Araeocorynus Jekel 1855 Araeosarus Walker 1859 Doticus Pascoe 1882 Metadoticus Olliff 1890 Neoxenus Valentine 1999, 1 sp., N. versicolor Valentine, 1999, Texas to Panama; four others in Central America and Antilles. Habroxenus Valentine 1999, 1 sp., H. politus Valentine, 1999, Maryland, Texas; four others in Central America and Antilles. Acaromimus Jordan 1907, 1 sp., A. americanus (Motschoulsky 1873), Florida, Alabama, and Texas; six others in Central America and Antilles. Acaropsis Jordan 1907 Xenorchestes Motschulsky 1873, not Wollaston 1854 Euxenus Blatchley 1920, not LeConte 1876 Cisanthribini Zimmerman 1994 Sicanthus Valentine 1999, 1 sp., S. rhizophorae Valentine 1999, Florida keys. Choragini Kirby 1819 Choragus Kirby 1819, 6 spp., eastern United States; 45 others almost worldwide. Alticopus Villa and Villa 1833 Pseudochoragus Petri 1912, 1 sp., P. nitens (LeConte 1884), Massachusetts and Oklahoma; another species in Europe. Choragus, of American authors, in part, not Kirby 1819 Holostilpna, of American authors, not Jordan 1907 Euxenus LeConte 1876, 3 spp., eastern United States; at least six more in Central America and Antilles. Holostilpna Jordan 1907 Euxenulus Valentine 1960, 1 sp., E. piceus (LeConte 1878), south Florida; also three undescribed in Antilles. Anthribinae Billberg 1820 Discotenini Lacordaire 1866 Discotenes Labram and Imhoff 1841, 2 spp., Texas; Arizona about 10 more south to Brazil. Phanosolena Schaeffer 1904 Ischnocerini Lacordaire 1866 Ischnocerus Schoenherr 1839, 3 spp., Maryland to Texas; Arizona; 10 more in Neotropics. Meconemus Labram and Imhoff 1839

8 Family 126. Anthribidae 699 Allandrini Pierce 1930 Allandrus LeConte 1876, 3 spp., United States except the desert southwest; also several in Palaearctic. Tropiderinus Reitter 1916 Stenocerini Kolbe 1897 Stenocerus Schoenherr 1826, 1 sp., S. longulus Jekel 1855, south Texas to Brazil; seven more Mexico to Argentina. Gymnognathini Valentine 1960 Gymnognathus Schoenherr 1826, 2 spp., south Texas; south Arizona; about 90 described and many undescribed Neotropical species. Analotes Schoenherr 1839 Tropiderini Lacordaire 1866 Gonotropis Leconte 1876, 1 sp., G. gibbosus LeConte 1876, Canada south to Pennsylvania and Colorado; plus three Holarctic. Tropideres, of European authors and Valentine 1960, in part, not Schoenherr 1823 Eurymycter LeConte 1876, 3 spp., United States except the desert southwest. Tropideres, of Valentine 1960, not Schoenherr 1923 Piesocorynini Valentine 1960 Piesocorynus Dejean 1834, 5 spp., eastern United States; Arizona; about 30 others in Neotropics and Antilles. Piezocorynus Schoenherr 1839 Camptotropis Jekel 1855 Brachycorynus Valentine 1999, 3 spp., eastern United States to Texas; a few others in Central America and Antilles. Tropideres, of LeConte 1876 and others, not Schoenherr1823 Brevibarra, of Valentine 1960, not Jordan 1906 Platyrhinini Imhoff 1856 Goniocloeus Jordan 1904, 1 sp., G. bimaculatus (Olivier 1795), eastern United States to Texas; 36 others south to Brazil. Tropideres, of various authors, in part, not Schoenherr 1823 Strabus Jekel 1860, not Gerstaecker 1860 [Curculionidae] Trachytropis Jordan 1904, 1 sp., T. arizonicus (Sleeper 1954), southeastern Arizona; two others in Central America. Platystomini Pierce 1916 Phoenicobiella Cockerell 1906, 2 spp., South Carolina to Lousiana, Florida and Bahamas; south Texas; a third species in Cuba. Phoenicobius LeConte 1876, not Moerch 1852 [Mollusca] Toxonotus Lacordaire 1866, 6 spp., eastern United States to New Mexico and Arizona; about 35 others throughout the Neotropics. Anthribus, of authors, in part, not Geoffroy 1762 Neanthribus Jordan 1906 Pseudanthribus Pierce 1930 Basitropidini Lacordaire 1866 Eugonus Schoenherr 1833, 1 sp., E. bicolor Valentine 1972, southeastern Arizona; about 30 more in Central and South America. Schimatocheilus Fahraeus 1839 Phaenisor Motschoulsky 1874 Phaenithon Schoenherr 1823, 1 sp., P. platanum (Schaeffer 1906), southeastern Arizona; about 65 others in Central and South America. Camaroderes Jekel 1855 Griburiosoma Schaeffer 1906 Zygaenodini Lacordaire 1866 Ormiscus G. R. Waterhouse 1845, 14 described and about 30 undescribed spp., eastern United States and desert southwest; probably 200 more throughout the Neotropics, mostly undescribed. Entomops Lacordaire 1866 Hormiscus Gemminger and Harold 1872 Toxotropis LeConte 1876 Gonops LeConte 1876 Eusphyrus LeConte 1876, 6 spp., eastern United States and desert southwest; 30 more in Central and South America. Opisthotropis Hoffmann and Tempere 1954 Araeoderes Schaeffer, 1906, 1 sp., A. texanus Schaeffer, 1906, southern Texas and southern Alabama. Trigonorhinini Valentine 1999 Trigonorhinus Wollaston 1861, 11 spp., southern Canada, entire United States; seven others south to Argentina, and in the Palearctic Region. Brachytarsus, of many authors, not Schoenherr 1823 Trigonorrhinus Gemminger and Harold 1872 Anthribulus LeConte 1876 Brachytarsoides Pierce 1930 Cratoparini LeConte 1876 Euparius Schoenherr 1823, 5 spp., eastern United States to Montana and Arizona; at least 70 others in Central and South America, and Japan to Australia. Cratoparis Dejean 1834 Caccorhinus Sharp 1891

9 700 Family 126. Anthribidae Anthribini Billberg 1820 Anthribus Geoffroy 1762, 2 adventive spp. from Europe, Atlantic coast and?california. Brachytarsus Schoenherr 1823 Anthotribus Gemminger and Harold 1872 Pseudobrachytarsus Pierce 1930 BIBLIOGRAPHY ANDERSON, W. H Larvae of some genera of Anthribidae (Coleoptera). Annals of the Entomological Society of America, 40: , pl. I-IV. BLATCHLEY, W. S. and C. W. LENG Rhynchophora or Weevils of North Eastern America. The Nature Company. Indianapolis, 682 pp., fig LECONTE, J. L The Rhynchophora of America, North of Mexico. Proceedings of the American Philosophical Society, XV (96): i-xvi PIERCE, W. D Studies of the North American weevils of the superfamily Platystomoidea. Proceedings of the United States National Museum, 77(no. 2840): 1-34, pls TING, P. C The mouthparts of the coleopterous group Rhynchophora. Microentomology, 1: VALENTINE, B. D The genera of the weevil family Anthribidae north of Mexico (Coleoptera). Transactions of the American Entomological Society, 86: VALENTINE, B. D Family Anthribidae. Pp In: M. H. Hatch. Beetles of the Pacific Northwest, Part V. University of Washington Publications in Biology, 16: i-xvi pl. I-LV. VALENTINE, B. D Notes on anthribid weevils. III. New species and records primarily from Arizona. Coleopterists Bulletin, 26: 1-11, fig.1-4. VALENTINE, B. D The Choragus-Holostilpna problem (Coleoptera: Anthribidae). Coleopterists Bulletin, 45: VALENTINE, B. D. 1998(1999). A review of Nearctic and some related Anthribidae (Coleoptera). Insecta Mundi, 12: WOLFRUM, P Anthribidae. Coleopterorum Catalogus, 26(102): WOLFRUM, P Anthribidae. Coleopterorum Catalogus, supplementum, 26(102): 1-63.

10 Family 127. Belidae BELIDAE Schönherr 1826 by Robert S. Anderson Family common name: The cycad weevils The only North American members of this family are odd weevils, recognized by their straight antennae, somewhat truncated elytra mostly exposing the last one or two tergites, and the short stout legs with femora which are 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. FIGURE Rhopalotria slossonae (Schaeffer) (from Bright 1993, reproduced with the permission of the Minister of Public Works and Government Services, 2001) Description. (based on Lawrence 1982). Shape elongate, slightly convex; length 3-6 mm; color pale to dark brown or black; vestiture of very fine short appressed pubescence. Rostrum moderately to very long and mostly narrow. Antennae straight, ending in a weak, loose club of three articles; articles 9 and 10 with deep apical pockets; antennal insertions ventral at or near the base of the rostrum. Labrum fused with clypeus. Labial palps with two articles and attached dorsally near the apex of the prementum. Gular sutures fused. Proventriculus lacks sclerotized plates. Pronotum with complete lateral edges. Procoxae contiguous and the procoxal cavities narrowly closed posteriorly. Elytra without an inner subcostal flange. Hindwing with fewer than five anal veins. Tarsal article 2 broadly lobed as article 3; tarsal claws simple. Visible sternites of the abdomen free; pygydium exposed by the elytra. Tegmen truncate or slightly emarginate apically and the median lobe with a distinct dorsal plate. Eggs are undescribed. Larvae (based in part on van Emden 1938, Lawrence 1982, Crowson 1986) short, broad, ventrally curved and subglabrous. Body lightly sclerotized. Legs absent. Head strongly retracted, endocarina indistinct, with very short antennae of one article and no epipharyngeal rods. Frontal sutures incomplete, not reaching articulating membrane of mandible. Maxilla with palp with two articles, palpiger absent. Thoracic spiracles on mesothorax. Abdomen with segments with two dorsal folds. Anal opening ventral. Pupae are undescribed. Habits and habitats. Within this family, only the genus Rhopalotria occurs in North America. It is restricted to south Florida where two species are associated with native and adventive species of Zamia (Cycadaceae). The biology has been well-studied by Norstog and Fawcett (1989). Adult weevils swarm on male cones of the cycads, where mating, feeding and oviposition occur. Larvae feed within the male cones. Weevils also visit female cones but do not feed. Pollen transport to the female cones occurs during these visits and these beetles (and their relatives elsewhere) may be obligate pollinators of the cycads. Status of the classification. The family-level classification of these beetles is somewhat controversial. Crowson (1986) considered them as a distinct family, the Allocorynidae. Kuschel (1995) places them as Belidae, subfamily Oxycoryninae. The most recent catalog (Alonso-Zarazaga and Lyal 1999), probably following Lawrence (1982) and Thompson (1992), places them as the subfamily Allocoryninae of the Oxycorynidae. Here they are placed as Belidae, subfamily Allocoryninae. Distribution. There is one genus with two species in North America. Both species occur only in Florida; one is adventive and one native. CLASSIFICATION OF THE NEARCTIC GENERA Belidae Schönherr 1826 Allocoryninae Sharp 1890 Rhopalotria Chevrolat 1878, 2 species, R. mollis (Sharp 1890), Florida (adventive on Zamia furfuracea L.) and R. slossonae (Schaeffer 1905), Florida (native on Zamia integrifolia L., coontie). Allocorynus Sharp 1890 BIBLIOGRAPHY ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL A world catalogue of families and genera of Curculionoidea (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae). Entomopraxis. Barcelona, Spain. CROWSON, R. A On the systematic position of Allocoryninae. Coleopterists Bulletin, 40: EMDEN, F. VAN On the taxonomy of Rhynchophora larvae (Coleoptera). Transactions of the Royal Entomological Society of London, 87: 1-37.

11 702 Family 127. Belidae KUSCHEL, G A phylogenetic classification of Curculionoidea to families and subfamilies. Memoirs of the Entomological Society of Washington, 14: LAWRENCE, J. L Coleoptera. Pp In: S. P. Parker, ed. Synopsis and Classification of Living Organisms. Volume 2. McGraw Hill. New York. NORSTOG, K. J. and P. K. S. FAWCETT Insect-cycad symbiosis and its relation to the pollination of Zamia furfuracea (Zamiaceae) by Rhopalotria mollis (Curculionidae). American Journal of Botany, 76: THOMPSON, R. T Observations on the morphology and classification of weevils (Coleoptera, Curculionoidea) with a key to major groups. Journal of Natural History, 26:

12 Family 128. Attelabidae 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 The members of this family are considered to be primitive weevils based on the straight antennae (non-geniculate), mouthpart structure, wing venation, etc. The family in the inclusive sense contains three subfamilies in America 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 following adult characters have been given by Thompson (1992) and Kuschel (1995) to support the single family status of Attelabidae: maxillary palps 3 or 4 segmented (if 3 segmented, elytral punctuation confused); tergite 8 concealed in both sexes; elytra separately and broadly rounded at apices, more or less exposing a pygidium (tergites 6 and 7); body setose, without broad scales; cuticle metallic or otherwise brightly colored in many; elytra with scutellary striole in many; abdominal ventrites decreasing in length, first 3 or 4 fused; pygidium visible in both sexes; male genitalia consisting of a free pedon and tectum and a FIGURE Merhynchites bicolor large tegmen (as in all groups (Fabricius) (from Bright 1993, with non-geniculate antennae); reproduced with the permission of intersegmental membrane between sternites 8 and 9 partly the Minister of Public Works and Government Services, 2001) or entirely sclerotized. The subfamily Attelabinae is characterized by the following: Adult with body more or less short, stout, glabrous or with appressed pubescence above, color variable; head rectangular to weakly triangular, elongate in males of some; eyes large, mostly reniform, longest diameter more or less vertical; rostrum short to moderate in length, distinctly widened apically, weakly arcuate in lateral view; lateral apical angle in some with tooth-like projection; postmentum in some males with pair of acuminate ventral projections; antennae inserted dorsolaterally, club 3-segmented, more or less compact, scrobe short, distinct, more or less vertical; mandibles robust, toothed on inner apical margin, outer margin more or less rounded; maxillary palps 4-segmented; labial palps indistinctly 1 or 2 segmented, located ventrally on prementum; elytra more or less quadrate, not much longer than wide, humeri simple to strongly protuberant, scutellary striole present or absent, striae becoming less distinct posteriorly in many; ventral abdominal sutures 1-3 more or less rigid; procoxae prominent, conical; prothoracic legs enlarged, profemora distinctly swollen, armed or unarmed; protibia longer, narrower and more arcuate in males; tibial apices uni-uncinate in males and bi-uncinate in females; tarsal claws connate. Larva with thoracic spiracle located in mesothorax or in intersegmental fold between prothorax and mesothorax; prementum and mentum fused, equally sclerotized; maxillary palps 2-segmented; prodorsal fold more convex than postdorsal fold, postdorsal fold sloping posteriorly; abdominal tergum 9 at least twice as long as sternum 9; anus subterminal or ventral; cuticular asperities more or less inconspicuous. The subfamily Rhynchitinae is characterized by the following: Adult with body more or less elongate, mostly with distinct semi-erect to erect setae above, color variable; head mostly triangular, widest at the base; eyes small to medium in size, rounded, longest diameter more or less horizonal; rostrum mostly slender, parallel-sided and longer than the head, more or less weakly arcuate in lateral view; antennae inserted laterally, club 3-segmented, more or less loosely united, scrobes shallow, longitudinally oblique, not well defined; mandibles flat, toothed on inner and outer margins; maxillary palps 4 segmented; labial palps 2 or 3 segmented, inserted laterally or apically on prementum; elytra elongate, mostly distinctly longer than wide, humeri simple, scutellary striole present or absent, striae mostly distinct through-

13 704 Family 128. Attelabidae out; ventral abdominal sutures distinct, suture between 1 and 2 rigid; all legs more or less equally developed; tibial apices mostly with one or two small straight mucros; tarsal claws with long clawlike inner processes; in some inner processes shorter, wider and more toothlike. Larva with thoracic spiracle located on postero-lateral portion of prothorax; prementum and mentum separate, mostly unequally sclerotized; maxillary palp 2 or 3 segmented; prodorsal and postdorsal folds (segments 6-8) more or less subequal in convexity; tergum and sternum of abdominal segment 9 subequal in length; anus terminal or subventral, transverse x-shaped or simple transverse cleft; cuticular asperities more or less conspicuous. The subfamily Pterocolinae is characterized by the following: Adult with body robust and dorsally convex, with appressed to semi-erect inconspicuous fine setae above, coloration metallic bluish-green to bluish-black; head weakly triangular; eyes large, oval, posteriorly more protuberant, anteriorly emarginate; rostrum short, as long as or shorter than the head, straight, in dorsal view more or less parallel-sided throughout, apically depressed, with beard like ventral setosity (more setose in males); antennae short, subequal to combined length of head and rostrum, inserted near basal 1/4 under basirostral ridge; scrobe fossa like; club large, strongly abrupt, compact; mandibles flat, toothed on inner and outer margins; maxillary palps 4-segmented, labial palps 3-segmented, inserted apically on prementum; pronotum with distinct lateral carinae; elytra short, apices individually rounded; pygidium and two complete abdominal terga visible; humeri simple; scutellary striole present; striae mostly distinct throughout; intervals more or less convex; all ventral abdominal sutures entire; mesepimeral side pieces strongly developed, visible in dorsal view between pronotum and elytra; front coxae small, globose, distinctly separated; middle and hind coxae widely separated; middle and hind femora larger than front femora; tibial apices with two blunt-tipped mucros; tarsal claws with broad basal tooth. Larva is undescribed. Attelabid eggs that have been examined are oval, creamy white to yellowish-white, shiny and without surface sculpture (Balduf 1959; Hamilton 1980, 1983, 1994). Only a few North American pupae in the subfamily Rhynchitinae have been described (Hamilton 1980, 1983; Hamilton and Kuritsky 1981). They possess taxonomically significant tuberculate setae and posterior processes associated with abdominal segment 9. Habits and habitats. All known members of the subfamily Attelabinae are leaf rollers and apparently cause no serious damage to their hosts although one Central American species of Hybolabus has been reported as a serious defoliator of its host tree, Cariniana pyriformis Miers (Lecythidaceae). All known females lay their egg(s) on leaves that they prepare by biting and cutting with their mandibles. The leaves are then rolled into a more or less barrel-shaped structure (nidus) that nourishes and protects the developing larvae. The enlarged front legs and the uncinate tibial apices manipulate the leaf tissue during nidus formation. Pupation occurs in the ground in the species that have been studied. Comments on the biology of three North American species are available. Packard (1890), Frost (1908), Blatchley and Leng (1916), and Lutz (1935) commented on the biology of Himatolabus pubescens (Say). Frost (1908) recorded a eulophid parasite bred from Attelabus rhois Boheman (=H. pubescens). Loding (1945) commented on the biology of Homoeolabus analis (Illiger). Murtfeldt (1872), Packard (1890), Girault (1904), and Edwards (1949) treated aspects of the biology of Attelabus bipustulatus Fabricius. Most of the information in these papers deals with taxonomy, host plants, nidus formation and distributions. Hopkins (1905) reported a trichogrammid egg parasite of A. bipustulatus. Van Emden (1938) included the larvae of H. pubescens, Homoeolabus analis and Attelabus nigripes LeConte in a key to the genera of Attelabini. Also, he associated the niduses of these species, and A. bipustulatus as well, with specific host plants. Vogt (1992) discussed the leaf rolling behavior, host plants and associated rhynchitid weevils (Pterocolinae) of attelabine weevils occurring from Canada to the Republic of Panama. Hamilton(1998), in a revision of the New World Pterocolinae, described 15 new species from Central and South America and clarified many attelabid-pterocoline-host plant associations initiated by Vogt. Hamilton (1994) provided a summary of known North American rhynchitine biology. The larvae of these weevils develop in living or dead leaves as leaf miners, in fruits, in cut flower heads, in cut terminal shoots, in terminal buds, in flower buds, and in cut leaf primordia. The cuts are made by the adult female with her mandibles after oviposition in the specific plant part. The flower heads, buds or leaf primordia are either completely cut from the plant or are partially cut and eventually drop to the ground. Information has been published on the life stages or life cycles of only eight North American rhynchitine weevils - Haplorhynchites aeneus (Boheman), Merhynchites bicolor (Fabricius), Merhynchites wickhami (Cockerell), Eugnamptus angustatus (Herbst), Auletobius cassandrae (LeConte), Temnocerus perplexus (Blatchley), Temnocerus naso (Casey) and Deporaus glastinus (LeConte). Boving and Craighead (1931) provided the first basic illustrations of H. aeneus larvae. Hamilton (1973, 1981) provided detailed illustrations of life stages, host plants, and behavior for H. aeneus. Chittenden (1901), Dickerson (1910), Blatchley and Leng (1916), Ewing (1915), and Essig (1958) comment on the biology of the eastern rose curculio, M. bicolor. Balduf (1959) published the most comprehensive study on the biology of M. bicolor to date. Hamilton and Kuritsky (1981) commented on the life cycle of M. bicolor and provided detailed descriptions of the larva and pupa. Cooley (1903), Lovett (1915), Robertson (1923), and Hoerner (1936) comment on the life cycle of the western rose curculio, M. wickhami. Boving and Craighead (1931) illustrated what appears to be the larva of E. angustatus but they did not rear it and they determined it as Orsodacne sp. by the method of elimination and locality. Hamilton (1980) provided notes on the biology of E. angustatus (Herbst) and described and illustrated the larva and pupa. Hamilton (1983) discussed the life cycles of A. cassandrae and T. perplexus and provided detailed descriptions and illustrations of their immature stages. Hamilton (1994) provided new

14 Family 128. Attelabidae 705 life cycle data for T. naso and D. glastinus on Quercus wislizenii A.D.&C. in southern California. Published biological information on other rhynchitid species is lacking except for brief comments by Kissinger (1964), given him by Vogt, stating that Eugnamptus spp. mine the dead leaves of various hardwoods and that females of Temnocerus aeratus (Say) lay eggs in and cut terminal oak twigs in which the larvae subsequently develop. Kissinger also indicated that, again according to Vogt, Involvulus hirtus (Fabricius) has habits similar to T. aeratus.van Emden (1938) included three North American species in a paper on the taxonomy of Rhynchophora larvae. He provided key characters for the larvae of H. aeneus, M. bicolor and Rhynchites velatus LeConte and included them in a key to select species of Rhynchitini. He also associated these three species with host plants and localities. Based on his host plant data, he examined the larva of H. aeneus not Rhynchites (Involvulus) hirtus (Fabricius) as listed. Haplorhynchites aeneus, M. bicolor and M. wickhami are the only North American rhynchitine species of known economic importance. Haplorhynchites aeneus attacks commercially grown sunflowers while M. bicolor and M. wickhami attack cultivated roses and sometimes damage blackberry and raspberry. Comments on the economic importance of M. bicolor have been made by Harris (1862), Chittenden (1901), Gates (1909), Dickerson (1910), Blatchley and Leng (1916) and Essig (1958). Cooley (1903), Lovett (1915), Robertson (1923), and Hoerner (1936), give information on the economic importance and control of M. wickhami. Schulz and Lipp (1969) and McBride and Oseto (1978) commented on the status of damage to sunflowers by H. aeneus. Vogt (1992) discussed the biology of the Pterocolinae and their attelabid hosts and coined the term thief weevils for their habit of taking over the niduses of their attelabid hosts. Although not specifically stated, his comments on the biology of these weevils were probably based mainly on observations of P. ovatus in Maryland. According to Vogt, female pterocolines force their way into freshly made attelabid leaf rolls where they eat or destroy the host egg and oviposit their own. The pterocoline larva develops rapidly feeding on the decaying leaf tissue and leaves the hollowed out leaf roll to pupate in the ground. Status of the classification. The subfamilies Rhynchitinae and Pterocolinae, here considered in the family Attelabidae, have been placed in the family Rhynchitidae by other workers. The family name Rhynchitidae is credited to Gistel 1856 (Alonso- Zarazaga and Lyal 1999). John L. LeConte (1876) was the first American entomologist to use the family name Rhynchitidae for weevils in the subfamilies Rhynchitinae and Pterocolinae. Subsequent workers (Sharp 1889, Pierce1909, Blatchley and Leng 1916, Ting 1936, Anderson 1991) grouped these weevils in the family Curculionidae. Pierce (1913) recognized the family Attelabidae including the Rhynchitinae and the new tribes Rhynchitini and Auletini. Voss ( ), in a worldwide taxonomic monograph series on the subfamilies Attelabinae, Rhynchitinae and Pterocolinae, consistently listed these groups in the family Curculionidae. Boving and Craighead (1931), in a synopsis of the larval forms of Coleoptera placed the Rhynchitinae and Attelabinae in the family Attelabidae. Crowson (1955) grouped these weevils in the family Attelabidae but suggested a possible alternative would be to establish a family Rhynchitidae for the Rhynchitinae and Pterocolinae. Lawrence (1982) also placed these weevils in the family Attelabidae. O Brien and Wibmer (1982), in an annotated checklist of North American weevils, recognized the families Rhynchitidae and Attelabidae but Wibmer and O Brien (1986), in their annotated checklist of South American weevils, regrouped these weevils into the single family Attelabidae. Thompson (1992) using abdominal characters, etc. lumped the three subfamilies in the family Attelabidae. More recently Kuschel (1995), in a phylogenetic approach, recognized only the family Attelabidae and Farrell (1998), in a molecular approach, also grouped the rhynchitines and attelabines in the family Attelabidae. Hamilton (1969 to 1998) has consistently used the family Rhynchitidae for the Rhynchitinae and Pterocolinae. Most recently, Alonso-Zarazaga and Lyal (1999) have recognized the family Rhynchitidae in their world catalogue of families and genera of Curculionoidea. The family name Attelabidae is credited to Billberg (1820). The genus Attelabus, on which the family name is based, was originally described by Linnaeus (1758). Linnaeus included only A. coryli Linnaeus in the genus and it was therefore the type by monotypy. Since the time of Linnaeus, many species from all over the world have been added to the genus Attelabus. Olivier (1807) decided that A. coryli was generically different from the other species that had been placed in Attelabus and described the new genus Apoderus with 13 species including A. coryli. Attelabus coryli is technically the type of Attelabus but was not recognized by Olivier, or anyone else, until Bedel (1888) pointed out that Attelabus should be used for those species placed by Olivier in Apoderus. Bedel proposed the name Cyphus for the species that were left in Attelabus by Olivier as well as many other species placed there by subsequent authors. Bedel s proprosal followed the International Code but subsequent workers, unaware of the change or unwilling to change the names of numerous species, have not followed Bedel. Silfverberg (1977) successfully appealed to the International Commission on Zoological Nomenclature to accept common usage of Attelabus in the interest of stability and to confirm the designation by Schoenherr (1823) of Attelabus curculionoides Linnaeus 1767 (= Curculio nitens Scopoli 1763) as typespecies. Two major groups of attelabids are now officially recognized - Apoderus with A. coryli as its type and Attelabus with Attelabus nitens Scopoli as its type. Jekel (1860) divided the genus Attelabus into 16 subgenera and Voss (1925) elevated most of them to the generic level. The genus Attelabus is applicable to only two species of attelabids in America, north of Mexico - A. bipustulatus Fabricius and A. nigripes LeConte. Distribution. The family includes 1,914 world species in 97 genera (Kuschel 1995). These weevils occur throughout the world but Kuschel points out that they do not occur in New Zealand, New Caledonia and the Pacific Islands. In the New World, 362 species have been described including 180 species of leaf rolling weevils in four subfamilies (Attelabinae, Euscelinae, Hybolabinae and Pilolabinae), 162 species of Rhynchitinae and 20 species of Pterocolinae. In America north of Mexico, there are 51 total species in 11 genera including 6 species of Attelabinae, 44 species of Rhynchitinae and one species of Pterocolinae.

15 706 Family 128. Attelabidae KEY TO NEARCTIC SUBFAMILIES AND GENERA 1. Tarsal claws appendiculate (Figs. 7 and 8); mandibles 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 Tarsal claws connate (Fig. 9); mandibles robust, not toothed on outer margin; front legs enlarged, distinctly larger than middle and hind legs; profemora distinctly swollen (Fig. 2); tibial apices uni-uncinate (male) or bi-uncinate (female) (Figs. 5 and 6) (Attelabinae) (1). Prothorax with distinct lateral carina (Fig. 3); propleura strongly excavated beneath carina; body small, robust; metallic bluish-green to bluish-black (Pterocolinae)... Pterocolus Prothorax not laterally carinate (Fig. 4); propleura not strongly excavated; body size, shape and color variable (Rhynchitinae) (2). Scutellary striole present... 4 Scutellary striole absent (3). Pygidium completely or almost completely covered by elytra; elytra with some erect setae; males with one tooth on outer edge of mandibles and females with two... Eugnamptus Pygidium mainly exposed, not completely or almost completely covered by elytra; elytra without erect setae; mandibular teeth similar in both sexes (4). Elytral striae distinct, more or less quadrate, moderately to deeply impressed; intervals narrower than width of striae, convex, smooth; interval punctures much smaller than striae; pubescence inconspicuous, more or less appressed; body dark colored, in some feebly metallic bronze or blue, less than 4 mm in length... Temnocerus Elytral striae more or less indistinct (more distinct in M. bicolor), weakly impressed, especially posteriorly; discernable intervals wider than width of striae, more or less flat, in some minutely rugose; interval punctures numerous, as large or nearly as large as striae; pubescence conspicuous, fine, semi-erect; body variable reddish-orange and black to brownish-black with faint bluish metallic luster, greater than 4 mm in length.... Merhynchites 6(3). Elytral striae distinctly rowed; intervals distinct, with punctures more or less smaller than striae; punctures not masked by pubescence... 7 Elytral striae not distinctly rowed; intervals indistinct, punctures as large or nearly as large as striae; punctures in some masked by pubescence (6). Elytra short, exposing pygidium and two abdominal terga; hind basitarsal segment longer than combined length of hind tarsal segments 3 and Deporaus 4 8 FIGURES Himatolabus pubescens (Say), male, lateral habitus; 3. Pterocolus ovatus (Fabricius), male, lateral habitus; 4. Rhynchites velatus LeConte, male, lateral habitus; 5. Himatolabus pubescens (Say), female, protibial apex; 6. Himatolabus pubescens (Say), male, protibial apex; 7. Pterocolus ovatus (Fabricius), tarsal claw; 8. Haplorhynchites aeneus (Boheman), tarsal claw; 9. Himatolabus pubescens (Say), tarsal claw. Elytra not short, covering all abdominal terga and part of the pygidium; hind basitarsal segment shorter or subequal to hind tarsal segments 3 and 4 combined (7). Elytral intervals wide, more or less flat, with numerous punctures, punctures in some as large or nearly as large as striae; males mostly more setose along elytral suture at declivity; antennal club symmetrical in both sexes Haplorhynchites Elytral intervals narrow, more or less convex, moderately punctured; punctures much smaller than striae; males not more setose along elytral suture at declivity; males with asymmetrical antennal club... Involvulus 9(6). Body with violaceous to greenish metallic luster; elytral pubescence of appressed whitish setae and widely rowed, dark, erect setae; males with anterolateral spine on each side of pronotum(fig. 4); length mostly greater than 4 mm Rhynchites Body without violaceous or greenish metallic luster; elytral pubescence not as above; males without anterolateral spines on each side of pronotum; length mostly less than 4 mm... Auletobius 10(1). Upper surface with appressed pubescence Himatolabus Upper surface without appressed pubescence, glabrous except for a few scattered fine erect setae

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

17 708 Family 128. Attelabidae Auletini Desbrochers 1908 Auletobius Desbrochers 1869, 9 spp.; known larvae develop in cut terminal leaf primordia; associated with a wide variety of hosts including species of Quercus, Comptonia, Potentilla, Eriogonum, etc.; regionally distributed in U.S. (key to Nearctic species, Pierce 1909; key to World species, Voss ). Deporaini Voss 1929 Deporaus Samouelle 1819, 1 sp., D. glastinus LeConte; larvae are leaf miners; associated with Q. wislizenii in southern CA and other Quercus spp. throughout its range; ranges throughout western U.S., east to CO and southeast to NM and west TX. Platyrhynchus Thunberg (1815) Pterocolinae Lacordaire 1866 Pterocolus Say 1831, 1 sp., P. ovatus (Fabricius) (Fig. 3); leaf roll thief, north of Mexico the larvae develop in leaf rolls prepared by three species of attelabine weevils (H. pubescens, A. bipustulatus and H. analis); distributed throughout eastern half of U.S., southwest to TX and AZ and south into Mexico (key to New World species, Hamilton 1998). BIBLIOGRAPHY ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL A world catalogue of families and genera of Curculionoidea (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae). Entomopraxis. Spain, 350 pp. ANDERSON, D. M Curculionidae (broad sense) (Curculionoidea). Pp In: F. W. Stehr, ed. Immature Insects. volume 2. Kendall/Hunt. Dubuque, IA. BALDUF, W. V Obligatory and facultative insects in rose hips, their recognition and bionomics. University of Illinois Press. Urbana, IL., 194 pp. BEDEL, L Faune des Coleopteres du Bassin de la Seine. Rhynchophora. Annales del la Société Entomologique de France, 6 (hors. Serie), 442 pp. BILLBERG, G. J Enumeratio Insectorum in Museo G. J. Billberg. Gadel, Stockholm, 138 pp. BLATCHLEY, W. S. and C. W. LENG Rhynchophora or weevils of North Eastern America. The Nature Publishing Company. Indianapolis, IN, 682 pp. BØVING, A. G. and F. C. CRAIGHEAD An illustrated synopsis of the principal larval forms of the order Coleoptera. Entomologica Americana, 11: BRIGHT, D. E The Insects and Arachnids of Canada. Part 21. The weevils of Canada and Alaska: Volume 1. Coleoptera: Curculionoidea, excluding Scolytidae and Curculionidae. Publication Research Branch, Agriculture Canada. Ottawa, Canada. CHITTENDEN, F. H Some insects injurious to violet, rose and other ornamental plants. United States Department of Agriculture Bulletin, 27: COOLEY, R. A Two insect pests: the rosebud curculio, Rhynchites bicolor Fabr.- the poplar leaf-folding sawfly, Pontania bozemani Cooley. Montana Experiment Station. Bulletin, 46: CROWSON, R. A The natural classification of the families of Coleoptera. E. W. Classey. Middlesex, England, 214 pp. DICKERSON, E. L Notes on Rhynchites bicolor Fabr. Journal of Economic Entomology, 3: EDWARDS, J. C Coleoptera or beetles east of the Great Plains. J. W. Edwards. Ann Arbor, MI, 181 pp. ESSIG, E. O Insects and mites of western North America. MacMillan. New York, NY, ix-xiii pp. EWING, H. E A case of persistent melanism. Biological Bulletin, 28: FARRELL, B.D Inordinate fondness explained: Why are there so many beetles? Science, 281: FROST, C.A Notes on Attelabus rhois and parasite. Psyche, 15: GATES, B. N The rose curculio (Rhynchites bicolor Fabr.) in Massachusetts. Journal of Economic Entomology, 2: GIRAULT, A. A Attelabus bipustulatus Fabr., the theory of oviposition and construction of nidus; misc. notes. Entomological News, 15: GISTEL, J Die Mysterien der Europaischen Insectenwelt. Dannheimer. Kempten, pp. HAMILTON, R.W Studies of the Rhynchophorous families Nemonychidae, Attelabidae and Rhynchitidae, with a revision of North American species of Attelabus Linnaeus, Rhynchites Schneider and Eugnamptus Schoenherr (Coleoptera: Curculionoidea). Unpublished Ph.D. dissertation, The Ohio State University, Columbus, Ohio, microfilm, 492 pp. HAMILTON, R.W The genus Pselaphorhynchites in America, north of Mexico (Coleoptera: Rhynchitidae). Annals of the Entomological Society of America, 64: HAMILTON, R.W Observations on the biology of Haplorhynchites aeneus (Boheman) (Coleoptera: Rhynchitidae). Coleopterists Bulletin, 27: HAMILTON, R.W The genus Haplorhynchites in America north of Mexico (Coleoptera: Rhynchitidae). Annals of the Entomological Society of America, 67: HAMILTON, R.W Taxonomic use of endophallic structures in some Attelabidae and Rhynchitidae of America north of Mexico with notes on nomenclature. Annals of the Entomological Society of America, 72: HAMILTON, R.W Notes on the biology of Eugnamptus collaris (Fabricius) (Coleoptera: Rhynchitidae) with descriptions of the larva and pupa. Coleopterists Bulletin, 34:

18 Family 128. Attelabidae 709 HAMILTON, R. W Description of the larva and pupa of Haplorhynchites aeneus (Boheman), (Coleoptera: Curculionoidea: Rhynchitidae). Journal of the Kansas Entomological Society, 54: HAMILTON, R. W Biological data on two North American rhynchitids (Coleoptera: Rhynchitidae) associated with sweet fern, Comptonia peregrina (Linnaeus): with descriptions, illustrations and comparisons of their immature stages. Journal of the Kansas Entomological Society, 56: HAMILTON, R. W The genus Merhynchites Sharp in America north of Mexico. Southwestern Entomologist, 10: HAMILTON, R. W A revision of the weevil genus Eugnamptus Schoenherr (Coleoptera: Rhynchitidae) in America north of Mexico. Transactions of the American Entomological Society, 115: HAMILTON, R. W Revision of the New World Genus Himatolabus Jekel (Coleoptera: Attelabidae) in North America. Transactions of the American Entomological Society, 118: HAMILTON, R. W New life cycle data for two western North American weevils (Coleoptera: Rhynchitidae) with a summary of North American rhynchitid biology. Coleopterists Bulletin, 48: HAMILTON, R. W Taxonomic revision of the New World Pterocolinae (Coleoptera: Rhynchitidae. Transactions of the American Entomological Society, 124: HAMILTON, R. W. and S. S. KURITSKY Description of the larva and pupa of Merhynchites bicolor (Fabricius). Coleopterists Bulletin, 35: HARRIS, T. W A treatise on some of the insects injurious to vegetation. Crosby and Nichols. Boston, MA, 640 pp. HOERNER, J. L Western Rose Curculio, Rhynchites bicolor wickhami Cockerell. Colorado Agricultural Experiment Station Bulletin, 432: HOPKINS, A. A (Secretarial Notes). Proceedings of the Entomological Society of Washington, 7: JEKEL, H Insecta saundersiana: or characters of undescribed insects in the collection of William Wilson Saunders, Esq. E. Newman. London, 242 pp. KISSINGER, D. G Curculionidae of America North of Mexico. A key to the genera. Taxonomic Publications. South Lancaster, MA, 143 pp. KUSCHEL, G A phylogenetic classification of Curculionoidea to families and subfamilies. Memoirs of the Entomological Society of Washington, 14: LAWRENCE, J. L Coleoptera. Pp In: S. P. Parker, ed. Synopsis and classification of living organisms. McGraw/ Hill. NY. LECONTE, J. L In J. L. LeConte and G. H. Horn, The Rhynchophora of America, north of Mexico. Proceedings of the American Philosophical Society, 15: LINNAEUS, C. V Systema naturae per regna tria naturae secundum classes, ordines, genera, species, cum characteribus, differentis, synonymis, locis, ed. 10. Salvius. Holmiae, vol. 1, 823 pp. LOVETT, A. L The rose curculio, Rhynchites bicolor Fabr. injures blackberry buds. Reports of the Department of Entomology, Oregon Agricultural Experiment Station Bulletin, pp LODING, H. P Catalogue of the beetles of Alabama. Wetumpka Printing. Wetumpka, AL, 172 pp. LUTZ, F. E Field book of insects, 3rd ed. G. P. Putnam and Sons. New York, NY, 510 pp. MCBRIDE, D. K. and C. Y. OSETO Sunflower insect pests. Cooperative Extension Service. North Dakota State University, Fargo, ND, Circular E-623, pp MURTFELDT, M. E Notes on Attelabus bipustulatus Fabr. Canadian Entomologist, 4: O BRIEN, C. W. and G. J. WIBMER Annotated checklist of the weevils (Curculionidae, sensu lato) of North America, Central America and the West Indies (Coleoptera: Curculionoidea). Memoirs of the American Entomological Institute, 34: ii-ix, OLIVIER, A. G Entomologie, ou histoire naturelle des insectes, avec leurs caractères génériques et spécifiques, leur description, leur synonymie et leur figure enluminée. Desray. Paris, Coleopteres, vol. 5, 612 pp. PACKARD, A. S Fifth report of the U.S. Entomological Commission on insects injurious to forest and shade trees. Government Printing Office. Washington, DC, 957 pp. PIERCE, W. D Studies of North American weevils. Proceedings of the United States National Museum, 37(1708): PIERCE, W. D Miscellaneous contributions to the knowledge of the weevils of the families Attelabidae and Brachyrhinidae. Proceedings of the United States National Museum, 45 (1988): ROBERTSON, H. A The rose curculio in Manitoba with notes on other insects affecting roses. Annual Reports of the Entomological Society of Ontario, 54: SCHOENHERR, C. J Tabula synoptica familiae curculionidum. Isis Oken, heft X, columns SCHULZ, T. J. and W. V. LIPP The status of the sunflower insect complex in the Red River Valley of North Dakota. Proceedings of the North Central Branch, Entomological Society of America, 24: SHARP, D. E Biologia Centrali Americana, Insecta, Coleoptera, Curculionidae: Attelabinae. 4: SILFVERBERG, H Attelabus Linnaeus, 1758 (Insecta, Coleoptera): Request for confirmation of designation of typespecies. Z. N. (S.) Bulletin of Zoological Nomenclature, 34: THOMPSON, R. T Observations on the morphology and classification of weevils (Coleoptera, Curculionoidea) with a key to major groups. Journal of Natural History, 26:

19 710 Family 128. Attelabidae TING, P.C The mouthparts of the coleopterous group Rhynchophora. Microentomology, 1: VAN EMDEN, F On the taxonomy of Rhynchophora larvae (Coleoptera). Transactions of the Royal Entomological Society of London, 87: VOGT, G Leaf rolling weevils (Coleoptera: Attelabidae) their host plants, and associated rhynchitid weevils in North America (Canada through the Republic of Panama): Summary of a long term field study. Pp In: D. Quainter and A. Aiello, eds. Insects of Panama and Mesoamerica: Selected Studies. Oxford University Press, vxii pp. VOSS, E Monographische Bearbeitung der Unterfamilie Rhynchitinae (Curc.) I, Teil, Nemonychini-Auletini. Archiv für Natrurgeschichte, 88: VOSS, E Die Unterfamilien Attelabinae und Apoderinae (Col. Curc.) (18. Beitrag zur Kenntnis der Curculioniden). Stettiner Entomologische Zeitung, 85: 1-78, VOSS, E Monographie der Rhynchitinen-Tribus Rhynchitini. 2. Gattungsgruppe: Rhynchitina. V. 1. Teil der Monographie der Rhynchitinae-Pterocolinae. (41. Beitrag zur Kenntnis der Curculioniden.) Koleopterologische Rundschau, 18: (1932); 19: (1933). VOSS, E Monographie der Rhynchitinen-Tribus Auletini. III. Teil der Monographie der Rhynchitinae-Pterocolinae. (37. Beitrag zur Kenntnis der Curculioniden.) Stettiner Entomologische Zeitung. 94: , (1933); 95: , (1934); 96: , (1935); 97: (1936); 98: (1937). VOSS, E Monographie der Rhynchitinen-Tribus Deporaini sowie der Unterfamilien Pterocolinae- Oxycoryninae (Allocorynini). VII. Teil der Monographie der Rhynchitinae- Pterocolinae. (73. Beitrag zur Kenntnis der Curculioniden) Stettiner Entomologische Zeitung, 99: , (1938); 103: (1942); 104: (1943). VOSS, E Monographie der Rhynchitinen-Tribus Rhynchitini. 2. Gattungsgruppe: Rhynchitina. V. 2. Teil der Monographie der Rhynchitinae-Pterocolinae. (45. Beitrag zur Kenntnis der Curculioniden), Koleopterologische Rundschau, 24: VOSS, E Monographie der Rhynchitinen-Tribus Rhinocartini sowie der Gattungsgruppe Eugnamptina der Tribus Rhynchitini. IV. Teil der Monographie der Rhynchitinae- Pterocolinae. (32. Beitrag zur Kenntnis der Curculioniden). Deutche Entomologische Zeitschrift, 1941: VOSS, E Monographie der Rhynchitinen-Tribus Rhynchitini. 2. Gattungsgruppe: Rhynchitina. V. 2. Teil der Monographie der Rhynchitinae-Pterocolinae. (45. Beitrag zur Kenntnis der Curculioniden). Mittheilungen Muenchener Entomologische Gesellschaft, 31: VOSS, E Die Unterfamilie Camarotinae, ihre Beziehungen zur Familie Attelabidae sowie ein Versuch, die phylogenetischen Zusammenhange innerhalb der letzteren zur Darstellung zu bringen (Coleoptera, Curculionidae) (191. Beitrag zur Kenntnis der Curculioniden). Entomologische Abhandlungen Staatliches Museum fur Tierkunde in Dresden, 32 : VOSS, E Monographie der Rhynchitinen-Tribus Rhynchitini. 2. Gattungsgruppe: Rhynchitina. V. 2. Teil der Monographie der Rhynchitinae-Pterocolinae. Entomologische Arbeiten, aus dem Museum G. Frey, 20: WIBMER, G. J. and C. W. O BRIEN Annotated checklist of the weevils (Curculionidae, sensu lato ) of South America (Coleoptera: Curculionoidea). Memoirs of the American Entomological Institute, 39: i-xvi,

20 Family 129. Brentidae BRENTIDAE Billberg 1820 Family common name: The straight-snouted weevils; pear shaped weevils by Robert S. Anderson and David G. Kissinger Family synonyms: Apionidae Schoenherr 1823; Cycladidae Schoenherr 1823; Eurhynchidae Lacordaire As 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 antennae. Apioninae and Nanophyinae are small pear-shaped weevils immediately recognized by a long cylindrical 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 Arrenodes minutus (Drury) (from Bright 1993, reproduced with the permission of the Minister of Public Works and Government Services, 2001) Description (based on Lawrence 1982). Shape variable elongate, narrow and parallelsided (Brentinae, Cyphagoginae, Trachelizinae), or stouter and more robust with expanded oval elytra and a distinctive pear-like shape (Apioninae, Nanophyinae); flat to convex; length mm; color pale brown to black, very rarely red or bicolored or with contrasting colors or markings; vestiture lacking, or of fine short to moderately long appressed or suberect pubescence; rarely with scales. Rostrum moderately to very long and usually narrow and cylindrical; sexual dimorphism in rostrum form (Brentinae, Cyphagoginae, Trachelizinae) or length (Apioninae) generally evident. Antennae straight or rarely geniculate (Nano-phyinae), funicle very narrow and slender, rarely filiform or moniliform (Brentinae, Cyphagoginae, Trachelizinae), ending in a weak, loose club (Nanophyinae) or a compact club (Apioninae, Cyladinae) of three articles. Antennal insertions lateral at the middle of the rostrum or rarely dorsal near the base. Maxillary palps of two or three articles. Labial palps small, of one or two articles, attached dorsally near the apex of the prementum, rarely in cavities. Proventriculus lacks sclerotized plates. Front coxae vary from contiguous to widely separated. Middle coxae and hind coxae narrowly to widely separated. Front tibiae rarely equipped with an antennacleaning organ. Hind wing usually with reduced anal venation. Tarsal claws variable. First two visible sternites of the abdomen connate and much longer than following two; pygydium concealed by the elytra. Tegmen bilobed or notched apically and the median lobe with a trough-like ventral plate and a narrow dorsal plate. Larvae (based on Lawrence 1982) when mature about mm in length, slender and subcylindrical (Brentinae) or broad and strongly curved (Apioninae). Small legs rarely present on thorax (Brentinae), usually absent, but with well-developed pedal lobes (Apioninae). Frontal sutures complete although rarely indistinct, reaching articulating membrane of mandible. Antenna of a single membranous article. Hypopharyngeal bracon present. Abdomen with first eight segments with two, three or four dorsal folds and bearing annular or rarely biforous spiracles. Habits and habitats. As noted, Brentidae is a composite of taxa of subfamily rank each of which has a rather different body form. Natural history aspects of Brentidae similarly are correlated with, and best discussed, at the subfamily level (Thomas 1996). Brentinae, Cyphagoginae and Trachelizinae are the traditional brentids. They are small to large, elongate weevils that are usually found under bark as both adults and larvae. Where known, females lay their eggs in holes they have excavated with their rostrum in living, dying or recently felled hardwood trees. Larvae tunnel deep into the heartwood and appear to feed on wood and fungus mycelia. Brentus anchorago Linnaeus is one of North America s largest weevils, with males reaching a body length of up to 40 mm. In some species, adult males guard females during egg-laying. Cyladinae includes only the adventive Cylas formicarius (Fabricius), the sweet potato weevil. Adults and larvae are associated with various Convolvulaceae (especially Ipomoea pescapreae L.) and are pests of sweet potatoes, Ipomoea batatas L. Larvae mine the stems. Little is known about Nearctic Nanophyinae. Most Palearctic Nanophyinae are associated with plants of the family Lythraceae, especially the genus Lythrum, but also Crassulaceae, Tamaricaceae and Ericaceae. Larvae are associated with the fruits, leaves and stems and many appear to cause galls. One species of Nanophyes Schoenherr has been introduced into North America for biological control of Lythrum salicaria L. (purple loosestrife; Lythraceae)

21 712 Family 129. Brentidae Trichapion centrale (Fall) (from Bright 1993, reproduced with the permission of the Minister of Public Works and Government Services, 2001) (Harris 2001) and a second is under consideration. Zeugonyx sabinae Notman has been reared from galls on Juniperus ashei Buchh. in Texas, but otherwise no information is available on the native North American fauna. Apioninae are associated with a wide variety of plants, mostly Fabaceae and Asteraceae, but also Malvaceae, Polygonaceae, Caprifioliaceae, and Umbelliferae. Many of the newly elevated genera, formerly subgenera of Apion sensu lato, are host specific on certain families of plants (e.g., Alocentron on Malvaceae; Perapion on Polygonaceae). Larvae mine stems, feed on developing pericarp of fruit, or develop in the seeds. A few species are adventive. Rhopalapion longirostre (Olivier) is associated with the ornamental hollyhock, Althea rosea Cav. (Tuttle 1954) and Ischnopterapion virens (Herbst) with clover, Trifolium (Hoebeke et al. 2000). Two species of Exapion have been introduced into the western United States as biological control agents for gorse, Ulex europaeus L. and scotch broom, Cytisus scoparius (L.) (both Fabaceae) (O Brien 1995) and Omphalapion hookerorum (Kirby) has 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 classification 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 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 wellknown in North America and their classification remains unchanged (Warner 1960; Kissinger 1968). Brentinae have 3 species in 3 genera, and Cyphagoginae, Trachelizinae and Cyladinae each 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, 3 4 FIGURES Front leg, Brentinae (schematic); 4. Front leg, Apioninae (schematic). there are now 5 genera and 5 species recognized in North America. This is based on phylogenetic evidence and follows the division of the once larger widespread genus Nanophyes Schoenherr by Alonso-Zarazaga (1989) and followed by Alonso-Zarazaga and Lyal (1999). The situation with Apioninae is even more complex with many of the various subgenera (e.g., Kissinger 1968) now elevated to generic rank, many generic concepts have been narrowed with some North American species transferred into previously Palearctic genera, and a number of new genera have been proposed (Kissinger 1989, 1990, 1992, 1998, 1999; Alonso- Zarazaga 1990). Alonso-Zarazaga (1990) treats the Palearctic classification of Apioninae in detail and is the basis for the development and understanding of the new system as applied in North America. At present in North America there are approximately 5 FIGURES Dorsal habitus, Cylas formicarius (Fabricius 1798); 6. Lateral view, pronotum and base of elytra, Cylas formicarius. 6

22 Family 129. Brentidae species placed in the 19 genera. A small number of North 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 (starting 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 1. Trochanter short and triangular, femur attached to the side of trochanter (Fig. 3); body more elongate 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) (1). Pronotum with a broad constricted collar (that is almost as long as the rest of the pronotum) before base (Fig. 5); mesothorax with a deep vertical sulcus near the posterior margin of the pronotum (Fig. 6); tarsal claws connate at base; elytra elongate-oval in dorsal view, lacking impressed striae (Fig. 5) (Cyladinae)... Cylas Pronotum with at most a very short constriction before base (Figs. 1, 7, 9, 10, 14); mesothorax lacking a deep lateral pit near the posterior margin of the pronotum; tarsal claws free; elytra elongate and parallel-sided in dorsal view, with distinctly or indistinctly impressed striae (2). All femora lacking distinct tooth (although slight angulation may be present on front femur); rostrum with ventral surface punctate and with dense microsetae or sparse elongate recumbent setae... 4 At least front femur with obvious tooth; rostrum with ventral surface smooth or coarsely punctate, not setose (Brentinae) (3). Elytra with stria 3 bent inwards at middle towards elytral suture and coalescent with stria 2 (Fig. 7); front tibia with inner edge incised in apical one- FIGURES Dorsal habitus, Stereodermus exilis Suffrian; 8. Front leg, Stereodermus exilis; 9. Dorsal habitus, Paratrachelizus uncimanus (Boheman); 10. Dorsal habitus, Brentus anchorago (Linnaeus), male; 11. Dorsal view, head and pronotum, Brentus anchorago, female; 12. Dorsal view, head, Arrenodes minutus (Drury), male; 13. Front leg, Arrenodes minutus.

23 714 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; funicle apparently with only 5 articles, articles 6 and 7 as wide as apical antennal article and forming a loose club (Nanophyinae)... 8 Antenna straight, not geniculate, scape shorter than combined length of first 3 articles of funicle; funicle with 7 articles, articles 6 and 7 narrow, not as wide as apical antennal article and distinct from compact apical club (Apioninae) (7). Tarsal claws equal in length and form... 9 Inner tarsal claw shorter and finer than outer claw FIGURE Dorsal habitus, Heterobrenthus texanus Schaeffer, male. half, incision surrounded by very dense setae (Fig. 8); rostrum with ventral surface with sparse elongate recumbent setae (Cyphagoginae) Stereodermus Elytra with stria 3 evident only as a row of shallow punctures, straight throughout length, not coalescent with stria 2 (Fig. 9); front tibia with inner edge straight, not incised; rostrum with ventral surface with dense microsetae in basal portion (Trachelizinae)... Paratrachelizus 5(4). Head longer than broad, not distinctly constricted behind eyes, lateral margins continuous, prolonged and convergent behind eyes (Figs. 10, 11); pronotum medially deeply sulcate in basal one-half (Figs. 10, 11); male with pronotum extremely elongated, lateral margins constricted near middle (Fig. 10)... Brentus Head short, width more or less equal to length, distinctly constricted behind eyes, lateral margins at most slightly prolonged but interrupted by constriction, not convergent behind eyes (Figs. 1, 12, 14); pronotum uniformly smooth dorsally, not sulcate (Figs. 1, 14); male with pronotum not elongated, and lateral margins not constricted near middle (5). Antenna with articles 2 to 11 more or less equal in length and width towards apex (Fig. 1); head constricted behind eyes, hind angles not projecting (Figs. 1, 12); front tibia with at most a swelling on inner margin, no distinct tooth evident (Fig. 13); rostrum very dissimilar in male (short and broad with large mandibles, Fig. 12) and female (narrow and cylindrical, Fig. 1)... Arrenodes Antenna with articles 2 to 11 increasing in length and width towards apex (Fig. 14); head strongly constricted behind eyes, hind angles projecting; front tibia with distinct tooth on inner margin (Fig. 14); rostrum dissimilar in male (dilated at apex) 9(8). Elytra with interstria 8 with a row of small, dark tubercles (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 mm Pseudotychius Elytra with interstria 8 smooth in basal one-quarter; elytral vestiture various; length mm (9). Elytra lacking fascia, setae appressed and uniformly arranged, each seta short, not or hardly exceeding the base of the following one (Fig. 16); pronotum and tibia lacking specialized erect setae, only one subhumeral seta on elytral interstria 9 (Fig. 16)... Microon Elytra with fascia, setae arcuate and disordered, each seta long, usually exceeding the base of the following one (Fig. 17); pronotum, tibia and odd elytral interstriae with specialized erect setae (Fig. 17)... Nanophyes 11(8). Antenna with basal 5 articles of funicle slender; elytra largely pale, with basal dark area; vestiture fine and short, consisting of whitish and dark scales... Nanodactylus Antenna with basal 4 articles of funicle slender; elytra uniformly testaceous; vestiture of long whitish setae... Zeugonyx 12(7). Middle coxae contiguous, not separated by junction between mesosternal process and intercoxal process of metasternum Middle coxae separated by junction between mesosternal process and intercoxal process of metasternum (12). Tarsal claws with acute basal tooth... Chrysapion Tarsal claws simple, at most swollen as base, not toothed (13). Rostrum straight, in side view not forming angle at junction with frons, apical half of rostrum usually tapered to apex (Fig. 18); eyes not or scarcely prominent; antennae inserted at basal of rostrum; scape at most as long as width of rostrum at midlength; prescutellar fovea conspicuous, lanceolate to sulciform; tarsi normal, front tarsus with first article X as long as wide; internal sac of aedeagus without baculi, other

24 Family 129. Brentidae FIGURES 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 redrawn after Alonso-Zarazaga 1990). structures may be present; associated with Polygonaceae... Perapion Rostrum strongly curved, in side view forming a conspicuous angle at junction with frons, apical half of rostrum parallel-sided to slightly widened to apex (Fig. 19); eyes prominent; antennae inserted at basal of rostrum; scape longer than width of rostrum at midlength; prescutellar fovea inconspicuous, similar to pronotal punctures; tarsi robust, front tarsus with first article X as long as wide; internal sac of aedeagus with two baculi and denticles; associated with Pinus (Pinaceae)... Podapion 15(12).* Front femur of male with polished (sometimes striate) area on ventral surface, the area generally limited posteriorly by a prominent longitudinal carina (Fig. 20); metasternum generally with a pair of spicules near the middle of the posterior margin... Fallapion Front femur of male without polished area on ventral surface (15). Pygydium of male lacking distinct, deep, transverse preapical sulcus, profile of pygydium not interrupted (Figs. 21, 22); abdominal ventrite 5 of male rounded apically; elytra with specialized seta on interstria 7 lacking Pygydium of male with distinct, deep, transverse preapical sulcus, profile of pygydium distinctly interrupted by sulcus (Figs. 23, 24, 27, 28); abdominal ventrite 5 of male truncate apically; elytra with specialized seta on interstria 7 at middle or near apical 1/3 present or lacking * From this point on the key is complex and relies extensively on characters of males. Males can be recognized by the shorter, more coarsely sculptured rostrum and (sometimes) by an upward deflection of the last abdominal ventrite. These are the taxa traditionally treated as Apion sensu lato and the reader may wish to consult Bright (1993) and Kissinger (1968) for species level identifications.

25 716 Family 129. Brentidae 17(16). Antennal club with sutures obsolete or absent; tarsal claws simple, not toothed; on Matricaria perforata (Asteraceae)... Omphalapion Antennal club with sutures distinct; tarsal claws toothed at base; on Malvaceae (17). Pronotum with vestiture directed away from the midline at the base, more or less parallel to the midline along the lateral margins, and perpendicular to the apical margin along the apical margin (Fig. 25); pronotum with basal flange well-developed (Fig. 25); on Malvaceae... Alocentron Pronotum with vestiture directed towards the midline, in some cases, totally transverse at the basal or apical margins (Fig. 26); pronotum with basal flange obsolete (Fig. 26); on Althea rosea (Malvaceae)... Rhopalapion 19(16). Elytra with transverse pattern of contrasting light and dark scales; pygydium of male with sulcus incomplete laterally, not reaching side margins of pygydium (Fig. 24); front tarsus with article 2 stout, about as wide as long; femora and tibiae robust; middle coxae widely separated ( x diameter of middle coxa); aedeagus with internal sac with large apically bifurcate structure; on Rutaceae, Caprifoliaceae... Neapion Elytra with scales more uniformly colored, not contrasting light and dark; pygydium of male with sulcus complete laterally, reaching side margins of pygydium (Figs. 27, 28); front tarsus with article 2 usually longer than wide; femora and tibiae slender; middle coxae moderately widely separated (<0.25 x diameter of middle coxa); aegeagus with internal sac without large structure; on Fabaceae and other plants (19). Genitalia of male with tegmen with prostegium articulated with free ring (Fig. 29) Genitalia of male with tegmen with prostegium fused with free ring (Fig. 30) (20). Tibiae of male simple, not mucronate Middle tibia (at least) of male mucronate (21). Scrobe with dorsal margin produced into a long, slender acute process projected ventrally; elytra with interstria 7 with specialized seta inserted near midlength of stria; metasternum very convex in lateral view; meso- and metasternal processes directed inwardly where meeting between hind coxae; on Fabaceae... Exapion Scrobe with dorsal margin simple, not produced into a long, slender acute process; elytra with interstria 7 or 9, or 7 and 9, with or without specialized seta inserted at or behind apical 1/3 of elytron; metasternum slightly convex in lateral view; mesoand metasternal processes either flat or with metasternal process more prominent where meeting between hind coxae; on various plants (22). Body color red throughout; lateral margins of pronotum parallel in basal one-half (Fig. 31); male genitalia with prostegium with high median crista projected basally; elytral interstria 2 prolonged outward at apex (Fig. 32); on Polygonaceae Apion Body color black or with metallic sheen, occasionally with light colored appendages; lateral margins of pronotum various; male genitalia with prostegium with at most moderately high basal median carina not projected basally; elytral interstria 2 only slightly prolonged outward at apex (Fig. 33); on Fabaceae (23). Elytra blue-green; elytral interstria 7 with one specialized seta in apical one-third; legs dark; lateral margins of pronotum parallel in basal one-half; rostrum not prominently expanded at point of insertion of antennae; eyes prominent Ischnopterapion Elytra black; elytral interstriae 7 and 9 with one specialized seta near apex; legs various, may be light in color; lateral margins of pronotum various in basal one-half; rostrum variously expanded at point of insertion of antennae; eyes prominent or not (24). Pronotum without basal flange, with lateral margins nearly evenly convergent from base to apex, subconical in dorsal form; rostrum prominently expanded at point of insertion of antennae; legs and article 1 of antenna dark... Apionion Pronotum with or without basal flange, with lateral margins various; rostrum smooth at point of insertion of antennae, not expanded; legs and article 1 of antenna dark or light (25). Pronotum with basal flange indistinct, subconical in dorsal form (Fig. 34); legs and article 1 of antenna light; aedeagus with median lobe and tegmen narrow, elongate, subcylindrical; internal sac with large teeth (>0.030 mm long)... Sayapion Pronotum without basal flange, lateral margins subparallel in basal one-half; legs and article 1 of antenna light or dark; aedeagus with median lobe slightly depressed, broader at base; internal sac with largest teeth (<0.020 mm long)... Kissingeria 27(21). Head with ventral surface with subcephalic ridges lacking or low, not extended basally on head to point equivalent to middle of eye; vestiture of interstriae 2-5 sparse, generally inconspicuous, consisting of one row of scales; one specialized seta on interstria 9; aedeagus with parameroid lobe with one or more macrochaetae; on Betula papyrifera (Betulaceae)... Betulapion Head with ventral surface with subcephalic ridges moderately to well developed, extended to or beyond the middle of the eye; vestiture of interstriae 2-5 relatively dense, consisting of two or more rows of scales; one specialized seta on each of interstriae 7 and 9; aedeagus with parameroid lobe generally lacking macrochaetae; associated mainly with Fabaceae and Asteraceae... Trichapion 28(20). Elytra blue in color, legs dark; tarsus with claw simple; rostrum of male markedly expanded laterally at point of antennal insertion (Fig. 35); frons flat, striate... Mesotrichapion Elytra not blue in color; rostrum of male smooth, not expanded laterally at point of antennal insertion (28). Male with metasternum tuberculate on median posterior margin and apical area of parameroid lobe

26 Family 129. Brentidae 717 bearing four or more macrochaetae more than 0.055mm long... Eutrichapion Male metasternum not tuberculate or parameroid lobe lacking macrochaetae more than 0.010mm long... Coelocephalapion CLASSIFICATION OF THE NEARCTIC GENERA Brentidae Billberg 1820 Brentinae Billberg 1820 Brentini Billberg 1820 Brentus Fabricius 1787, 1 sp., B. anchorago (Linnaeus 1758), Florida. Adults are often found under bark of Bursera simaruba (L.) Sarg. Brenthus Illiger 1801 Arrhenodini Lacordaire 1866 Arrenodes Schoenherr 1823, 1 sp., A. minutus (Drury 1770), generally distributed in eastern United States, into extreme southern Canada. Adults are found under bark especially of oaks; larvae bore into the wood (Buchanan 1960). Males are territorial and guard females during egg-laying (Sanborne 1983). Brentus Panzer 1788, not Fabricius 1787 Arrhenodes Schoenherr 1826 Eupsalis Lacordaire 1866 Platysystrophus Kleine 1917 Heterobrenthus Sharp 1895, 1 sp., H. texanus Schaeffer 1915, southern Texas. Cyphagoginae Kolbe 1892 Stereodermini Sharp 1895 Stereodermus Lacordaire 1866, 1 sp., S. exilis Suffrian 1870, southern Florida. Trachelizinae Lacordaire 1866 Trachelizini Lacordaire 1866 Paratrachelizus Kleine 1921, 1 sp., P. uncimanus (Boheman 1839), southern Florida. Cyladinae Schoenherr 1823 Cylas Latreille 1802, 1 sp., C. formicarius (Fabricius 1798), generally distributed in southern United States. Adventive on Convolvulaceae including Ipomoea batata (L.) Lam. (sweet potato). Cylanus Rafinesque 1815 Protocylas Pierce 1941 Apioninae Schoenherr 1823 Apionini Schoenherr 1823 Apion Herbst 1797, 1 sp., identity uncertain, Alberta. Adults have been collected around edges of high elevation snowfields. Associated with Polygonaceae in Palearctic Region. Apiolum Kirby 1808 Apionus Rafinesque 1814 Apius Billberg 1820, not Panzer 1806 Apium Agassiz 1846 Oxystomum Gistel 1856 Oxeostomum Gistel 1856 Erythrapion Schilsky 1906 Aplemonini Kissinger 1968 Perapion Wagner 1907, 4 spp., generally distributed. Associated with Polygonaceae. Eroosapion Ehret 1994 (valid subgenus) Hemiperapion Wagner 1930 (valid subgenus) Rhaphidoplectron Alonso-Zarazaga 1990 (valid subgenus) Podapion Riley 1883, 1 sp., P. gallicola Riley 1883, generally distributed in eastern United States into extreme southern Canada; also in California and Oregon. Associated with Pinus spp. Aspidapiini Alonso-Zarazaga 1990 Alocentron Schilsky 1901, 6 spp., generally distributed. Associated with Malvaceae. Bulborhinapion Schatzmayr 1926 (valid subgenus) Nearctalox Alonso-Zarazaga 1990 (valid subgenus) Ceratapiini Alonso-Zarazaga 1990 Omphalapion Schilsky 1901, 1 sp., O. hookerorum (Kirby 1808), British Columbia, Alberta, Saskatchewan, Manitoba and Nova Scotia; adventive. This species was first discovered in Nova Scotia (Peschken et al. 1993) but has recently been introduced into western Canada for biological control of Matricaria perforata Mérat (scentless chamomile; Asteraceae) (Harris and McClay 2001). Exapiini Alonso-Zarazaga 1990 Exapion Bedel 1997, 2 spp., California, Oregon and Washington. Adventive; introduced for biological control of gorse, Ulex europaeus L. and scotch broom, Cytisus scoparius (L.) (Fabaceae) (O Brien 1995). Ulapion Ehret 1997 (valid subgenus) Ixapiini Alonso-Zarazaga 1990 Neapion Alonso-Zarazaga 1990, 6 spp., generally distributed in eastern North America west to southern Texas, north into ex-

27 718 Family 129. Brentidae treme southern Canada. Subgenus Neotropion, 1 sp., N. xanthoxyli (Fall), associated with Rutaceae (genus Zanthoxylum); subgenus Neapion, 5 spp., associated with Caprifoliaceae (genus Viburnum). Xixias Kissinger 1990 Neotropion Alonso-Zarazaga 1990 (valid subgenus) Malvapiini Alonso-Zarazaga 1990 Rhopalapion Schilsky 1906, 1 sp., R. longirostre (Olivier 1807), generally distributed. Adventive on Althea rosea Cav. (hollyhock; Malvaceae). Oxystomatini Alonso-Zarazaga 1990 Oxystomatina Alonso-Zarazaga 1990 Eutrichapion Reitter 1916, 3 spp., generally distributed. Associated with Fabaceae. Cnemapion Bokor Leconteapion Alonso-Zarazaga 1990 (valid subgenus) Phalcrolobus Alonso-Zarazaga 1990 (valid subgenus) Psilocalymma Alonso-Zarazaga 1990 (valid subgenus) Mesotrichapion Györffy 1956, 1 sp., M. cyanitinctum (Fall 1927), Alaska to northern Quebec, south to southern Manitoba. Association with Fabaceae (genus Astragalus). Trichapiina Alonso-Zarazaga 1990 Betulapion Ehret 1994, 1 sp., B. simile (Kirby 1811), generally distributed. Adventive on Betula papyrifera Marsh. (paper birch; Salicaceae); larvae develop in flowers. Kissingeria Alonso-Zarazaga 1990, 6 spp., generally distributed. Associated with Fabaceae. Trichapion Wagner 1912, 45 spp., generally distributed. Associated mostly with Fabaceae and Asteraceae. Synapiina Alonso-Zarazaga 1990 Ischnopterapion Bokor 1923, 1 sp., I. virens (Herbst 1797), New York, Pennsylvania, Maryland, New Jersey, Connecticut, Delaware and Virginia. Adventive on various species of clover, Trifolium (Fabaceae) (Hoebeke et al. 2000). Piezotrachelini Voss 1959 Chrysapion Kissinger 1968, 2 spp., Arizona, California and Texas. Fallapion Kissinger 1968, 30 spp., generally distributed. Associated mostly with Asteraceae and Umbelliferae. Incertae sedis Apionion Kissinger 1998, 2 spp., A. crassum (Fall 1898), eastern United States; A. dilatatum (Smith 1884), Arizona. Associated with Fabaceae. Coelocephalapion Wagner 1914, 22 spp., generally distributed. Associated mostly with Fabaceae and Asteraceae. Sayapion Kissinger 1999, 5 spp., eastern United States, Texas and Arizona. One species has been associated with Fabaceae. Nanophyinae Gistel 1856 Nanophyini Gistel 1856 Microon Alonso-Zarazaga 1989, 1 sp., M. canadense (Brown 1944), western United States south to Arizona and western Canada east to Manitoba. At least one Palearctic species is associated with the genus Lythrum (Lythraceae). Nanodactylus Blatchely 1922, 1 sp., N. obesulus Blatchley 1922, Illinois, Indiana, Texas. Nanophyes Schoenherr 1838, 1 sp., N. marmoratus (Goeze 1777), Manitoba, New York; introduced into Canada in 1997 for the biological control of Lythrum salicaria L. (Lythraceae) (Harris 2001). This weevil species is also approved for introduction into the United States and is established in the vicinity of Ithaca, New York. A second species, N. brevis Boheman 1845, also is under consideration for introduction. Pseudotychius Blatchley 1922, 1 sp., P. watsoni Blatchley 1922, eastern United States, north into Ontario. Zeugonyx Notman 1922, 1 sp., Z. sabinae Notman 1922, Texas. This species was reared from galls on twigs of Juniperus ashei Buchh. (Cupressaceae). BIBLIOGRAPHY ALONSO-ZARAZAGA, M. A Revision of the supraspecific taxa in the Palaearctic Apionidae Schoenherr, Introduction and subfamily Nanophyinae Seidlitz, 1891 (Coleoptera, Curculionoidea). Fragmenta Entomologica, 21: ALONSO-ZARAZAGA, M. A Revision of the supraspecific taxa in the Palaearctic Apionidae Schoenherr, 1823 (Coleoptera, Curculionoidea). 2. Subfamily Apioninae Schoenherr, 1823: Introduction, keys and descriptions. Graellsia, 46: ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL A world catalogue of families and genera of Curculionoidea (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae). Entomopraxis. Barcelona, Spain.

28 Family 129. Brentidae 719 BRIGHT, D. E The Insects and Arachnids of Canada. Part 21. The weevils of Canada and Alaska: Volume 1. Coleoptera: Curculionoidea, excluding Scolytidae and Curculionidae. Publication Research Branch, Agriculture Canada. Ottawa, Canada. BUCHANAN, W. D Biology of the oak timberworm, Arrhenodes minutus. Journal of Economic Entomology, 53: HARRIS, P Nanophyes marmoratus (Goeze). Flower-feeding weevil. ananmar.htm HARRIS, P. and A. MCCLAY Omphalapion hookeri Kirby. Seed-head weevil. agents/aomphook.htm HOEBEKE, E. R., R. A. BYERS, M. A. ALONSO-ZARAZAGA, and J. F. STIMMEL Ischnopterapion (Chlorapion) virens (Herbst) (Coleoptera: Curculionoidea: Brentidae: Apioninae), a Palearctic clover pest new to North America: recognition features, distribution, and bionomics. Proceedings of the Entomological Society of Washington, 102: KISSINGER, D. G Curculionidae subfamily Apioninae of North and Central America with reviews of the world genera of Apioninae and world subgenera of Apion Herbst (Coleoptera). Taxonomic Publications. South Lancaster, Massachusetts. KISSINGER, D. G Apionidae from North and Central America. Part 1. Notes on the classification of the Apion subgenus Trichapion Wagner with description of new species from the United States of America (Coleoptera). Insecta Mundi, 3: KISSINGER, D. G Apionidae from North and Central America. Part 2. Description of a new subgenus and two new species of Apion from Mexico. (Coleoptera). Insecta Mundi, 4: KISSINGER, D. G Apionidae from North and Central America. Part 4. Generic classification and introduction to the genus Coelocephalapion Wagner, with new species from Mexico and Venezuela (Coleoptera). Insecta Mundi, 6: KISSINGER, D. G Apionidae from North and Central America. Part 5. Description of the genus Apionion and 4 new species (Coleoptera). Insecta Mundi, 12: KISSINGER, D. G Description of a new genus, Sayapion, from North and Central America (Coleoptera: Apionidae). Insecta Mundi, 13: 72. LAWRENCE, J.L Coleoptera, Pp In: S. P. Parker, ed. Synopsis and Classification of Living Organisms. Volume 2. McGraw Hill. New York. LAWRENCE. J. F. and A. F. NEWTON, Jr Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). Pp In: J. Pakaluk and S. A. Slipinski, eds., Biology, phylogeny and classification of Coleoptera: Papers celebrating the 80th birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw. O BRIEN, C. W Curculionidae, premiere biological control agents. Memoirs of the Entomological Society of Washington, 14: O BRIEN, C. W. and G. J. WIBMER Annotated checklist of the weevils (Curculionidae sensu lato) of North America, Central America, and the West Indies (Coleoptera: Curculionoidea). Memoirs of the American Entomological Institute, 34: i-ix, PESCHKEN, D. P., K. C. SAWCHYN and D. E. BRIGHT First record of Apion hookeri Kirby (Coleoptera: Curculionidae) in North America. Canadian Entomologist, 125: SANBORNE, M Some observations on the behaviour of Arrhenodes minutus (Drury) (Coleoptera: Brentidae). Coleopterists Bulletin, 37: THOMAS, M. C The primitive weevils of Florida (Coleoptera: Brentidae: Brentinae). Florida Department of Agriculture. Division of Plant Industry. Entomology Circular No pp. TUTTLE, D. M Notes on the bionomics of six species of Apion (Curculionidae, Coleoptera). Annals of the Entomological Society of America, 47: WARNER, R. E The genus Stereodermus new to America north of Mexico with a revised key to the genera of Brentidae. Coleopterists Bulletin, 14: 29.

29 720 Family 130. Ithyceridae 130. ITHYCERIDAE Schönherr 1823 by Robert S. Anderson Family common name: The New York weevil Among 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 pubescence. FIGURE Ithycerus noveboracensis (Forster) (from Bright 1993, reproduced with the permission of the Minister of Public Works and Government Services, 2001) Description (based on Lawrence 1982): Body large, stout. Pubescence of scale-like bristles, distinct. Head with single gular suture; pregular sutures absent. Rostrum broad, stout, not sexually dimorphic; antennal insertions lateral. Maxilla without lacinia; maxillary palp of three articles, rigid, partially retracted into large palpifer; labial palp of three articles; labrum absent; mandible stout. Antennae straight, moniliform; antennal club compact, of three articles. Proventriculus without sclerotized plates. Notosternal suture complete. Procoxae contiguous, mexocoxae narrowly separated and metacoxae moderately widely separated. Hind wing with four anal veins. Legs with trochanter short; triangular; femur attached to side of trochanter. Tarsal claws toothed. Abdomen with ventrites 1 and 2 fused together but separated by distinct suture. 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 mm. Subspherical, longer than wide. External surface of chorion of hexagonal facets with impressed borders. Surface strongly punctate, with numerous aeropyles. Larva (based in part on Sanborne 1981, Lawrence 1991) when mature about mm in length. Body relatively short and broad, subcylindrical, strongly c-shaped. Body yellowish, with heavily sclerotized mouth frame and mandibles. Vestiture of moderately long, scattered setae. Head protracted and hypognathous, about as long as wide. Frontal sutures complete, reaching articulating membrane of mandible. Clypeus distinguishable from frons and completely separated from labrum. Three pairs of stemmata on each side. Labrum free, bearing one sensillum and four pairs of setae. Antenna of a single dome-like article bearing a sensorium, a bifurcate appendage and several setae. Mandible with two apical teeth, and three blunt teeth on inner edge, accessory ventral process and mola absent. Hypopharyngeal bracon present. Maxilla with palp with 2 articles, palpiger present. Labial palp of two articles; palps widely separated. Premental sclerite subtriangular. Thorax with pronotal sclerite moderately pigmented. Legs very small, widely separated, subconical, two or three jointed. Abdomen with first four segments with three dorsal folds, segments 5-8 with two dorsal folds. Spiracles annular. Pupa (based on Sanborne 1981) with distinct labrum present; mandibles with one pair of short setae; setosity extensive. Habits and habitats. Adult I. noveboracensis are associated with various species of Fagaceae, Betulacae and Juglandaceae. Adults appear to prefer white oak (Quercus alba L.) and American beech (Fagus grandifolia Ehrh.). They feed mainly on the bark of shoots, leaf petioles, leaf buds and acorn buds. Eggs are laid in the ground and larvae feed on the roots of the same host plants. Pupation takes place in the soil. A detailed study of the biology has been published by Sanborne (1981). Status of the classification. Once enigmatic in its placement there now seems to be a consensus that this species belongs in a separate and distinct family (Lawrence 1982, Thompson 1992, Alonso-Zarazaga and Lyal 1999). Kuschel (1995) however, placed it in Curculionidae. Relationships to other Curculionoidea are still disputed. Distribution. This family contains only the species Ithycerus noveboracensis (Forster). It is found throughout eastern North America. CLASSIFICATION OF THE NEARCTIC GENERA Ithyceridae Schönherr 1823 Ithycerus Schönherr 1823, 1 sp., I. noveboracensis (Forster 1771), eastern United States into extreme southern Canada. Associated with various species of Fagaceae, Betulacae and Juglandaceae. Pachyrhynchus Kirby 1837 BIBLIOGRAPHY ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL A world catalogue of families and genera of Curculionoidea (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae). Entomopraxis. Barcelona, Spain.