125.€NEMONYCHIDAE€Bedel€1882franz/WeevilCourseResources/... · 2012. 7. 7. ·...

692 · Family 125. Nemonychidae

Description (based onLawrence 1982). Shape elon-gate, slightly convex; length3.0-5.5 mm; color pale brownto black; vestiture of fine shortto moderately long appressedor suberect pubescence. Ros-trum moderately to very longand mostly narrow. Antennaestraight, ending in a weak,loose club of three articles; an-tennal insertions lateral at themiddle or near the apex of therostrum. Labrum distinct, notfused with clypeus. Mandibleswith a small but distinct mola.Maxillae with separate galea andlacinia and maxillary palps flex-ible. Labial palps attached ven-trally near the base of theprementum. Gular sutureswell-developed and separate.Proventriculus lacks sclerotizedplates. Procoxae contiguousand the procoxal cavities nar-

rowly closed posteriorly. Mesocoxal cavities either not closed lat-erally or narrowly so. Elytra without an inner subcostal flange.Hind wing mostly with four anal veins or fewer. Tarsal claws ofsome cleft. Visible sternites of the abdomen are free; pygydiumconcealed by the elytra. Tegmen simple or bilobed apically and themedian lobe with a distinct dorsal plate.

Eggs are undescribed.Larvae (based on Anderson 1991) when mature about 4.0-

4.8 mm in length, of moderate thickness throughout length,strongly �C�-shaped. Body white, covered with mixture of longand short setae. Minute legs present on thorax. Head hypogna-thous, rounded at sides, pigmented, with few to many setae onfrons and epicranium. Frontal sutures complete, reaching articu-lating membrane of mandible. Clypeus not distinguishable fromfrons and incompletely separated from labrum. One pair of ante-rior stemmata. Labrum short, the anterior margin rounded, bear-

ing four pairs of setae. Antenna of a single membranous articlebearing an accessory appendage. Mandible with two apical teeth,an obtuse protuberance on cutting edge, a distinctly producedmolar area with a flattened grinding surface, and one pair of setae.Hypopharyngeal bracon present. Maxillary palp with three ar-ticles, palpiger present or absent. Labial palp of two articles.Premental sclerite present, may be divided medially. Thorax withpronotal sclerite transverse, lightly pigmented or unpigmented,sparsely covered with setae. Legs very small, subconical, of two orthree segments, with or without a terminal claw. Abdomen withfirst eight segments with two dorsal folds and bearing annular orbicameral 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 NorthAmerica, adults are found on male flowers of several pine speciesvery early in the season, often while snow is still on the ground.Adults feed on pollen. After mating, females lay eggs in theflowers where the larvae also feed on pollen. Mature larvae dropfrom the flowers to the ground and pupate in the soil. Pupationcan take from a few months to two years. In some instanceslarvae have been observed feeding on plant parts other thanpollen (Thomas and Herdy 1961). In other areas of the worldgymnosperms such as Araucariaceae and Podocarpaceae, and someprimitive angiosperms (Fagaceae and Ranunculacaeae) also serveas host plants. Whereas pines are the only documented hosts inNorth America, other genera of conifers may serve as hosts.

Status of the classification. This family is bipolar in distri-bution with approximately similar numbers of taxa found in thesouthern temperate zones of South America, New Zealand andAustralia, and in the northern Holarctic Region. Whereas thefamily was once considered to be absent in tropical areas, a fewspecies have recently been collected in Panama and Venezuela onPodocarpus and appear to represent an undescribed genus. TheNorth American fauna has recently been revised (Kuschel 1989)and is well-known. A catalog of the North American species wasprepared by Hamilton (1994).

Distribution. There are 5 genera and 15 species in NorthAmerica. Two additional species in the genus Atopomacer are knownfrom Pinus at high elevations in far northern Mexico. NorthAmerican nemonychid species are generally distributed in the

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 elongaterostrum combined with the presence of a distinct labrum. Adults are found in association with the male pollen-bearing flowers of Pinus species.

FIGURE 1.125. Cimberis compta(LeConte) (from Bright 1993,reproduced with the permission ofthe Minister of Public Works andGovernment Services, 2001)

Family 125. Nemonychidae · 693

western montane and boreal regions and likely occur anywherewhere pines are present.

KEY TO THE NEARCTIC GENERA

1. Second tarsomere truncate at middle, not projectedover base of third (Fig. 2); hind tibia with singleapical spur; tarsal claw with broad basal flange(Fig. 4); elytra with punctures arranged into indis-tinct striae (Rhinorhynchinae) ......... Atopomacer

� Second tarsomere lobed at middle, projected overbase of third (Fig. 3); hind tibia with two apicalspurs; tarsal claw simple, lacking broad basalflange (Fig. 5); elytra with punctures not arrangedinto striae (Cimberidinae) ................................. 2

2(1). Mandibles inserted dorsally, the sockets fully ex-posed in dorsal view (Fig. 6); in lateral view, man-dibles directed obliquely downwards in relationto rostral plane; antennae situated at middle orafter middle of rostrum, as distant from mandibu-lar sockets as combined length of first three ar-ticles (Doydirhynchini) .................... Lecontellus

� Mandibles inserted laterally, the sockets only par-tially exposed in dorsal view (Figs. 7-9); in lateralview, mandibles continuous with rostral plane;antennae situated at middle or before middle ofrostrum, distinctly closer to mandibular socketsthan combined length of first three articles(Cimberidini) ..................................................... 3

3(2). Mandibles evenly curved on outer margin, eacharmed with well-developed tooth on inner margin(Fig. 8) .................................................. Cimberis

� Mandibles angulate on outer margin, unarmed oninner margin (Figs. 7, 9) .................................... 4

4(3). Labrum trapezoidal or broadly rounded at apex, withsix peg-like apical setae in addition to three pairsof dorsal setae (Fig. 7); base of rostrum moder-ately saddled against obliquely rising frons (Fig.10) in dorsal view, with 1-3 low carinae flankedwith rows of confluent punctures; females withsetiferous patches mostly present on one or twoventrites ........................................... Pityomacer

� Labrum triangular, pointed at apex, without any peg-like setae in addition to three pairs of dorsal se-tae (Fig. 9); base of rostrum very deeply saddledagainst vertically rising frons (Fig. 11) in dorsalview, 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), Brit-ish Columbia, Alberta, Washington, Oregon, California, Idahoand Nevada. Adults collected on Pinus species.

Cimberis Gozis 1881, 7 spp., generally distributed. Adults col-lected on Pinus species.

FIGURES 2.125-11.125. 2. Atopomacer orites Kuschel 1989, tarsus; 3. Cimberis elongata (LeConte 1876), tarsus; 4. Atopomacer orites Kuschel1989, tarsal claw; 5. Cimberis turbans Kuschel 1989, tarsal claw; 6. Lecontellus pinicola Kuschel 1989, rostrum apex; 7. Pityomacer carmelitesKuschel 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.

2 35

6

7 8 9 10 11

4

694 · Family 125. Nemonychidae

Pityomacer Kuschel 1989, 3 spp., British Columbia, Alberta, Wash-ington, Oregon, California and Montana. Adults collected onPinus species.

Doydirhynchini Pierce 1916

Lecontellus Kuschel 1989, 3 spp., Washington, Oregon, Californiaand Nevada. Adults collected on Pinus species.

Rhinorhynchinae Voss 1922

Rhinorhynchini Voss 1922

Atopomacer Kuschel 1989, 1 sp., A. ites Kuschel 1989, Coloradoand Arizona. Adults collected on Pinus species apparently at highelevations.

BIBLIOGRAPHY

ANDERSON, D. M. 1991. Nemonychidae (Curculionoidea). Pp.585-586. In: F. W. Stehr, ed. Immature Insects. Volume 2.Kendall/Hunt. Dubuque, Iowa.

BRIGHT, D. E. 1993. The Insects and Arachnids of Canada, Part.21. The Weevils of Canada and Alaska: Volume 1. Centre forLand and Biological Research. Ottawa, 217 pp.

HAMILTON, R. W. 1994. A catalog of the Coleoptera of Americanorth of Mexico. Family: Nemonychidae. USDA AgricultureHandbook 529-134, x + 8 pp.

KUSCHEL, G. 1989. The Nearctic Nemonychidae (Coleoptera:Curculionoidea). Entomologica Scandinavica, 20: 121-171.

LAWRENCE, J. L. 1982. Coleoptera. Pp. 482-553. In: S. P. Parker,ed. Synopsis and Classification of Living Organisms. Volume2. McGraw Hill. New York.

THOMAS, J. B. and H. HERDY. 1961. A note on the life historyof Cimberis elongatus (LeC.) (Coleoptera: Anthribidae). Cana-dian Entomologist, 93: 406-408.

Family 126. Anthribidae · 695

Description: Shape un-usually diverse; elongate anddepressed, to oval, convex, andmite-like, mostly more or lesselongate and convex above;length 0.4 to 40 mm, in U.S.0.4 to 16 mm; vestiture rarelyabsent, of hair-like scalesmostly in mixed shades ofwhite, gray, straw, brown orblack, in the tropics some withbright pink, red, yellow, orgreen. In collections, species areoften confused with ceram-bycids, chrysomelids, ciids,scolytids, and mites.

Head large, rarely retractile into prothorax; if rostrum presentmostly 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 gen-iculate, most with an apical club of three antennomeres, clubfaint or absent in species with elongate antennae, and involvingfrom two to eight antennomeres in some tropical species; in-serted either on the lateral or ventral surface of the rostrum, or onthe face between or below the eyes. Labrum distinct, mostly semi-circular, setose; mandibles unusually large for a weevil, stout,curved, upper surface flattened and with a setose groove, theapices acute, mostly with a postmedian tooth; maxillary palpislender, with four palpomeres, the first very short, the fourthlong; gular sclerite and sutures invisible; postmentum (fusedmentum and submentum) large, deeply emarginate in front,heavily sclerotized; ligula large, corneous, mostly notched to deeplyemarginate in front, rarely entire; labial palpi with three palpomeres,the second shortest, both pairs of palpi elongate, cylindrical, flex-ible, and with acute (very rarely truncate) apices. Eyes dorsal, dorso-lateral, or lateral, mostly large, ranging from entire to deeply emar-ginate, the facets very variable in size and number, and withoutintermixed setae.

Pronotum apex narrower than the maximum head width(except those choragine genera with retractile heads); some exoticgenera with the eyes out on broad stalks; base mostly nearly aswide as the elytra, constricted or produced laterad in some; shape

mostly trapezoidal, apex obliquely truncate, rarely somewhat pro-duced over the head, base truncate with an antebasal transverseridge, this mostly turned forward at the sides forming a short ormore rarely complete side margin (called the lateral carina), thetransverse ridge is always antebasal but it is referred to as basal inthose species in which the surface behind it is more or less vertical;surface smooth, punctate, reticulate, rugose, or tuberculate; pleu-ral region broad, supra-coxal sutures well developed; prosternumshort to long in front of coxae, the intercoxal process narrow(rarely broader than a coxa); the coxal cavities closed behind. Me-sosternum short, the process separating the mesocoxae ex-tremely variable in shape and proportions. Metasternum mostlylong, in some so short that meso- and metacoxae are barely sepa-rate, a transverse groove or suture near posterior margin, and inmany a midventral longitudinal depression or groove. Legs ro-bust to slender; trochantins not exposed; procoxae globular,mostly protruding, not touching or only barely so, and groovedon their inner faces for reception of the intercoxal process ofprosternum; mesocoxae globular, separate; metacoxae elongateand transverse (except Sicanthus Valentine and CisanthribusZimmerman where they are globular or broadly tear-drop shaped),separate, not reaching elytral margin; trochanters moderate withthe apices oblique; femora swollen postmedially, mostly withoutteeth or spines; tibiae slender, without movable spurs, but withan apical hook or tooth in males of some genera; tarsal formula5-5-5, apparently 4-4-4, the second apically emarginate, the thirddeeply bilobed and tomentose beneath, partially embraced by thesecond, and rarely with the lobes fused, the fourth very small andinvisible in dorsal view, the fifth elongate, with claws mostlytoothed or cleft. Scutellum small, varied in shape, hidden in someflightless species. Elytral apices always locking into a longitudinalgroove in pygidial base, pygidial disc always partially exposed inposterior view but may be invisible from above; normally 10elytral striae or groups of punctures plus an abbreviated scutellarrow, rarely 12, 14 or 19-21 rows of punctures; epipleural foldpresent, narrowed or obsolete apically. Wing venation and fold-ing pattern approach that of the Chrysomelidae.

Abdomen with five visible sterna, the four sutures entire,fused, and immovable (rarely the fourth suture flexible); surfacewithout coarse sculpturing. Male genitalia with the penis mostlyslender, curved, with a movable dorsal plate which when raisedgives an open �bird�s beak� appearance, with paired slender basal

126. ANTHRIBIDAE Billberg 1820

by Barry D. ValentineFamily common name: The fungus weevils

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

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 isdirected anteriad, and the elytra usually have an abbreviated scutellar stria.

FIGURE 1.126. Toxonotus cornutus(Say)

696 · Family 126. Anthribidae

struts; parameres and pars basalis fused, the latter with apex mostlyhooded, rarely flattened, the ventral struts fused into a singlemedian structure. Female genitalia sclerotized, the valvifers withlong apodemes, the coxites apically toothed or ridged, the stylilateral, the spermatheca C-shaped.

Larvae are crescent-shaped, subcylindrical, fleshy, widest inthe mid-abdominal region; size 4 to 12 mm in length (the major-ity still undescribed); body with lateral fleshy protuberances and afew to many short or long setae scattered over various segmentsand the head; color near white. Head exserted, hypognathous,setiferous, or rarely retracted into the prothorax, narrower thanprothorax, with epicranial suture present. Antennae reduced, one-segmented 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 galeasetiferous, lacinia acute and inconspicuous; labium withsubmentum, mentum, ligula, and one-segmented palpi. Ocelliabsent. Thorax with legs absent or if present, l-, 2-, or 3-seg-mented, without tarsunguli. Abdomen nine-segmented, withtwo or three plicae per segment, the ninth segment smaller thanthe eighth. Spiracles annular or annular-biforous. Urogomphiabsent. Anderson (1947) provides a partial key.

Habits and habitats. Distributions and biologies of Nearc-tic species are summarized by B. D. Valentine (1999). In general,adults may be found on the larval food plants. Those whoselarvae feed in the stems or receptacles of various weeds appear tofeed upon the pollen of the same plants; species with fungivorouslarvae mostly are found feeding on the surface of the same fungi;those with wood-boring larvae occur on dead or dying tree trunksor branches, and in some cases at least feed upon bark. Adults ofthe majority of anthribid species in the United States are rare inmuseum collections. The best all-around collecting technique isbeating dead or diseased branches, clumps of dead twigs or leaves,or tangles of dead vines. The beating cloth should be as closeunder the plants as possible and must be examined at once, foranthribids recover and fly off much more rapidly than otherweevils. Some of the choragine genera complicate things by beingactive jumpers, and are as a result exasperatingly difficult to catch.Another technique is to range the woods examining the trunksof trees, especially the thin-barked ones, for signs of infirmity,and also all dead branches both above and on the ground. Lookin particular for smooth gray or black patches flush with thewood surface, or small, protruding, black, brown or reddishlumps. These are pyrenomycete fungi, often in the generaHypoxylon and Biscogniauxia, which are ignored by most collec-tors; they are very hard and when cut with a knife have a drycrumbly or charcoal-like consistency. All our species of Eurymycter,Piesocorynus, Choragus, and Euxenus are associated with thesegrowths. Sweeping weedy fields usually produces onlyTrigonorhinus spp., the species T. tomentosus (Say) on commonragweed, T. limbatus (Say) and T. griseus (LeConte) on sneeze- orbitter-weed and other composites, T. alternatus (Say) onfungus-infested morning glory, T. sticticus (Boheman) on smuttygrasses 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 hostsof T. ornatus (Schaeffer), T. strigosus (Jordan), and T. nigromaculatus(Schaeffer) are not known. Sweeping bushy, overgrown areas willsometimes reveal species of Ormiscus or Eusphyrus, especially inearly spring, but beating is better for these and all other membersof our fauna except species of Euparius, which occur on polyporefungi. Larvae of all our species feed on vegetation; they can befound in the twigs and branches of trees, in hard or polyporefungi, or under bark of dead or dying trees. One species, thecoffee bean weevil, Araecerus fasciculatus (DeGeer 1775), lives inseeds and all sorts of dried plant materials from banana flour tostrychnine. A related species in Australia and New Zealand occurson dried fruit, especially apples. Our introduced species of theEuropean genus Anthribus (formerly Brachytarsus) have larvaethat feed on scale insects of the subfamily Lecaniinae; the femalelays each of her eggs under a female scale that has laid her eggs,the weevil larvae then develop exclusively at the expense of theeggs of the scale insect. This is a startling deviation from thephytophagous habits of the Rhynchophora.

Status of the classification. This family has been subjectto considerable nomenclatorial confusion and lack of studies ofhigher taxa beyond all proportions consistent with the rest of theorder. The genera need considerable study. The world cataloguesof Wolfrum (1929, 1953) need many additions and revisions.The Nearctic fauna is summarized by Valentine (1999) who pro-vides a synonymic checklist and keys to most genera. Other basicreferences 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 speciesknown from all regions; 88 described and 32 undescribed speciesoccur in North America. Of 30 North American genera, Araecerusis cosmopolitan; Choragus, Eurymycter, Gonotropis, Allandrus, andTrigonorhinus are holarctic (although the latter extends south toArgentina); Sicanthus is endemic but probably neotropical;Euxenulus, Phoenicobiella, and Araeoderes appear to be North Ameri-can but are obviously related to Latin American groups; and allremaining genera are Neotropical, forming the northern fringe ofan extensive Antillean, Central, or South American fauna.

KEY TO THE NEARCTIC TRIBES AND GENERA

1. Antennae inserted on anterior surface of rostrumor head (Fig. 2); antennal club never with fourantennomeres (Choraginae) ............................. 2

� Antennae inserted on lateral or ventral surface ofrostrum (Fig. 3); or antennal club sometimes withfour antennomeres. (Anthribinae) .................. 10

2(1). Eyes rounded, upper edges not closer together thanlower ................................................................. 3

� Eyes elongate-oval, upper ends closer to each otherthan lower (Choragini) ...................................... 7

3(2). Hind coxae elongate-transverse, almost reachingelytral edge (Araecerini) ................................... 4

� Hind coxae globular or short tear-drop shaped(Cisanthribini) ...................................... Sicanthus


4(3). Head not retractile into prothorax, the eyes too wide;elytra with a scutellar plus 10 striae or rows ofpunctures ......................................................... 5

� Head capable of being retracted into prothorax pastthe eyes (Fig. 2); elytra with 12 or more rows ofpunctures ......................................................... 6

5(4). Lateral prothoracic carina present; transversepronotal carina basal .......................... Araecerus

� Lateral prothoracic carina absent; transversepronotal carina antebasal ....................Neoxenus

6(4). Lateral prothoracic carina double, not upturned atapex; elytra with 12-14 rows of punctures ................................................................ Habroxenus

� Lateral prothoracic carina single, mostly with apexupturned; elytra with 19-21 rows of punctures ........................................................ Acaromimus

7(2). Head not retractile into prothorax, the eyes too wide;pronotum with a raised reticulum forming a honey-comb pattern .................................................... 8

� Head retractile into prothorax past the eyes;pronotum punctate, not reticulate .................. 9

8(7). Antennae with 11 antennomeres ........... Choragus� Antennae with 9 or 10 antennomeres ...................

.................................................. Pseudochoragus

9(7). Elytra with 10 rows or fields of punctures, plus ashort scutellar row ................................ Euxenus

� Elytra with 11 to 15 rows of punctures, one or moremostly incomplete ............................. Euxenulus

10(1). Each elytron partially enclosing anterior part ofscutellum, resulting scutellar notch shaped likean inverted omega (Fig. 4); eyes large, entire, veryfinely faceted (Gymnognathini) ........................................................................... Gymnognathus

� Scutellar notch forming a simple V or U; eyes asabove, or small, or notched, or coarsely faceted....................................................................... 11

11(10). Rostrum with a dorsal, median carina interrupted atbase by an abrupt, small, very deep pit or trans-verse groove (Platystomini) ........................... 12

� Rostrum carinate or not, at most with a basal punc-ture ................................................................. 13

12(11). Lobes of all third tarsomeres separate .. Toxonotus� Lobes of third tarsomeres fused..... Phoenicobiella

13(11). Mandibles with strongly toothed ventral cuttingedge (Fig. 3), as well as normal dorsal edge(Cratoparini) ........................................... Euparius

� Mandibles without a toothed ventral cutting edge....................................................................... 14

14(13). Eye entire, or truncate, or faintly sinuate on ante-rior face .......................................................... 15

� Eye strongly notched or emarginate on anterior face(nearest insertions of antennae) .................... 24

15(14). Entire face with conspicuous white pubescence;center of pronotum with a pit or groove; our spe-cies with a conspicuous patch or band of whitecrossing the suture (Tropiderini) ................... 16

� White pubescence, if present, confined to spots, ifconspicuous, pronotum will have the disc con-cave with a central swelling, and elytra will havemultiple tufts of erect pubescence ............... 17

16(15). Pronotum with a smooth, longitudinal, shallowgroove from central pit to antebasal carina; elytralpale patch antemedian and not reaching sidemargins .............................................. Gonotropis

� Pronotum with a sinuous, transverse, mediangroove; elytral pale band postmedian and reach-ing the side margins ........................ Eurymycter

17(15). Antennal club with 4 antennomeres; rostrum thick,width at apex less than twice depth; scrobesdorso-lateral, interscrobal distance mostly lessthan interoccular distance (Discotenini) ..................................................................... Discotenes

� Antennal club with 1 or 3 antennomeres; rostrumdepressed apically, width at apex more thantwice depth; interscrobal distance more thaninteroccular distance ..................................... 18

18(17). Eyes finely faceted, more than 26 rows across maxi-mum width ...................................................... 19

� Eyes coarsely faceted, 26 or fewer rows acrossmaximum width (Piesocorynini) ...................... 23

19(18). Sides of rostral apex abruptly widened to coverthe laterally protruding mandibular bases; ourspecies with a small post-ocular tooth on apex ofprothorax (Ischnocerini) ................. Ischnocerus

� Sides of rostral apex with mandibular sheathsweakly or not wider than rostral dorsum; no smallpost-ocular teeth on prothoracic apex .......... 20

20(19). Merger of rostrum with venter of head capsule form-ing a broad curve; rostrum long, flattened, andapically flared ................................................. 21

� Merger of rostrum with venter of head capsule indi-cated by a transverse groove or abrupt angle;rostrum shorter, not apically flared (Platyrhinini)....................................................................... 22

21(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 theupper ends of the eyes; lateral prothoracic ca-rina with an apical tubercle ............Trachytropis

� Face with at most a weak concave area at rostralbase; lateral prothoracic carina not toothed atapex.................................................. Goniocloeus

FIGURES 2.126-4.126. 2. Euxenus sp., head, anterior view; 3.Euparius marmoreus (Olivier), head, dorsal view; 4. Gymnognathus sp.,scutellum, dorsal view.

2 3 4


23(18). Eyes with 14 or more rows of facets across maxi-mum width ......................................Piesocorynus

� Eyes with 12 or fewer rows of facets across maxi-mum width .................................. Brachycorynus

24(14). All third tarsomeres with the lobes fused down themidline (Anthribini) .............................. Anthribus

� All third tarsomeres with the lobes separate ..... 25

25(24). Rostrum quadrate, or with rounded apical angles ........................................................................ 26

� Rostrum, excluding mandibles, narrowed from baseto apex; rostral apex with central third longer thansides (Trigonorhinini) ................... Trigonorhinus

26(25). Intercoxal process of mesosternum laterallyangulate, or swollen, or bent; scrobes sulciform,continued across the rostral sides and endingbelow the eyes (Basitropidini) ........................ 27

� Intercoxal process of mesosternum simple; scrobesfoveiform, more or less rounded and not contin-ued transversly or ventrally around and underthe rostrum (Zygaenodini) .............................. 28

27(26). Lateral prothoracic carina extending to apex whereit is toothed; antennal scrobes strongly producedtoward ventral midline ........................... Eugonus

� Lateral prothoracic carina not reaching anterior mar-gin, nor apically toothed, antennal scrobes widelyseparated on venter ......................... Phaenithon

28(26). Lateral prothoracic carina absent or if present notreaching anterior margin ................................ 29

� Lateral prothoracic carina extending to apex ................................................................ Araeoderes

29(28). Transverse pronotal carina clearly antebasal, inca-pable of contacting the elytral base; pronotal hindangles not projecting laterad of the humerae ............................................................... Ormiscus

� Transverse pronotal carina subbasal or basal, ca-pable of contacting elytral base at some point;pronotal hind angles often projecting laterad ofthe humerae ....................................... Eusphyrus


Suprageneric taxa are diagnosed in the generic key. NOTE: Manyundescribed species are included in the comments on distribu-tion. As used below, the designation Neotropical includesAntillean species; Central and South America do not.

Anthribidae Billberg 1820

Choraginae Kirby 1819

Araecerini Lacordaire 1876

Araecerus Schoenherr 1823, 2 introduced spp., including the cof-fee bean weevil, A. fasciculatus (DeGeer 1775): Worldwide. Alsoabout 75 Indopacific species.

Araeocerus Schoenherr 1839Araeocorynus Jekel 1855

Araeosarus Walker 1859Doticus Pascoe 1882Metadoticus Olliff 1890

Neoxenus Valentine 1999, 1 sp., N. versicolor Valentine, 1999, Texasto Panama; four others in Central America and Antilles.

Habroxenus Valentine 1999, 1 sp., H. politus Valentine, 1999, Mary-land, 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 andAntilles.

Acaropsis Jordan 1907Xenorchestes Motschulsky 1873, not Wollaston 1854Euxenus 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 othersalmost worldwide.

Alticopus Villa and Villa 1833

Pseudochoragus Petri 1912, 1 sp., P. nitens (LeConte 1884), Massa-chusetts and Oklahoma; another species in Europe.

Choragus, of American authors, in part, not Kirby 1819Holostilpna, of American authors, not Jordan 1907

Euxenus LeConte 1876, 3 spp., eastern United States; at least sixmore in Central America and Antilles.

Holostilpna Jordan 1907

Euxenulus Valentine 1960, 1 sp., E. piceus (LeConte 1878), southFlorida; also three undescribed in Antilles.

Anthribinae Billberg 1820

Discotenini Lacordaire 1866

Discotenes Labram and Imhoff 1841, 2 spp., Texas; Arizona about10 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


Allandrini Pierce 1930

Allandrus LeConte 1876, 3 spp., United States except the desertsouthwest; also several in Palaearctic.

Tropiderinus Reitter 1916

Stenocerini Kolbe 1897

Stenocerus Schoenherr 1826, 1 sp., S. longulus Jekel 1855, southTexas to Brazil; seven more Mexico to Argentina.

Gymnognathini Valentine 1960

Gymnognathus Schoenherr 1826, 2 spp., south Texas; south Ari-zona; about 90 described and many undescribed Neotropical spe-cies.

Analotes Schoenherr 1839

Tropiderini Lacordaire 1866

Gonotropis Leconte 1876, 1 sp., G. gibbosus LeConte 1876, Canadasouth 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 desertsouthwest.

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 1839Camptotropis Jekel 1855

Brachycorynus Valentine 1999, 3 spp., eastern United States toTexas; a few others in Central America and Antilles.

Tropideres, of LeConte 1876 and others, not Schoenherr1823Brevibarra, of Valentine 1960, not Jordan 1906

Platyrhinini Imhoff 1856

Goniocloeus Jordan 1904, 1 sp., G. bimaculatus (Olivier 1795), east-ern United States to Texas; 36 others south to Brazil.

Tropideres, of various authors, in part, not Schoenherr 1823Strabus Jekel 1860, not Gerstaecker 1860 [Curculionidae]

Trachytropis Jordan 1904, 1 sp., T. arizonicus (Sleeper 1954), south-eastern 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 NewMexico and Arizona; about 35 others throughout the Neotropics.

Anthribus, of authors, in part, not Geoffroy 1762Neanthribus Jordan 1906Pseudanthribus Pierce 1930

Basitropidini Lacordaire 1866

Eugonus Schoenherr 1833, 1 sp., E. bicolor Valentine 1972, south-eastern Arizona; about 30 more in Central and South America.

Schimatocheilus Fahraeus 1839Phaenisor Motschoulsky 1874

Phaenithon Schoenherr 1823, 1 sp., P. platanum (Schaeffer 1906),southeastern Arizona; about 65 others in Central and SouthAmerica.

Camaroderes Jekel 1855Griburiosoma Schaeffer 1906

Zygaenodini Lacordaire 1866

Ormiscus G. R. Waterhouse 1845, 14 described and about 30undescribed spp., eastern United States and desert southwest;probably 200 more throughout the Neotropics, mostlyundescribed.

Entomops Lacordaire 1866Hormiscus Gemminger and Harold 1872Toxotropis LeConte 1876Gonops LeConte 1876

Eusphyrus LeConte 1876, 6 spp., eastern United States and desertsouthwest; 30 more in Central and South America.

Opisthotropis Hoffmann and Tempere 1954

Araeoderes Schaeffer, 1906, 1 sp., A. texanus Schaeffer, 1906, southernTexas and southern Alabama.

Trigonorhinini Valentine 1999

Trigonorhinus Wollaston 1861, 11 spp., southern Canada, entireUnited States; seven others south to Argentina, and in the Pale-arctic Region.

Brachytarsus, of many authors, not Schoenherr 1823Trigonorrhinus Gemminger and Harold 1872Anthribulus LeConte 1876Brachytarsoides Pierce 1930

Cratoparini LeConte 1876

Euparius Schoenherr 1823, 5 spp., eastern United States to Mon-tana and Arizona; at least 70 others in Central and South America,and Japan to Australia.

Cratoparis Dejean 1834Caccorhinus Sharp 1891


Anthribini Billberg 1820

Anthribus Geoffroy 1762, 2 adventive spp. from Europe, Atlanticcoast and ?California.

Brachytarsus Schoenherr 1823Anthotribus Gemminger and Harold 1872Pseudobrachytarsus Pierce 1930

BIBLIOGRAPHY

ANDERSON, W. H. 1947. Larvae of some genera of Anthribidae(Coleoptera). Annals of the Entomological Society of America,40: 489-517, pl. I-IV.

BLATCHLEY, W. S. and C. W. LENG. 1916. Rhynchophora orWeevils of North Eastern America. The Nature Company.Indianapolis, 682 pp., fig. 1-155.

LECONTE, J. L. 1876. The Rhynchophora of America, North ofMexico. Proceedings of the American Philosophical Society,XV (96): i-xvi + 1-455.

PIERCE, W. D. 1930. Studies of the North American weevils ofthe superfamily Platystomoidea. Proceedings of the UnitedStates National Museum, 77(no. 2840): 1-34, pls. 1-5.

TING, P. C. 1936.The mouthparts of the coleopterous groupRhynchophora. Microentomology, 1: 93-114.

VALENTINE, B. D. 1960. The genera of the weevil familyAnthribidae north of Mexico (Coleoptera). Transactions ofthe American Entomological Society, 86: 41-85.

VALENTINE, B. D. 1971. Family Anthribidae. Pp. 243-248. In:M. H. Hatch. Beetles of the Pacific Northwest, Part V. Univer-sity of Washington Publications in Biology, 16: i-xvi + 1-662.pl. I-LV.

VALENTINE, B. D. 1972. Notes on anthribid weevils. III. Newspecies and records primarily from Arizona. ColeopteristsBulletin, 26: 1-11, fig.1-4.

VALENTINE, B. D. 1991. The Choragus-Holostilpna problem(Coleoptera: Anthribidae). Coleopterists Bulletin, 45: 301-307.

VALENTINE, B. D. 1998(1999). A review of Nearctic and somerelated Anthribidae (Coleoptera). Insecta Mundi, 12: 251-296.

WOLFRUM, P. 1929. Anthribidae. Coleopterorum Catalogus,26(102): 1-145.

WOLFRUM, P. 1953. Anthribidae. Coleopterorum Catalogus,supplementum, 26(102): 1-63.

Family 127. Belidae · 701

Description. (based onLawrence 1982). Shape elon-gate, slightly convex; length 3-6 mm; color pale to dark brownor black; vestiture of very fineshort appressed pubescence.Rostrum moderately to verylong and mostly narrow. An-tennae straight, ending in aweak, loose club of three ar-ticles; articles 9 and 10 withdeep apical pockets; antennalinsertions ventral at or near thebase of the rostrum. Labrumfused with clypeus. Labial palpswith two articles and attacheddorsally near the apex of theprementum. Gular suturesfused. Proventriculus lacks scle-rotized plates. Pronotum withcomplete lateral edges.Procoxae contiguous and theprocoxal cavities narrowlyclosed posteriorly. Elytra with-

out an inner subcostal flange. Hindwing with fewer than five analveins. Tarsal article 2 broadly lobed as article 3; tarsal claws simple.Visible sternites of the abdomen free; pygydium exposed by theelytra. Tegmen truncate or slightly emarginate apically and themedian 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 andno epipharyngeal rods. Frontal sutures incomplete, not reachingarticulating membrane of mandible. Maxilla with palp with twoarticles, palpiger absent. Thoracic spiracles on mesothorax. Ab-domen with segments with two dorsal folds. Anal opening ven-tral.

Pupae are undescribed.Habits and habitats. Within this family, only the genus

Rhopalotria occurs in North America. It is restricted to south Floridawhere two species are associated with native and adventive species

of Zamia (Cycadaceae). The biology has been well-studied byNorstog and Fawcett (1989). Adult weevils swarm on male conesof the cycads, where mating, feeding and oviposition occur. Lar-vae feed within the male cones. Weevils also visit female conesbut do not feed. Pollen transport to the female cones occursduring these visits and these beetles (and their relatives elsewhere)may be obligate pollinators of the cycads.

Status of the classification. The family-level classificationof these beetles is somewhat controversial. Crowson (1986) con-sidered them as a distinct family, the Allocorynidae. Kuschel (1995)places them as Belidae, subfamily Oxycoryninae. The most recentcatalog (Alonso-Zarazaga and Lyal 1999), probably followingLawrence (1982) and Thompson (1992), places them as the sub-family Allocoryninae of the Oxycorynidae. Here they are placed asBelidae, subfamily Allocoryninae.

Distribution. There is one genus with two species in NorthAmerica. Both species occur only in Florida; one is adventive andone native.


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. 1999. A worldcatalogue of families and genera of Curculionoidea (Insecta:Coleoptera) (Excepting Scolytidae and Platypodidae).Entomopraxis. Barcelona, Spain.

CROWSON, R. A. 1986. On the systematic position ofAllocoryninae. Coleopterists Bulletin, 40: 243-244.

EMDEN, F. VAN. 1938. On the taxonomy of Rhynchophora larvae(Coleoptera). Transactions of the Royal Entomological Soci-ety of London, 87: 1-37.

127. BELIDAE Schönherr 1826


Family common name: The cycad weevils

The only North American members of this family are odd weevils, recognized by their straight antennae, some-what truncated elytra mostly exposing the last one or two tergites, and the short stout legs with femora whichare 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 1.127. Rhopalotriaslossonae (Schaeffer) (from Bright1993, reproduced with thepermission of the Minister ofPublic Works and GovernmentServices, 2001)

702 · Family 127. Belidae

KUSCHEL, G. 1995. A phylogenetic classification ofCurculionoidea to families and subfamilies. Memoirs of theEntomological Society of Washington, 14: 5-33.

LAWRENCE, J. L. 1982. Coleoptera. Pp. 482-553. In: S. P. Parker,ed. Synopsis and Classification of Living Organisms. Volume2. McGraw Hill. New York.

NORSTOG, K. J. and P. K. S. FAWCETT. 1989. Insect-cycadsymbiosis and its relation to the pollination of Zamia furfuracea(Zamiaceae) by Rhopalotria mollis (Curculionidae). AmericanJournal of Botany, 76: 1380-1394.

THOMPSON, R. T. 1992. Observations on the morphology andclassification of weevils (Coleoptera, Curculionoidea) with akey to major groups. Journal of Natural History, 26: 835-891.

Family 128. Attelabidae · 703

Description: The follow-ing adult characters have beengiven by Thompson (1992)and Kuschel (1995) to supportthe single family status ofAttelabidae: maxillary palps 3or 4 segmented (if 3 seg-mented, elytral punctuationconfused); tergite 8 concealedin both sexes; elytra separatelyand broadly rounded at apices,more or less exposing a py-gidium (tergites 6 and 7); bodysetose, without broad scales;cuticle metallic or otherwisebrightly colored in many; elytrawith scutellary striole in many;abdominal ventrites decreasingin length, first 3 or 4 fused;pygidium visible in both sexes;male genitalia consisting of afree pedon and tectum and alarge tegmen (as in all groupswith non-geniculate antennae);intersegmental membrane be-tween sternites 8 and 9 partlyor entirely sclerotized.

The subfamily Attelabinae is characterized by the following:Adult with body more or less short, stout, glabrous or withappressed pubescence above, color variable; head rectangular toweakly triangular, elongate in males of some; eyes large, mostlyreniform, longest diameter more or less vertical; rostrum short tomoderate in length, distinctly widened apically, weakly arcuate inlateral view; lateral apical angle in some with tooth-like projection;

postmentum in some males with pair of acuminate ventral pro-jections; antennae inserted dorsolaterally, club 3-segmented, moreor less compact, scrobe short, distinct, more or less vertical; man-dibles robust, toothed on inner apical margin, outer margin moreor less rounded; maxillary palps 4-segmented; labial palps indis-tinctly 1 or 2 segmented, located ventrally on prementum; elytramore or less quadrate, not much longer than wide, humeri simpleto strongly protuberant, scutellary striole present or absent, striaebecoming less distinct posteriorly in many; ventral abdominalsutures 1-3 more or less rigid; procoxae prominent, conical; pro-thoracic legs enlarged, profemora distinctly swollen, armed orunarmed; 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 in-tersegmental fold between prothorax and mesothorax;prementum and mentum fused, equally sclerotized; maxillarypalps 2-segmented; prodorsal fold more convex than postdorsalfold, postdorsal fold sloping posteriorly; abdominal tergum 9 atleast twice as long as sternum 9; anus subterminal or ventral;cuticular asperities more or less inconspicuous.

The subfamily Rhynchitinae is characterized by the follow-ing: Adult with body more or less elongate, mostly with distinctsemi-erect to erect setae above, color variable; head mostly trian-gular, widest at the base; eyes small to medium in size, rounded,longest diameter more or less horizonal; rostrum mostly slen-der, parallel-sided and longer than the head, more or less weaklyarcuate in lateral view; antennae inserted laterally, club 3-segmented,more or less loosely united, scrobes shallow, longitudinally ob-lique, not well defined; mandibles flat, toothed on inner andouter margins; maxillary palps 4 segmented; labial palps 2 or 3segmented, inserted laterally or apically on prementum; elytraelongate, mostly distinctly longer than wide, humeri simple,scutellary striole present or absent, striae mostly distinct through-

128. ATTELABIDAE Billberg 1820by 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-genicu-late), mouthpart structure, wing venation, etc. The family in the inclusive sense contains three subfamilies inAmerica 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 theinner margin, the front legs enlarged and the tarsal claws connate. In the Rhynchitinae (tooth nose snout beetles), thebody is more or less elongate, the rostrum mostly narrow and elongate, the mandibles flat and toothed on the inner andouter margins, the legs subequal in size and the tarsal claws appendiculate. In the Pterocolinae (thief weevils), the bodyis short and compact, the short rostrum apically depressed and parallel-sided, the mandibles flat and toothed on theinner and outer margins (as in the Rhynchitinae), the middle and hind femora larger than the front femora, and the tarsalclaws have a broad basal tooth. The Pterocolinae are a unique New World group that are clearly related to the Rhynchitinaealthough the rhynchitine link has not yet been determined.

FIGURE 1.128. Merhynchites bicolor(Fabricius) (from Bright 1993,reproduced with the permission ofthe Minister of Public Works andGovernment Services, 2001)

704 · Family 128. Attelabidae

out; ventral abdominal sutures distinct, suture between 1 and 2rigid; all legs more or less equally developed; tibial apices mostlywith one or two small straight mucros; tarsal claws with longclawlike inner processes; in some inner processes shorter, widerand more toothlike.

Larva with thoracic spiracle located on postero-lateral por-tion of prothorax; prementum and mentum separate, mostlyunequally sclerotized; maxillary palp 2 or 3 segmented; prodorsaland postdorsal folds (segments 6-8) more or less subequal inconvexity; tergum and sternum of abdominal segment 9 subequalin length; anus terminal or subventral, transverse x-shaped orsimple transverse cleft; cuticular asperities more or less conspicu-ous.

The subfamily Pterocolinae is characterized by the following:Adult with body robust and dorsally convex, with appressed tosemi-erect inconspicuous fine setae above, coloration metallic blu-ish-green to bluish-black; head weakly triangular; eyes large, oval,posteriorly more protuberant, anteriorly emarginate; rostrumshort, as long as or shorter than the head, straight, in dorsal viewmore or less parallel-sided throughout, apically depressed, withbeard like ventral setosity (more setose in males); antennae short,subequal to combined length of head and rostrum, inserted nearbasal 1/4 under basirostral ridge; scrobe fossa like; club large,strongly abrupt, compact; mandibles flat, toothed on inner andouter margins; maxillary palps 4-segmented, labial palps 3-seg-mented, inserted apically on prementum; pronotum with dis-tinct lateral carinae; elytra short, apices individually rounded; py-gidium and two complete abdominal terga visible; humeri simple;scutellary striole present; striae mostly distinct throughout; inter-vals more or less convex; all ventral abdominal sutures entire;mesepimeral �side pieces� strongly developed, visible in dorsalview 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 apiceswith 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 subfamilyRhynchitinae have been described (Hamilton 1980, 1983;Hamilton and Kuritsky 1981). They possess taxonomically sig-nificant tuberculate setae and posterior processes associated withabdominal segment 9.

Habits and habitats. All known members of the subfamilyAttelabinae are leaf rollers and apparently cause no serious dam-age to their hosts although one Central American species ofHybolabus has been reported as a serious defoliator of its hosttree, Cariniana pyriformis Miers (Lecythidaceae). All known femaleslay their egg(s) on leaves that they prepare by biting and cuttingwith their mandibles. The leaves are then rolled into a more orless barrel-shaped structure (nidus) that nourishes and protectsthe developing larvae. The enlarged front legs and the uncinatetibial apices manipulate the leaf tissue during nidus formation.Pupation occurs in the ground in the species that have been stud-

ied. Comments on the biology of three North American speciesare available. Packard (1890), Frost (1908), Blatchley and Leng(1916), and Lutz (1935) commented on the biology ofHimatolabus pubescens (Say). Frost (1908) recorded a eulophid para-site 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 bipustulatusFabricius. Most of the information in these papers deals withtaxonomy, 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 thegenera of Attelabini. Also, he associated the niduses of thesespecies, and A. bipustulatus as well, with specific host plants. Vogt(1992) discussed the leaf rolling behavior, host plants and asso-ciated rhynchitid weevils (Pterocolinae) of attelabine weevils oc-curring from Canada to the Republic of Panama. Hamilton(1998),in a revision of the New World Pterocolinae, described 15 newspecies from Central and South America and clarified manyattelabid-pterocoline-host plant associations initiated by Vogt.

Hamilton (1994) provided a summary of known NorthAmerican rhynchitine biology. The larvae of these weevils de-velop in living or dead leaves as leaf miners, in fruits, in cutflower heads, in cut terminal shoots, in terminal buds, in flowerbuds, and in cut leaf primordia. The cuts are made by the adultfemale with her mandibles after oviposition in the specific plantpart. The flower heads, buds or leaf primordia are either com-pletely cut from the plant or are partially cut and eventually dropto the ground. Information has been published on the life stagesor 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). Bovingand Craighead (1931) provided the first basic illustrations of H.aeneus larvae. Hamilton (1973, 1981) provided detailed illustra-tions 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 theeastern rose curculio, M. bicolor. Balduf (1959) published the mostcomprehensive 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 appearsto be the larva of E. angustatus but they did not rear it and theydetermined it as Orsodacne sp. by the �method of elimination andlocality.� Hamilton (1980) provided notes on the biology of E.angustatus (Herbst) and described and illustrated the larva andpupa. Hamilton (1983) discussed the life cycles of A. cassandraeand T. perplexus and provided detailed descriptions and illustra-tions of their immature stages. Hamilton (1994) provided new


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 islacking except for brief comments by Kissinger (1964), given him byVogt, stating that Eugnamptus spp. mine the dead leaves of varioushardwoods and that females of Temnocerus aeratus (Say) lay eggs inand 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) in-cluded three North American species in a paper on the taxonomy ofRhynchophora larvae. He provided key characters for the larvae of H.aeneus, M. bicolor and Rhynchites velatus LeConte and included them ina key to select species of Rhynchitini. He also associated these threespecies 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 onlyNorth American rhynchitine species of known economic impor-tance. Haplorhynchites aeneus attacks commercially grown sunflow-ers while M. bicolor and M. wickhami attack cultivated roses andsometimes damage blackberry and raspberry. Comments on theeconomic 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 informationon the economic importance and control of M. wickhami. Schulzand Lipp (1969) and McBride and Oseto (1978) commented onthe status of damage to sunflowers by H. aeneus.

Vogt (1992) discussed the biology of the Pterocolinae andtheir attelabid hosts and coined the term �thief weevils� for theirhabit of taking over the niduses of their attelabid hosts. Al-though not specifically stated, his comments on the biology ofthese weevils were probably based mainly on observations of P.ovatus in Maryland. According to Vogt, female pterocolines forcetheir way into freshly made attelabid leaf rolls where they eat ordestroy the host egg and oviposit their own. The pterocolinelarva develops rapidly feeding on the decaying leaf tissue andleaves the hollowed out leaf roll to pupate in the ground.

Status of the classification. The subfamilies Rhynchitinaeand Pterocolinae, here considered in the family Attelabidae, havebeen placed in the family Rhynchitidae by other workers. Thefamily name Rhynchitidae is credited to Gistel 1856 (Alonso-Zarazaga and Lyal 1999). John L. LeConte (1876) was the firstAmerican entomologist to use the family name Rhynchitidae forweevils in the subfamilies Rhynchitinae and Pterocolinae. Subse-quent workers (Sharp 1889, Pierce1909, Blatchley and Leng 1916,Ting 1936, Anderson 1991) grouped these weevils in the familyCurculionidae. Pierce (1913) recognized the family Attelabidaeincluding the Rhynchitinae and the new tribes Rhynchitini andAuletini. Voss (1922-1969), in a worldwide taxonomic mono-graph series on the subfamilies Attelabinae, Rhynchitinae andPterocolinae, consistently listed these groups in the familyCurculionidae. Boving and Craighead (1931), in a synopsis of thelarval forms of Coleoptera placed the Rhynchitinae and Attelabinaein the family Attelabidae. Crowson (1955) grouped these weevils

in the family Attelabidae but suggested a possible alternativewould be to establish a family Rhynchitidae for the Rhynchitinaeand Pterocolinae. Lawrence (1982) also placed these weevils in thefamily Attelabidae. O�Brien and Wibmer (1982), in an annotatedchecklist of North American weevils, recognized the familiesRhynchitidae and Attelabidae but Wibmer and O�Brien (1986),in their annotated checklist of South American weevils, regroupedthese weevils into the single family Attelabidae. Thompson (1992)using abdominal characters, etc. lumped the three subfamilies inthe family Attelabidae. More recently Kuschel (1995), in a phylo-genetic approach, recognized only the family Attelabidae and Farrell(1998), in a molecular approach, also grouped the rhynchitinesand attelabines in the family Attelabidae. Hamilton (1969 to 1998)has consistently used the family Rhynchitidae for the Rhynchitinaeand Pterocolinae. Most recently, Alonso-Zarazaga and Lyal (1999)have recognized the family Rhynchitidae in their world catalogueof families and genera of Curculionoidea.

The family name Attelabidae is credited to Billberg (1820). Thegenus Attelabus, on which the family name is based, was originallydescribed by Linnaeus (1758). Linnaeus included only A. coryliLinnaeus in the genus and it was therefore the type by monotypy.Since the time of Linnaeus, many species from all over the worldhave been added to the genus Attelabus. Olivier (1807) decided thatA. coryli was generically different from the other species that had beenplaced in Attelabus and described the new genus Apoderus with 13species including A. coryli. Attelabus coryli is technically the type ofAttelabus but was not recognized by Olivier, or anyone else, untilBedel (1888) pointed out that Attelabus should be used for thosespecies placed by Olivier in Apoderus. Bedel proposed the name Cyphusfor the species that were left in Attelabus by Olivier as well as manyother species placed there by subsequent authors. Bedel�s proprosalfollowed the International Code but subsequent workers, unawareof the change or unwilling to change the names of numerous spe-cies, have not followed Bedel. Silfverberg (1977) successfully appealedto the International Commission on Zoological Nomenclature toaccept common usage of Attelabus in the interest of stability and toconfirm the designation by Schoenherr (1823) of Attelabuscurculionoides Linnaeus 1767 (= Curculio nitens Scopoli 1763) as type-species. Two major groups of attelabids are now officially recognized- Apoderus with A. coryli as its type and Attelabus with Attelabus nitensScopoli as its type. Jekel (1860) divided the genus Attelabus into 16subgenera and Voss (1925) elevated most of them to the genericlevel. The genus Attelabus is applicable to only two species of attelabidsin America, north of Mexico - A. bipustulatus Fabricius and A. nigripesLeConte.

Distribution. The family includes 1,914 world species in 97genera (Kuschel 1995). These weevils occur throughout the worldbut Kuschel points out that they do not occur in New Zealand,New Caledonia and the Pacific Islands. In the New World, 362species have been described including 180 species of leaf rollingweevils in four subfamilies (Attelabinae, Euscelinae, Hybolabinaeand Pilolabinae), 162 species of Rhynchitinae and 20 species ofPterocolinae. In America north of Mexico, there are 51 total spe-cies in 11 genera including 6 species of Attelabinae, 44 species ofRhynchitinae and one species of Pterocolinae.


KEY TO NEARCTIC SUBFAMILIES AND GENERA

1. Tarsal claws appendiculate (Figs. 7 and 8); man-dibles depressed, toothed on inner and outermargin; front legs subequal to middle and hindlegs or middle and hind femora larger than frontfemora; tibial apices unarmed or with small spursor mucros ......................................................... 2

� Tarsal claws connate (Fig. 9); mandibles robust, nottoothed on outer margin; front legs enlarged, dis-tinctly larger than middle and hind legs; profemoradistinctly swollen (Fig. 2); tibial apices uni-unci-nate (male) or bi-uncinate (female) (Figs. 5 and 6)(Attelabinae) ................................................... 10

2(1). Prothorax with distinct lateral carina (Fig. 3);propleura strongly excavated beneath carina;body small, robust; metallic bluish-green to blu-ish-black (Pterocolinae) ...................... Pterocolus

� Prothorax not laterally carinate (Fig. 4); propleuranot strongly excavated; body size, shape andcolor variable (Rhynchitinae) ........................... 3

3(2). Scutellary striole present .................................... 4� Scutellary striole absent ..................................... 6

4(3). Pygidium completely or almost completely coveredby elytra; elytra with some erect setae; maleswith one tooth on outer edge of mandibles andfemales with two ............................ Eugnamptus

� Pygidium mainly exposed, not completely or almostcompletely covered by elytra; elytra withouterect setae; mandibular teeth similar in both sexes......................................................................... 5

5(4). Elytral striae distinct, more or less quadrate, moder-ately to deeply impressed; intervals narrowerthan width of striae, convex, smooth; intervalpunctures much smaller than striae; pubescenceinconspicuous, more or less appressed; bodydark colored, in some feebly metallic bronze orblue, less than 4 mm in length ........ Temnocerus

� Elytral striae more or less indistinct (more distinctin M. bicolor), weakly impressed, especially pos-teriorly; discernable intervals wider than widthof striae, more or less flat, in some minutely rug-ose; interval punctures numerous, as large ornearly as large as striae; pubescence conspicu-ous, fine, semi-erect; body variable reddish-or-ange and black to brownish-black with faint blu-ish metallic luster, greater than 4 mm in length ...................................................... Merhynchites

6(3). Elytral striae distinctly rowed; intervals distinct, withpunctures more or less smaller than striae; punc-tures not masked by pubescence ................... 7

� Elytral striae not distinctly rowed; intervals indis-tinct, punctures as large or nearly as large asstriae; punctures in some masked by pubescence......................................................................... 9

7(6). Elytra short, exposing pygidium and two abdominalterga; hind basitarsal segment longer than com-bined length of hind tarsal segments 3 and 4 ............................................................... Deporaus

� Elytra not short, covering all abdominal terga andpart of the pygidium; hind basitarsal segmentshorter or subequal to hind tarsal segments 3and 4 combined ................................................ 8

8(7). Elytral intervals wide, more or less flat, with numer-ous punctures, punctures in some as large ornearly as large as striae; males mostly more se-tose along elytral suture at declivity; antennalclub symmetrical in both sexes ......................................................................... Haplorhynchites

� Elytral intervals narrow, more or less convex, mod-erately punctured; punctures much smaller thanstriae; males not more setose along elytral sutureat declivity; males with asymmetrical antennalclub ..................................................... Involvulus

9(6). Body with violaceous to greenish metallic luster;elytral pubescence of appressed whitish setaeand widely rowed, dark, erect setae; males withanterolateral spine on each side of pronotum(Fig.4); length mostly greater than 4 mm ........................................................................... Rhynchites

� Body without violaceous or greenish metallic lus-ter; elytral pubescence not as above; males with-out 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, gla-brous except for a few scattered fine erect setae....................................................................... 11

2 3

5

6

7

8

94

FIGURES 2.128-9.128. 2. Himatolabus pubescens (Say), male, lateralhabitus; 3. Pterocolus ovatus (Fabricius), male, lateral habitus; 4. Rhynchitesvelatus LeConte, male, lateral habitus; 5. Himatolabus pubescens (Say),female, protibial apex; 6. Himatolabus pubescens (Say), male, protibialapex; 7. Pterocolus ovatus (Fabricius), tarsal claw; 8. Haplorhynchitesaeneus (Boheman), tarsal claw; 9. Himatolabus pubescens (Say), tarsalclaw.


11(10). Profemora unarmed in both sexes; submentum inmales with pair of ventrally projecting acuminatespines; ventral rostral apex without median coni-cal prominence .............................................. 12

� Profemora in males armed with one or two bluntspine like projections; profemora in females un-armed or armed with a single peg like projection;submentum in males without pair of ventral pro-jecting accuminate spines; ventral rostral apexin lateral view with median conical prominence(more pronounced in females) ............ Attelabus

12(11). Body uniformly shiny black to brownish-black; headin both sexes subequal in size and shape; sur-face of abdominal sternites smooth, without tu-bercles; pronotum with pair of basilateral pitlikedepressions ..................................... Xestolabus

� Body bicolored; pronotum, elytra and abdomen redto reddish-orange; legs, sterna and head black;head elongate in males; females with abdominalsternites 1-3 with pair of acute tubercles;pronotum without pair of pitlike pronotal depres-sions .............................................. 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 leafrolls; A. bipustulatus is associated with Quercus spp. and Carpinussp. and ranges from SE Canada throughout the eastern U.S. andsouthwest to OK and TX; A. nigripes is associated with Rhuscopallina L. and other Rhus sp. and ranges throughout the easternhalf of the U.S. and south into Mexico; the two species can bedistinguished by color. A. bipustulatus is black with reddish hu-meral maculae while A. nigripes is reddish throughout with vari-able darker areas (western form of A. nigripes is darker overallwith pale areas in basal half of elytra).

Homoeolabus Jekel 1860, 1 sp., H. analis (Illiger); females are leafrollers; larvae develop in leaf rolls; associated with many Quercussp.; ranges from SE Canada throughout the eastern half of theU.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 leafrolls; both species are associated with Quercus spp.; H. pubescensalso rolls the leaves of Alnus incana (L.) and Corylus americanaWalt. in northeastern North America; H. pubescens ranges fromSE Canada and the northeastern U.S. southwest into TX, NMand AZ and south through Mexico; H. axillaris is recorded in thesouthwestern states of AZ and UT and ranges south into Mexico;The two species can easily be separated by color. H. pubescens isreddish- brown throughout (with blackish-brown areas and darkerextremities in some) and H. axillaris is black to dilute black with

reddish humeral maculae; (key to New World species Hamilton1992).

Xestolabus Jekel 1860, 1 sp., X. constrictipennis (Chittenden); fe-males are leaf rollers; larvae develop in leaf rolls; occurs in thesouthwestern states of AZ and NM and south into Mexico;associated with Rhus toxicodendron L. and Rhus spp. in the south-western U.S.

Rhynchitinae Gistel 1856

Rhynchitini Gistel 1856

Eugnamptus Schoenherr 1839, 9 spp.; known larvae are leaf min-ers; associated with species of Quercus, Sassafras, Juglans, Carya,Cornus, Liquidambar, etc.; distributed in the eastern half of theU.S. and southwestern states of AZ and NM; Eugnamptusangustatus (Herbst) is the most abundant species in North Americaand occurs in the eastern half of the U. S. in 4 sympatric colorforms (key to Nearctic species, Hamilton 1990).

Haplorhynchites Voss 1938, 6 spp., larvae develop in flower headscut by females; associated with various composite species ofHelianthus, Silphium, Coreopsis, Viquiera, etc.; H. aeneus is the mostcommon North American species and is widely distributed fromsouth central Canada throughout the middle and eastern U.S.;the other species occur in the southwestern states (key to Nearcticspecies, Hamilton 1974).

Involvulus Schrank 1798, 1 sp., I. hirtus (Fabricius); larvae developin cut terminal shoots of Quercus spp.according to Vogt inKissinger (1964); ranges throughout northeastern U.S. and south-west 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 rangesthroughout the western half of the U.S. and southwesternCanada; 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 cutterminal shoots, buds and leaf primordia; associated with a widevariety of hosts including species of Quercus, Comptonia, Acacia,Manzanita, etc.; regionally distributed in U.S. (key to Nearctic spe-cies, Hamilton 1971).

Pselaphorhynchites Schilsky 1903

Rhynchites Schneider 1791, 1 sp., R. velatus LeConte (Fig. 4); larvaedevelop in fruits of the desert plum, Prunus andersonii Gray; re-corded only from CA and NV.


Auletini Desbrochers 1908

Auletobius Desbrochers 1869, 9 spp.; known larvae develop in cutterminal leaf primordia; associated with a wide variety of hostsincluding species of Quercus, Comptonia, Potentilla, Eriogonum, etc.;regionally distributed in U.S. (key to Nearctic species, Pierce 1909;key to World species, Voss 1933-37).

Deporaini Voss 1929

Deporaus Samouelle 1819, 1 sp., D. glastinus LeConte; larvae areleaf miners; associated with Q. wislizenii in southern CA andother Quercus spp. throughout its range; ranges throughout west-ern 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 rollthief, north of Mexico the larvae develop in leaf rolls prepared bythree species of attelabine weevils (H. pubescens, A. bipustulatusand H. analis); distributed throughout eastern half of U.S., south-west to TX and AZ and south into Mexico (key to New Worldspecies, Hamilton 1998).

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VOSS, E. 1941. Monographie der Rhynchitinen-Tribus Rhynchitini.2. Gattungsgruppe: Rhynchitina. V. 2. Teil der Monographieder Rhynchitinae-Pterocolinae. (45. Beitrag zur Kenntnis derCurculioniden). Mittheilungen Muenchener EntomologischeGesellschaft, 31: 628-680.

VOSS, E. 1965. Die Unterfamilie Camarotinae, ihre Beziehungenzur Familie Attelabidae sowie ein Versuch, diephylogenetischen Zusammenhange innerhalb der letzterenzur Darstellung zu bringen (Coleoptera, Curculionidae) (191.Beitrag zur Kenntnis der Curculioniden). EntomologischeAbhandlungen Staatliches Museum fur Tierkunde in Dresden,32 : 221-244.

VOSS, E. 1969. Monographie der Rhynchitinen-Tribus Rhynchitini.2. Gattungsgruppe: Rhynchitina. V. 2. Teil der Monographieder Rhynchitinae-Pterocolinae. Entomologische Arbeiten,aus dem Museum G. Frey, 20: 117-375.

WIBMER, G. J. and C. W. O�BRIEN. 1986. Annotated checklistof the weevils (Curculionidae, sensu lato ) of South America(Coleoptera: Curculionoidea). Memoirs of the AmericanEntomological Institute, 39: i-xvi, 1- 563.

Family 129. Brentidae · 711

Description (based onLawrence 1982). Shape variableelongate, narrow and parallel-sided (Brentinae, Cyphag-oginae, Trachelizinae), orstouter and more robust withexpanded oval elytra and a dis-tinctive �pear-like� shape(Apioninae, Nanophyinae); flatto convex; length 1.0-40.0 mm;color pale brown to black, veryrarely red or bicolored or withcontrasting colors or markings;vestiture lacking, or of fineshort to moderately long ap-pressed or suberect pubescence;rarely with scales.

Rostrum moderately tovery long and usually narrowand cylindrical; sexual dimor-phism in rostrum form(Brentinae, Cyphagoginae,Trachelizinae) or length(Apioninae) generally evident.Antennae straight or rarely gen-iculate (Nano-phyinae), funiclevery narrow and slender, rarelyfiliform or moniliform

(Brentinae, Cyphagoginae, Trachelizinae), ending in a weak, looseclub (Nanophyinae) or a compact club (Apioninae, Cyladinae) ofthree articles. Antennal insertions lateral at the middle of therostrum or rarely dorsal near the base. Maxillary palps of two orthree articles. Labial palps small, of one or two articles, attacheddorsally near the apex of the prementum, rarely in cavities. Proven-triculus lacks sclerotized plates. Front coxae vary from contiguousto widely separated. Middle coxae and hind coxae narrowly towidely separated. Front tibiae rarely equipped with an antenna-cleaning 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 con-cealed by the elytra. Tegmen bilobed or notched apically and themedian lobe with a �trough-like� ventral plate and a narrow dor-sal plate.

Larvae (based on Lawrence 1982) when mature about 2.0-18.0 mm in length, slender and subcylindrical (Brentinae) or broadand strongly curved (Apioninae). Small legs rarely present onthorax (Brentinae), usually absent, but with well-developed pedallobes (Apioninae). Frontal sutures complete although rarely in-distinct, reaching articulating membrane of mandible. Antennaof a single membranous article. Hypopharyngeal bracon present.Abdomen with first eight segments with two, three or four dor-sal folds and bearing annular or rarely biforous spiracles.

Habits and habitats. As noted, Brentidae is a composite oftaxa of subfamily rank each of which has a rather different bodyform. Natural history aspects of Brentidae similarly are correlatedwith, and best discussed, at the subfamily level (Thomas 1996).

Brentinae, Cyphagoginae and Trachelizinae are the traditionalbrentids. They are small to large, elongate weevils that are usuallyfound under bark as both adults and larvae. Where known, fe-males lay their eggs in holes they have excavated with their ros-trum in living, dying or recently felled hardwood trees. Larvaetunnel deep into the heartwood and appear to feed on wood andfungus mycelia. Brentus anchorago Linnaeus is one of NorthAmerica�s largest weevils, with males reaching a body length ofup to 40 mm. In some species, adult males guard females duringegg-laying.

Cyladinae includes only the adventive Cylas formicarius (Fab-ricius), the sweet potato weevil. Adults and larvae are associatedwith various Convolvulaceae (especially Ipomoea pescapreae L.) andare pests of sweet potatoes, Ipomoea batatas L. Larvae mine thestems.

Little is known about Nearctic Nanophyinae. Most PalearcticNanophyinae are associated with plants of the family Lythraceae,especially the genus Lythrum, but also Crassulaceae, Tamaricaceaeand Ericaceae. Larvae are associated with the fruits, leaves andstems and many appear to cause galls. One species of NanophyesSchoenherr has been introduced into North America for biologi-cal control of Lythrum salicaria L. (purple loosestrife; Lythraceae)

129. BRENTIDAE Billberg 1820

by Robert S. Anderson and David G. Kissinger

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

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

As used here, Brentidae is a composite of primitive weevil forms, not recognizable by any one or a fewcharacters. Nearly all of them, with the exception of only the Nanophyinae, have straight, non-geniculateantennae. Apioninae and Nanophyinae are small �pear-shaped� weevils immediately recognized by a long cylin-

drical trochanter with the femur attached at its apex, a character state that is known in no other beetles. Brentinae,Cyphagoginae and Trachelizinae have elongated, parallel-sided bodies and a long, generally straight rostrum that oftenexhibits marked sexual dimorphism. Cyladinae are the sweet potato weevils, recognizable by their unique habitus.

1.129. Arrenodes minutus (Drury)(from Bright 1993, reproducedwith the permission of the Ministerof Public Works and GovernmentServices, 2001)

712 · Family 129. Brentidae

(Harris 2001) and a second isunder consideration. Zeugonyxsabinae Notman has beenreared from galls on Juniperusashei Buchh. in Texas, but oth-erwise no information is avail-able on the native NorthAmerican fauna.

Apioninae are associatedwith a wide variety of plants,mostly Fabaceae and Aster-aceae, but also Malvaceae,Polygonaceae, Caprifioliaceae,and Umbelliferae. Many of thenewly elevated genera, for-merly subgenera of Apion sensulato, are host specific on certainfamilies 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 spe-cies are adventive. Rhopalapion longirostre (Olivier) is associatedwith the ornamental hollyhock, Althea rosea Cav. (Tuttle 1954)and Ischnopterapion virens (Herbst) with clover, Trifolium (Hoebekeet al. 2000). Two species of Exapion have been introduced intothe western United States as biological control agents for gorse,Ulex europaeus L. and scotch broom, Cytisus scoparius (L.) (bothFabaceae) (O�Brien 1995) and Omphalapion hookerorum (Kirby) hasbeen introduced for biological control of scentless chamomile,Matricaria perforata Mérat (Harris and McClay 2001). Podapiongallicola Riley 1883 has been associated with Pinus, larvae occur ingalls on twigs (Bright 1993).

Status of the classification. At present there is continuingcontroversy over the classification of the members of this family.Alonso-Zarazaga and Lyal (1999) recognize Brentidae (includingBrentinae, Cyphagoginae, Trachelizinae and Cyladinae) as distinctfrom Apionidae and Nanophyidae and accord each family levelstatus. This system is different from others such as Lawrence(1982) which recognizes Brentidae (including Brentinae,Cyphagoginae and Trachelizinae) and Apionidae (includingApioninae, Nanophyinae and Cyladinae). The family level classi-fication used here follows Lawrence and Newton (1995) andgroups all of these taxa within Brentidae; however, at thesubfamilial level, within Brentidae, the classification followsAlonso-Zarazaga and Lyal (1999). O�Brien and Wibmer (1982)provide a checklist of the North American species of Nanophyinaeand species of Apioninae (all as Apion sensu lato).

Brentinae, Cyphagoginae, Trachelizinae and Cyladinae are well-known in North America and their classification remains un-changed (Warner 1960; Kissinger 1968). Brentinae have 3 speciesin 3 genera, and Cyphagoginae, Trachelizinae and Cyladinae eachhave only 1 genus and 1 species represented. Apioninae andNanophyinae on the other hand have undergone substantialchanges at the generic level over the last few years. In Nanophyinae,

there are now 5 genera and 5 species recognized in North America.This is based on phylogenetic evidence and follows the divisionof the once larger widespread genus Nanophyes Schoenherr byAlonso-Zarazaga (1989) and followed by Alonso-Zarazaga andLyal (1999). The situation with Apioninae is even more complexwith many of the various subgenera (e.g., Kissinger 1968) nowelevated to generic rank, many generic concepts have been nar-rowed with some North American species transferred into previ-ously Palearctic genera, and a number of new genera have beenproposed (Kissinger 1989, 1990, 1992, 1998, 1999; Alonso-Zarazaga 1990). Alonso-Zarazaga (1990) treats the Palearctic clas-sification of Apioninae in detail and is the basis for the develop-ment and understanding of the new system as applied in NorthAmerica. At present in North America there are approximately

FIGURES 3.129-4.129. 3. Front leg, Brentinae (schematic); 4. Frontleg, Apioninae (schematic).

3 4

2.129. Trichapion centrale (Fall)(from Bright 1993, reproducedwith the permission of the Ministerof Public Works and GovernmentServices, 2001)

FIGURES 5.129-6.129. 5. Dorsal habitus, Cylas formicarius (Fabricius1798); 6. Lateral view, pronotum and base of elytra, Cylas formicarius.

5

6


140 species placed in the 19 genera. A small number of NorthAmerican species have not yet been assigned to a genus in thisnew system. Identifying Apioninae is no easy task and the key tothe genera of Apioninae formerly placed in Apion sensu lato (start-ing at couplet 15) is based extensively on male characters only.Also, many of the characters used in establishing the new genericclassification are based on male genitalia and require dissectionsand detailed study.

Distribution. Collectively this family is widely distributedin North America, but some of the constituent subfamilies havevery restricted distributions. Apioninae are widespread in NorthAmerica and found in most habitat types from sea level to thealpine zone. Nanophyinae are also widespread but do not extendas far north as do Apioninae, nor do they include the diversity ofhabitats of the latter group, but then there are also many fewerspecies. Excluding Cyladinae, which has only the one adventivespecies, Cylas formicarius Fabricius, Brentinae, Cyphagoginae andTrachelizinae are generally tropical in their distributions and 4 ofthe 5 species that occur in North America only do so in extremesouthern Florida (Brentus anchorago (Linnaeus), Stereodermus exilisSuffrian, and Paratrachelizus uncimanus (Boheman)) or southernTexas (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 tothe side of trochanter (Fig. 3); body more elon-gate and narrow, length greater than 3.0 mm, mostgreater than 10.0 mm (Figs. 1, 5, 14) ................ 2

� Trochanter long and somewhat cylindrical, femurattached to the apex of trochanter (Fig. 4); body�pear-shaped�, length less than 5.0 mm, most lessthan 3.0 mm (Figs. 2, 16, 31) ............................. 7

2(1). Pronotum with a broad constricted collar (that isalmost as long as the rest of the pronotum) be-fore base (Fig. 5); mesothorax with a deep verti-cal sulcus near the posterior margin of thepronotum (Fig. 6); tarsal claws connate at base;elytra elongate-oval in dorsal view, lacking im-pressed striae (Fig. 5) (Cyladinae) ............. Cylas

� Pronotum with at most a very short constriction be-fore base (Figs. 1, 7, 9, 10, 14); mesothorax lack-ing a deep lateral pit near the posterior margin ofthe pronotum; tarsal claws free; elytra elongateand parallel-sided in dorsal view, with distinctlyor indistinctly impressed striae ....................... 3

3(2). All femora lacking distinct tooth (although slight an-gulation may be present on front femur); rostrumwith ventral surface punctate and with densemicrosetae or sparse elongate recumbent setae......................................................................... 4

� At least front femur with obvious tooth; rostrum withventral surface smooth or coarsely punctate, notsetose (Brentinae) ............................................ 5

4(3). Elytra with stria 3 bent inwards at middle towardselytral suture and coalescent with stria 2 (Fig. 7);front tibia with inner edge incised in apical one-

FIGURES 7.129-13.129. 7. Dorsal habitus, Stereodermus exilis Suffrian;8. Front leg, Stereodermus exilis; 9. Dorsal habitus, Paratrachelizusuncimanus (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.

7

8

9 10

11

12

13


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

7(1). Antenna geniculate, scape distinctly longer thancombined length of first 3 articles of funicle; fu-nicle apparently with only 5 articles, articles 6and 7 as wide as apical antennal article and form-ing a loose club (Nanophyinae) ....................... 8

� Antenna straight, not geniculate, scape shorter thancombined length of first 3 articles of funicle; fu-nicle with 7 articles, articles 6 and 7 narrow, notas wide as apical antennal article and distinctfrom compact apical club (Apioninae) ............ 12

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

....................................................................... 11

9(8). Elytra with interstria 8 with a row of small, dark tu-bercles (Fig. 15); elytra largely pale, with dark,oblique fascia extended from basal one-third atmiddle to humeri; vestiture consisting of sparse,erect setae and long, decumbent, moderatelydense scales intermixed; length 2.0-2.3 mm ......................................................... Pseudotychius

� Elytra with interstria 8 smooth in basal one-quarter;elytral vestiture various; length 1.2-1.9 mm .. 10

10(9). Elytra lacking fascia, setae appressed and uniformlyarranged, each seta short, not or hardly exceed-ing the base of the following one (Fig. 16);pronotum and tibia lacking specialized erect se-tae, only one subhumeral seta on elytral interstria9 (Fig. 16) ............................................... Microon

� Elytra with fascia, setae arcuate and disordered,each seta long, usually exceeding the base ofthe following one (Fig. 17); pronotum, tibia andodd elytral interstriae with specialized erect se-tae (Fig. 17) ...................................... Nanophyes

11(8). Antenna with basal 5 articles of funicle slender;elytra largely pale, with basal dark area; vestiturefine and short, consisting of whitish and darkscales .......................................... Nanodactylus

� Antenna with basal 4 articles of funicle slender;elytra uniformly testaceous; vestiture of longwhitish setae ...................................... Zeugonyx

12(7). Middle coxae contiguous, not separated by junc-tion between mesosternal process and intercoxalprocess of metasternum ................................ 13

� Middle coxae separated by junction betweenmesosternal process and intercoxal process ofmetasternum................................................... 15

13(12). Tarsal claws with acute basal tooth .... Chrysapion� Tarsal claws simple, at most swollen as base, not

toothed........................................................... 14

14(13). Rostrum straight, in side view not forming angle atjunction with frons, apical half of rostrum usuallytapered to apex (Fig. 18); eyes not or scarcelyprominent; antennae inserted at basal 0.29-0.47of rostrum; scape at most as long as width of ros-trum at midlength; prescutellar fovea conspicu-ous, lanceolate to sulciform; tarsi normal, fronttarsus with first article 1.25-1.60 X as long as wide;internal sac of aedeagus without baculi, other

half, incision surrounded by very dense setae(Fig. 8); rostrum with ventral surface with sparseelongate recumbent setae (Cyphagoginae) ........................................................... Stereodermus

� Elytra with stria 3 evident only as a row of shallowpunctures, straight throughout length, not coa-lescent with stria 2 (Fig. 9); front tibia with inneredge straight, not incised; rostrum with ventralsurface with dense microsetae in basal portion(Trachelizinae) .......................... Paratrachelizus

5(4). Head longer than broad, not distinctly constrictedbehind eyes, lateral margins continuous, pro-longed and convergent behind eyes (Figs. 10,11); pronotum medially deeply sulcate in basalone-half (Figs. 10, 11); male with pronotum ex-tremely elongated, lateral margins constrictednear middle (Fig. 10) .............................. Brentus

� Head short, width more or less equal to length, dis-tinctly constricted behind eyes, lateral marginsat most slightly prolonged but interrupted by con-striction, not convergent behind eyes (Figs. 1,12, 14); pronotum uniformly smooth dorsally, notsulcate (Figs. 1, 14); male with pronotum not elon-gated, and lateral margins not constricted nearmiddle ............................................................... 6

6(5). Antenna with articles 2 to 11 more or less equal inlength and width towards apex (Fig. 1); head con-stricted behind eyes, hind angles not projecting(Figs. 1, 12); front tibia with at most a swelling oninner margin, no distinct tooth evident (Fig. 13);rostrum very dissimilar in male (short and broadwith large mandibles, Fig. 12) and female (narrowand cylindrical, Fig. 1) ........................ Arrenodes

� Antenna with articles 2 to 11 increasing in lengthand width towards apex (Fig. 14); head stronglyconstricted behind eyes, hind angles projecting;front tibia with distinct tooth on inner margin (Fig.14); rostrum dissimilar in male (dilated at apex)

FIGURE 14.129. Dorsal habitus, Heterobrenthus texanus Schaeffer,male.


structures may be present; associated withPolygonaceae ...................................... Perapion

� Rostrum strongly curved, in side view forming aconspicuous angle at junction with frons, apicalhalf of rostrum parallel-sided to slightly widenedto apex (Fig. 19); eyes prominent; antennae in-serted at basal 0.10-0.19 of rostrum; scape longerthan width of rostrum at midlength; prescutellarfovea inconspicuous, similar to pronotal punc-tures; tarsi robust, front tarsus with first article0.85-1.00 X as long as wide; internal sac ofaedeagus with two baculi and denticles; associ-ated with Pinus (Pinaceae) .................... Podapion

15(12).* Front femur of male with polished (sometimes stri-ate) area on ventral surface, the area generallylimited posteriorly by a prominent longitudinalcarina (Fig. 20); metasternum generally with a pairof spicules near the middle of the posterior mar-gin ........................................................ Fallapion

� Front femur of male without polished area on ven-tral surface ..................................................... 16

16(15). Pygydium of male lacking distinct, deep, transversepreapical sulcus, profile of pygydium not inter-rupted (Figs. 21, 22); abdominal ventrite 5 of malerounded apically; elytra with specialized seta oninterstria 7 lacking ......................................... 17

� Pygydium of male with distinct, deep, transversepreapical sulcus, profile of pygydium distinctlyinterrupted by sulcus (Figs. 23, 24, 27, 28); ab-dominal ventrite 5 of male truncate apically; elytrawith specialized seta on interstria 7 at middle ornear apical 1/3 present or lacking ................ 19

* From this point on the key is complex and relies extensively oncharacters of males. Males can be recognized by the shorter, morecoarsely sculptured rostrum and (sometimes) by an upward deflectionof the last abdominal ventrite. These are the taxa traditionally treatedas Apion sensu lato and the reader may wish to consult Bright (1993)and Kissinger (1968) for species level identifications.

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

15 16 17

18 19

20

21 22

25

2324

26 27 2829

30 3132 33 34 35


17(16). Antennal club with sutures obsolete or absent; tar-sal claws simple, not toothed; on Matricariaperforata (Asteraceae) ................. Omphalapion

� Antennal club with sutures distinct; tarsal clawstoothed at base; on Malvaceae ...................... 18

18(17). Pronotum with vestiture directed away from the mid-line at the base, more or less parallel to the mid-line along the lateral margins, and perpendicularto 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 mid-line, in some cases, totally transverse at the basalor apical margins (Fig. 26); pronotum with basalflange obsolete (Fig. 26); on Althea rosea(Malvaceae) .................................... Rhopalapion

19(16). Elytra with transverse pattern of contrasting lightand dark scales; pygydium of male with sulcusincomplete laterally, not reaching side marginsof pygydium (Fig. 24); front tarsus with article 2stout, about as wide as long; femora and tibiaerobust; middle coxae widely separated (0.25-0.30x diameter of middle coxa); aedeagus with inter-nal sac with large apically bifurcate structure; onRutaceae, Caprifoliaceae ...................... Neapion

� Elytra with scales more uniformly colored, not con-trasting light and dark; pygydium of male withsulcus complete laterally, reaching side marginsof pygydium (Figs. 27, 28); front tarsus with ar-ticle 2 usually longer than wide; femora and tibiaeslender; middle coxae moderately widely sepa-rated (<0.25 x diameter of middle coxa); aegeaguswith internal sac without large structure; onFabaceae and other plants ............................. 20

20(19). Genitalia of male with tegmen with prostegium ar-ticulated with free ring (Fig. 29) ..................... 21

� Genitalia of male with tegmen with prostegium fusedwith free ring (Fig. 30) .................................... 28

21(20). Tibiae of male simple, not mucronate ................ 22� Middle tibia (at least) of male mucronate ........... 27

22(21). Scrobe with dorsal margin produced into a long,slender acute process projected ventrally; elytrawith interstria 7 with specialized seta insertednear midlength of stria; metasternum very con-vex in lateral view; meso- and metasternal pro-cesses directed inwardly where meeting betweenhind coxae; on Fabaceae ...................... Exapion

� Scrobe with dorsal margin simple, not produced intoa long, slender acute process; elytra with interstria7 or 9, or 7 and 9, with or without specializedseta inserted at or behind apical 1/3 of elytron;metasternum slightly convex in lateral view; meso-and metasternal processes either flat or withmetasternal process more prominent where meet-ing between hind coxae; on various plants .. 23

23(22). Body color red throughout; lateral margins ofpronotum parallel in basal one-half (Fig. 31); malegenitalia with prostegium with high median cristaprojected basally; elytral interstria 2 prolongedoutward at apex (Fig. 32); on Polygonaceae ........................................................................ Apion

� Body color black or with metallic sheen, occasion-ally with light colored appendages; lateral mar-

gins of pronotum various; male genitalia withprostegium with at most moderately high basalmedian carina not projected basally; elytralinterstria 2 only slightly prolonged outward atapex (Fig. 33); on Fabaceae ........................... 24

24(23). Elytra blue-green; elytral interstria 7 with one spe-cialized seta in apical one-third; legs dark; lateralmargins of pronotum parallel in basal one-half;rostrum not prominently expanded at point of in-sertion of antennae; eyes prominent ................................................................. Ischnopterapion

� Elytra black; elytral interstriae 7 and 9 with one spe-cialized seta near apex; legs various, may be lightin color; lateral margins of pronotum various inbasal one-half; rostrum variously expanded atpoint of insertion of antennae; eyes prominent ornot .................................................................. 25

25(24). Pronotum without basal flange, with lateral marginsnearly evenly convergent from base to apex,subconical in dorsal form; rostrum prominentlyexpanded at point of insertion of antennae; legsand article 1 of antenna dark ...............Apionion

� Pronotum with or without basal flange, with lateralmargins various; rostrum smooth at point of inser-tion of antennae, not expanded; legs and article1 of antenna dark or light ............................... 26

26(25). Pronotum with basal flange indistinct, subconicalin dorsal form (Fig. 34); legs and article 1 of an-tenna light; aedeagus with median lobe and teg-men narrow, elongate, subcylindrical; internal sacwith large teeth (>0.030 mm long) ...... Sayapion

� Pronotum without basal flange, lateral marginssubparallel in basal one-half; legs and article 1 ofantenna light or dark; aedeagus with median lobeslightly depressed, broader at base; internal sacwith largest teeth (<0.020 mm long) .... Kissingeria

27(21). Head with ventral surface with subcephalic ridgeslacking or low, not extended basally on head topoint equivalent to middle of eye; vestiture ofinterstriae 2-5 sparse, generally inconspicuous,consisting of one row of scales; one specializedseta on interstria 9; aedeagus with parameroidlobe with one or more macrochaetae; on Betulapapyrifera (Betulaceae) .................... Betulapion

� Head with ventral surface with subcephalic ridgesmoderately to well developed, extended to orbeyond the middle of the eye; vestiture ofinterstriae 2-5 relatively dense, consisting of twoor more rows of scales; one specialized seta oneach of interstriae 7 and 9; aedeagus withparameroid lobe generally lacking macrochaetae;associated mainly with Fabaceae and Asteraceae.......................................................... Trichapion

28(20). Elytra blue in color, legs dark; tarsus with clawsimple; rostrum of male markedly expanded later-ally at point of antennal insertion (Fig. 35); fronsflat, striate ................................. Mesotrichapion

� Elytra not blue in color; rostrum of male smooth, notexpanded laterally at point of antennal insertion....................................................................... 29

29(28). Male with metasternum tuberculate on median pos-terior margin and apical area of parameroid lobe


bearing four or more macrochaetae more than0.055mm long ................................ Eutrichapion

� Male metasternum not tuberculate or parameroidlobe lacking macrochaetae more than 0.010mmlong ........................................ Coelocephalapion


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), gener-ally distributed in eastern United States, into extreme southernCanada. Adults are found under bark especially of oaks; larvaebore into the wood (Buchanan 1960). Males are territorial andguard females during egg-laying (Sanborne 1983).

Brentus Panzer 1788, not Fabricius 1787Arrhenodes Schoenherr 1826Eupsalis Lacordaire 1866Platysystrophus Kleine 1917

Heterobrenthus Sharp 1895, 1 sp., H. texanus Schaeffer 1915, south-ern Texas.

Cyphagoginae Kolbe 1892

Stereodermini Sharp 1895

Stereodermus Lacordaire 1866, 1 sp., S. exilis Suffrian 1870, south-ern 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), generallydistributed in southern United States. Adventive onConvolvulaceae including Ipomoea batata (L.) Lam. (sweet potato).

Cylanus Rafinesque 1815Protocylas Pierce 1941

Apioninae Schoenherr 1823

Apionini Schoenherr 1823

Apion Herbst 1797, 1 sp., identity uncertain, Alberta. Adults havebeen collected around edges of high elevation snowfields. Asso-ciated with Polygonaceae in Palearctic Region.

Apiolum Kirby 1808Apionus Rafinesque 1814Apius Billberg 1820, not Panzer 1806Apium Agassiz 1846Oxystomum Gistel 1856Oxeostomum Gistel 1856Erythrapion Schilsky 1906

Aplemonini Kissinger 1968

Perapion Wagner 1907, 4 spp., generally distributed. Associatedwith 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 distrib-uted in eastern United States into extreme southern Canada; alsoin California and Oregon. Associated with Pinus spp.

Aspidapiini Alonso-Zarazaga 1990

Alocentron Schilsky 1901, 6 spp., generally distributed. Associatedwith 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), Brit-ish 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 west-ern 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, Ulexeuropaeus 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 ineastern North America west to southern Texas, north into ex-


treme southern Canada. Subgenus Neotropion, 1 sp., N. xanthoxyli(Fall), associated with Rutaceae (genus Zanthoxylum); subgenusNeapion, 5 spp., associated with Caprifoliaceae (genus Viburnum).

Xixias Kissinger 1990Neotropion Alonso-Zarazaga 1990 (valid subgenus)

Malvapiini Alonso-Zarazaga 1990

Rhopalapion Schilsky 1906, 1 sp., R. longirostre (Olivier 1807), gen-erally distributed. Adventive on Althea rosea Cav. (hollyhock;Malvaceae).

Oxystomatini Alonso-Zarazaga 1990

Oxystomatina Alonso-Zarazaga 1990

Eutrichapion Reitter 1916, 3 spp., generally distributed. Associ-ated with Fabaceae.

Cnemapion BokorLeconteapion 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. Asso-ciation with Fabaceae (genus Astragalus).

Trichapiina Alonso-Zarazaga 1990

Betulapion Ehret 1994, 1 sp., B. simile (Kirby 1811), generally dis-tributed. 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. Associ-ated mostly with Fabaceae and Asteraceae.

Synapiina Alonso-Zarazaga 1990

Ischnopterapion Bokor 1923, 1 sp., I. virens (Herbst 1797), NewYork, Pennsylvania, Maryland, New Jersey, Connecticut, Dela-ware and Virginia. Adventive on various species of clover, Trifo-lium (Fabaceae) (Hoebeke et al. 2000).

Piezotrachelini Voss 1959

Chrysapion Kissinger 1968, 2 spp., Arizona, California and Texas.

Fallapion Kissinger 1968, 30 spp., generally distributed. Associ-ated mostly with Asteraceae and Umbelliferae.

Incertae sedis

Apionion Kissinger 1998, 2 spp., A. crassum (Fall 1898), easternUnited States; A. dilatatum (Smith 1884), Arizona. Associatedwith Fabaceae.

Coelocephalapion Wagner 1914, 22 spp., generally distributed. As-sociated mostly with Fabaceae and Asteraceae.

Sayapion Kissinger 1999, 5 spp., eastern United States, Texas andArizona. 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 eastto Manitoba. At least one Palearctic species is associated with thegenus Lythrum (Lythraceae).

Nanodactylus Blatchely 1922, 1 sp., N. obesulus Blatchley 1922, Illi-nois, Indiana, Texas.

Nanophyes Schoenherr 1838, 1 sp., N. marmoratus (Goeze 1777),Manitoba, New York; introduced into Canada in 1997 for thebiological control of Lythrum salicaria L. (Lythraceae) (Harris 2001).This weevil species is also approved for introduction into theUnited States and is established in the vicinity of Ithaca, NewYork. A second species, N. brevis Boheman 1845, also is underconsideration for introduction.

Pseudotychius Blatchley 1922, 1 sp., P. watsoni Blatchley 1922, east-ern 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 asheiBuchh. (Cupressaceae).

BIBLIOGRAPHY

ALONSO-ZARAZAGA, M. A. 1989. Revision of the supraspecifictaxa in the Palaearctic Apionidae Schoenherr, 1823. 1. Intro-duction and subfamily Nanophyinae Seidlitz, 1891 (Co-leoptera, Curculionoidea). Fragmenta Entomologica, 21: 205-262.

ALONSO-ZARAZAGA, M. A. 1990. Revision of the supraspecifictaxa in the Palaearctic Apionidae Schoenherr, 1823 (Co-leoptera, Curculionoidea). 2. Subfamily ApioninaeSchoenherr, 1823: Introduction, keys and descriptions.Graellsia, 46: 19-156.

ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL. 1999. Aworld catalogue of families and genera of Curculionoidea(Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae).Entomopraxis. Barcelona, Spain.


BRIGHT, D. E. 1993. The Insects and Arachnids of Canada. Part21. The weevils of Canada and Alaska: Volume 1. Coleoptera:Curculionoidea, excluding Scolytidae and Curculionidae.Publication 1882. Research Branch, Agriculture Canada. Ot-tawa, Canada.

BUCHANAN, W. D. 1960. Biology of the oak timberworm,Arrhenodes minutus. Journal of Economic Entomology, 53:510-513.

HARRIS, P. 2001. Nanophyes marmoratus (Goeze). Flower-feedingweevil. http://res2.agr.ca/lethbridge/weedbio/agents/ananmar.htm

HARRIS, P. and A. MCCLAY. 2001. Omphalapion hookeri Kirby.Seed-head weevil. http://res2.agr.ca/lethbridge/weedbio/agents/aomphook.htm

HOEBEKE, E. R., R. A. BYERS, M. A. ALONSO-ZARAZAGA,and J. F. STIMMEL. 2000. Ischnopterapion (Chlorapion) virens(Herbst) (Coleoptera: Curculionoidea: Brentidae: Apioninae),a Palearctic clover pest new to North America: recognitionfeatures, distribution, and bionomics. Proceedings of theEntomological Society of Washington, 102: 151-161.

KISSINGER, D. G. 1968. Curculionidae subfamily Apioninae ofNorth and Central America with reviews of the world generaof Apioninae and world subgenera of Apion Herbst (Co-leoptera). Taxonomic Publications. South Lancaster, Massa-chusetts.

KISSINGER, D. G. 1989. Apionidae from North and CentralAmerica. Part 1. Notes on the classification of the Apionsubgenus Trichapion Wagner with description of new speciesfrom the United States of America (Coleoptera). InsectaMundi, 3: 271-227.

KISSINGER, D. G. 1990. Apionidae from North and CentralAmerica. Part 2. Description of a new subgenus and two newspecies of Apion from Mexico. (Coleoptera). Insecta Mundi,4: 33-40.

KISSINGER, D. G. 1992. Apionidae from North and CentralAmerica. Part 4. Generic classification and introduction to thegenus Coelocephalapion Wagner, with new species from Mexicoand Venezuela (Coleoptera). Insecta Mundi, 6: 65-77.

KISSINGER, D. G. 1998. Apionidae from North and CentralAmerica. Part 5. Description of the genus Apionion and 4 newspecies (Coleoptera). Insecta Mundi, 12: 93-102.

KISSINGER, D. G. 1999. Description of a new genus, Sayapion,from North and Central America (Coleoptera: Apionidae).Insecta Mundi, 13: 72.

LAWRENCE, J.L. 1982. Coleoptera, Pp. 482-553. In: S. P. Parker,ed. Synopsis and Classification of Living Organisms. Volume2. McGraw Hill. New York.

LAWRENCE. J. F. and A. F. NEWTON, Jr. 1995. Families andsubfamilies of Coleoptera (with selected genera, notes, refer-ences and data on family-group names). Pp 779-1006. In: J.Pakaluk and S. A. Slipinski, eds., Biology, phylogeny andclassification of Coleoptera: Papers celebrating the 80th birth-day of Roy A. Crowson. Muzeum i Instytut Zoologii PAN,Warsaw.

O�BRIEN, C. W. 1995. Curculionidae, premiere biological controlagents. Memoirs of the Entomological Society of Washing-ton, 14: 129-136.

O�BRIEN, C. W. and G. J. WIBMER. 1982. Annotated checklistof the weevils (Curculionidae sensu lato) of North America,Central America, and the West Indies (Coleoptera:Curculionoidea). Memoirs of the American EntomologicalInstitute, 34: i-ix, 1-382.

PESCHKEN, D. P., K. C. SAWCHYN and D. E. BRIGHT. 1993.First record of Apion hookeri Kirby (Coleoptera: Curculionidae)in North America. Canadian Entomologist, 125: 629-631.

SANBORNE, M. 1983. Some observations on the behaviour ofArrhenodes minutus (Drury) (Coleoptera: Brentidae). Coleop-terists Bulletin, 37: 106-113.

THOMAS, M. C. 1996. The primitive weevils of Florida (Co-leoptera: Brentidae: Brentinae). Florida Department of Agri-culture. Division of Plant Industry. Entomology Circular No.375. 3 pp.

TUTTLE, D. M. 1954. Notes on the bionomics of six species ofApion (Curculionidae, Coleoptera). Annals of the Entomo-logical Society of America, 47: 301-307.

WARNER, R. E. 1960. The genus Stereodermus new to Americanorth of Mexico with a revised key to the genera of Brentidae.Coleopterists Bulletin, 14: 29.

720 · Family 130. Ithyceridae

Description (based onLawrence 1982): Body large,stout. Pubescence of scale-likebristles, distinct.

Head with single gular su-ture; pregular sutures absent.Rostrum broad, stout, notsexually dimorphic; antennalinsertions lateral. Maxilla with-out lacinia; maxillary palp ofthree articles, rigid, partially re-tracted into large palpifer; labialpalp of three articles; labrumabsent; mandible stout. An-tennae straight, moniliform;antennal club compact, of threearticles. Proventriculus withoutsclerotized plates.

Notosternal suture com-plete. Procoxae contiguous, mexocoxae narrowly separated andmetacoxae moderately widely separated. Hind wing with fouranal veins. Legs with trochanter short; triangular; femur attachedto side of trochanter. Tarsal claws toothed. Abdomen withventrites 1 and 2 fused together but separated by distinct suture.Cap piece of tegmen strongly bilobed; the median lobe with abroad ventral plate and a narrow dorsal plate.

Egg (based on Sanborne 1981) with length 1.2-2.2 mm.Subspherical, longer than wide. External surface of chorion ofhexagonal facets with impressed borders. Surface strongly punc-tate, with numerous aeropyles.

Larva (based in part on Sanborne 1981, Lawrence 1991) whenmature about 20-25 mm in length. Body relatively short andbroad, subcylindrical, strongly c-shaped. Body yellowish, withheavily sclerotized mouth frame and mandibles. Vestiture ofmoderately long, scattered setae. Head protracted and hypogna-thous, about as long as wide. Frontal sutures complete, reachingarticulating membrane of mandible. Clypeus distinguishable fromfrons and completely separated from labrum. Three pairs of st-emmata on each side. Labrum free, bearing one sensillum andfour pairs of setae. Antenna of a single dome-like article bearinga sensorium, a bifurcate appendage and several setae. Mandiblewith two apical teeth, and three blunt teeth on inner edge, acces-sory ventral process and mola absent. Hypopharyngeal braconpresent. Maxilla with palp with 2 articles, palpiger present. Labial

palp of two articles; palps widely separated. Premental scleritesubtriangular. Thorax with pronotal sclerite moderately pig-mented. Legs very small, widely separated, subconical, two orthree jointed. Abdomen with first four segments with three dor-sal 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 associatedwith various species of Fagaceae, Betulacae and Juglandaceae.Adults appear to prefer white oak (Quercus alba L.) and Americanbeech (Fagus grandifolia Ehrh.). They feed mainly on the bark ofshoots, leaf petioles, leaf buds and acorn buds. Eggs are laid inthe ground and larvae feed on the roots of the same host plants.Pupation takes place in the soil. A detailed study of the biologyhas been published by Sanborne (1981).

Status of the classification. Once enigmatic in its place-ment there now seems to be a consensus that this species be-longs in a separate and distinct family (Lawrence 1982, Thomp-son 1992, Alonso-Zarazaga and Lyal 1999). Kuschel (1995) how-ever, placed it in Curculionidae. Relationships to otherCurculionoidea are still disputed.

Distribution. This family contains only the species Ithycerusnoveboracensis (Forster). It is found throughout eastern NorthAmerica.


Ithyceridae Schönherr 1823

Ithycerus Schönherr 1823, 1 sp., I. noveboracensis (Forster 1771),eastern United States into extreme southern Canada. Associatedwith various species of Fagaceae, Betulacae and Juglandaceae.

Pachyrhynchus Kirby 1837

BIBLIOGRAPHY

ALONSO-ZARAZAGA, M. A. and C. H. C. LYAL. 1999. Aworld catalogue of families and genera of Curculionoidea(Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae).Entomopraxis. Barcelona, Spain.

130. ITHYCERIDAE Schönherr 1823


Family common name: The New York weevil

Among the primitive weevils with straight, non-geniculate antennae, this enigmatic family contains only onespecies, whose adults can be recognized by their large size (12-18 mm long), stout form, and distinct pubes-cence.

FIGURE 1.130. Ithycerusnoveboracensis (Forster) (from Bright1993, reproduced with thepermission of the Minister ofPublic Works and GovernmentServices, 2001)

Family 130. Ithyceridae · 721

KUSCHEL, G. 1995. A phylogenetic classification ofCurculionoidea to families and subfamilies. Memoirs ofthe Entomological Society of Washington, 14: 5-33.

LAWRENCE, J.L. 1982. Coleoptera. Pp. 482-553. In: S. P. Parker,ed. Synopsis and Classification of Living Organisms. Volume2. McGraw Hill. New York.

LAWRENCE, J. L. 1991. Ithyceridae (Curculionoidea). Pp. 590-591. In: F. W. Stehr, ed. Immature Insects. Volume 2.Kendall/Hunt. Dubuque, Iowa.

SANBORNE, M. 1981. Biology of Ithycerus noveboracensis (Forster)(Coleoptera) and weevil phylogeny. Evolutionary Mono-graphs, 4: 1-80.

THOMPSON, R.T. 1992. Observations on the morphology andclassification of weevils (Coleoptera, Curculionoidea) with akey to major groups. Journal of Natural History, 26: 835-891.

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