722 · Family 131.

Curculionidae

131. CURCULIONIDAE Latreille 1802

by Robert S. Anderson

Family common name: The weevils or snout beetles

W
eevils are one of the most diverse groups of organisms. Over 60,000 species have been described world
wide and their diversity in North America is challenged among beetles perhaps only by Staphylinidae. Wee-
vils are associated with virtually all kinds of plants and plant parts. Most feed on living plants but some are
saprophagous. Weevils are immediately recognizable by their elongate rostrum (or snout), with mouthparts situated at
the apex, geniculate antennae and compact antennal club. Some weevils in the subfamilies Entiminae, Cossoninae and
Scolytinae have the rostrum reduced in form and not markedly produced anteriorly. Traditional considerations of the
weevils do not include Scolytinae and Platypodinae but increasing evidence suggests these beetles are derived from
within Curculionidae.

Description (based on in some, dorsal. Mandibles of some bearing a scar at apex or

Lawrence 1982): Shape very deciduous process. Maxillae in some concealed by expanded men-
variable, broadly oval to elon- tum, a few with distinct galea and lacinia. Labial palpi of one or
gate, slightly flattened to mark- two articles, rarely absent; in some weevils palpi inserted in cavi-
edly convex, most covered with ties on the ventral surface of the prementum. Proventriculus of
recumbent or appressed some lacking sclerotized plates. Front coxae contiguous or sepa-
vestiture of scales, some with rated, middle and hind coxae variable. Tarsi of 5 articles but article
metallic sheen or forming con- 4 very small and hidden between lobes of article 3 (exception,
trasting patterns, some Raymondionyminae with only 4 articles); tarsal claws of some
subglabrous or with erect or connate and simple or with a basal process or tooth. Abdomen
suberect hairs only; length with first two ventrites connate, very rarely free. Pygydium formed
from 1-40 mm (most 2-20 by tergite VII or VIII, in most concealed beneath apex of elytra,
mm); color variable, typically exposed and/or sulcate in some. Cap piece of tegmen may be
black or dark brown, more reduced and may or may not be bilobed, occasionally absent;
rarely of other colors. aedeagus with a trough-like ventral plate and membranous dor-
Eyes present, may be re- sally; in some aedeagus with separate pedon and tectum.
duced or absent. Rostrum vary- Larvae (based on Lawrence 1982) subcylindrical, slightly
ing from very short and indis- curved; lightly sclerotized and grublike; usually with very fine
FIGURE 1.131. Sphenophorus tinct to very long and narrow; hairs. Head hypognathous and free, rarely retracted into protho-
pertinax (Olivier)
most sexually dimorphic with rax. Frontal arms “v-shaped” and not reaching mandibular ar-
female rostrum longer, finer and with position of antennal inser- ticulations, endocarina usually present. Stemmata absent in most.
tion more basal. Antennae geniculate (very few exceptions where Antennae of 1 or 2 articles and apical article sometimes a conical
scape is very short and position of antennal insertion on rostrum sensorium. Frontoclypeal suture present. Labrum free, usually
is basal); club of three articles (sometimes with one), compact, in with 4 pairs of setae. Maxillae with galea and lacinia fused to form
some the apical articles recessed in glabrous basal article; funicle mala, maxillary palpi usually of 2 articles. Labial palpi of 1 or
of 5-7 articles, slender; point of antennal insertion on rostrum is rarely, and indistinctly, of 2 articles. Abdominal tergites usually
various, mostly between midlength and apex, mostly lateral but with 3 or 4 transverse plicae. Thoracic spiracles found on the
prothorax or between prothorax and mesothorax. Legs absent.
Acknowledgments: This chapter includes major contributions by Habits and habitats. The habits and habitats of Scolytinae,
Robert J. Rabaglia (Scolytinae), Henry A. Hespenheide (Conoderinae), long treated as a separate family, are summarized under that sub-
Boris A. Korotyaev (Ceutorhynchinae), and Anne T. Howden family heading.
(Entiminae), to whom I am deeply grateful. I also thank the many Weevils can be found associated with just about any kind of
people who contributed to this chapter in other ways, by providing plant in any terrestrial or freshwater habitat. Most species are
specimens, literature or advice, by checking text or keys, or by answering strictly phytophagous as adults and larvae and usually have a
my many questions. These people are Charles W. O’Brien, Horace R. narrow range of suitable host plants. Most species are associated
Burke, Boris A. Korotyaev, Miguel Alonso-Zarazaga, Anne T. Howden,
with angiosperms but a few are associated with gymnosperms,
Donald E. Bright, Chris Lyal, Enzo Colonnelli, and Steve Lingafelter.
Line illustrations are by Nadine Dupérré of Laval, Quebec. Paul
mainly the various conifers in the Pinaceae. Adult and larval feed-
Skelley and Mike Thomas provided the needed push to get it done. ing habits vary extensively but can loosely be classified into two
groups: one in which both adults and larvae are polyphagous
 Family 131. Curculionidae · 723

FIGURES 2.131-3.131. 2. Lateral view of a generalized curculionid head; schematic; 3. Lateral habitus of a generalized curculionid; schematic
(both after Kissinger 1964)

(Entiminae), and one in which adults and larvae have a more or near water. Most of these taxa are best found at night when
restricted range of host plants (other subfamilies). Among the adults come up onto the plants to feed. A few weevils are found
polyphagous species, the larvae feed externally in the soil on roots in intertidal situations (e.g., many Cossoninae, Emphyastes, and
whereas the adults feed generally on foliage. Species with more Thalasselephas) where they develop in driftwood or seaweed. There
restricted ranges of hosts usually feed little as adults (often visit- are many weevils in arid habitats such as deserts (Entiminae) and
ing flowers) or feed on foliage or reproductive structures, and grasslands (Baridinae and Ceutorhynchinae), likely because of
their larvae feed internally in the stems, roots, leaves or reproduc- their associations with the plants that dominate those habitats. A
tive structures of a few congeric or confamilial plant taxa. Some great number of weevils that have immigrated to North America
weevil larvae in the Hyperinae and Ceutorhynchinae feed exter- from Europe are likely associated with imported ornamental plants
nally on foliage and reproductive structures. Pupation usually or amongst ballast brought by ships at the turn of the 19th cen-
takes place in the host plant or in the soil but species of Hypera tury.
and Cionus construct a loosely woven cocoon that is attached to Various groups of weevils are also common as Quaternary
the host plant. fossils in northern North America and are important in recon-
Adults of some weevil species (Raymondionyminae and structing the late Cenozoic history of northern habitats (Matthews
Molytinae) have reduced eyes or are eyeless and live in the soil or 1982).
leaf litter. Some weevil species in the Conoderinae, Cossoninae, Obviously a more complete summary of the natural history
Cryptorhynchinae and Molytinae feed in dead plant material, usu- of Curculionidae is beyond the scope of these few notes.
ally wood. Some species appear to live in association with ants, Status of the classification. The classification of the wee-
although this appears an obligate relationship only for vils was regarded by Crowson in 1955 as the last great problem to
Liometophilus (Cryptorhynchinae). Some species of Entiminae are be clarified within the Coleoptera. While there have been many
parthenogenetic. Most Entiminae as well as some Crypto- advances in the classification, much still remains to be resolved.
rhynchinae and Molytinae are flightless. The classification used herein largely is that of Alonso-Zarazaga
Curculionidae are a very important group economically. Some and Lyal (1999) with a few changes in placement and ranking of
species are serious pests of ornamental, agricultural and forestry certain taxa. A total of 18 subfamilies are recognized. Lawrence
plants and have well-known common names (e.g., boll weevil, and Newton (1995), the classification at the family level adopted
white pine weevil, strawberry root weevil, black vine weevil, etc.). for this book, recognize only 6 subfamilies within Curculionidae,
Recently, species have become increasingly used in the biological demoting many subfamilies to tribes within their Curculioninae.
control of introduced pest plants (e.g., Neochetina, Hylobius, They also refer to the Entiminae as Brachycerinae although the
Cyphocleonus, Eustenopus, etc.) particularly in western North Ameri- constitution remains basically the same. They recognize
can grasslands and southeastern aquatic habitats (O’Brien 1995). Dryophthorinae as a separate family but not Raymondionyminae
An excellent review of the biology of Anthonomini is by Burke and Erirhininae, all three of which are recognized as families in
(1976). the classifications of Thompson (1992) and Alonso-Zarazaga
Some subfamilies as Erirhininae, Bagoinae, Cyclominae and and Lyal (1999). These authors consider these as having family
Ceutorhynchinae have a number of genera and species associated level status because they do not share the same derived male
with freshwater macrophytes. Many of these species are very good genitalic structure as the Curculionidae sensu stricto. Herein all are
swimmers (Morris 1995) and adults spend most of their time in considered subfamilies within Curculionidae. The classification
724 · Family 131. Curculionidae

of Kuschel (1995) is very similar to that of Lawrence and New- the different names being used for different degrees of develop-
ton (1995) but includes Ithyceridae within the subfamily ment and positioning of the apical tooth. Associated with this is
Brachycerinae of Curculionidae. the use of the term ‘apical comb of setae’ which I use to apply to
Catalogs are available for some groups of Curculionidae in the row of setae that may be across the apex of the hind tibia or
North America (e.g., Howden 1993; O’Brien 1986, 1989, 1996, in some weevils is displaced by a change in position of the apical
1997) and an annotated checklist (and supplements) with full tooth to be oriented longitudinally to the main axis of the tibia.
synonyms, information about keys, and distributions has been We do use ‘corbel’ and related terms in the keys to Entiminae,
published (O’Brien and Wibmer 1982, 1984; Wibmer and O’Brien contrary to the recommendations of Thompson (1992). See
1989). A review of the state of knowledge about immatures is by Thompson (1992) for details.
Burke and Anderson (1976). Excellent (but outdated) regional
works to the species level are those of Hatch (1971) for the Pacific CLASSIFICATION OF THE NEARCTIC SUBFAMILIES AND TRIBES
Northwest and Downie and Arnett (1996) for northeastern North
America. Blatchley and Leng (1916) remains an old standard. Curculionidae Latreille 1802
Many of the keys used herein are modified from Kissinger (1964). I. Dryophthorinae
Distribution. Curculionids are found just about everywhere 1. Dryophthorini
in North America. Diversity is greatest in the southern United 2. Orthognathini
States but no recent regional counts are available. The last tabula- 3. Rhynchophorini
tion for the Nearctic Region as a whole was in 1978 by O’Brien II. Erirhininae
and Wibmer who counted 239 genera and 2388 species. Bousquet 4. Erirhinini
(1991) recorded almost 700 species in Canada and Alaska. Ander- III. Raymondionyminae
son (1993a) counted 249 species in 115 genera in extreme south- 5. Raymondionymini
ern Florida alone. Many recent additions to the fauna are the IV. Curculioninae
result of deliberate introductions for biological control purposes 6. Curculionini
but also, a number of taxa recently added to the North American 7. Acalyptini
fauna are from extreme southern Florida or Texas and are recent 8. Anthonomini
discoveries. The species Isochnus arcticus (Korotyaev 1976) is found 9. Cionini
as far north as Ellesmere Island at almost 82 degrees north lati- 10. Derelomini
tude. 11. Ellescini
Some weevil species are routinely intercepted at ports of 12. Mecinini
entry of foreign materials (especially agricultural products) into 13. Otidocephalini
the United States and Canada. Some of these taxa have tradition- 14. Rhamphini
ally or occasionally been considered as part of the North Ameri- 15. Smicronychini
can fauna and included in keys and faunal lists. At present, there 16. Storeini
is no evidence to suggest they are established in North America 17. Tychiini
and they are not included in the key. These genera are: Diocalandra V. Bagoinae
Faust 1894; Dynatopechus Marshall 1931; Sternochetus Pierce 1917; VI. Baridinae
Liophloeus Germar 1817; Euophryum Broun 1909. 18. Baridini
Terminology. In general, standard terms for beetle anatomy 19. Madarini
are used in the keys and text (see Figs. 2, 3). Generally known and 20. Madopterini
readily visible characters are used where possible but in some 21. Nertinini
instances specialized characters requiring high magnification or VII. Ceutorhynchinae
dissections are required. Simply put, some weevil groups are dif- 22. Ceutorhynchini
ficult to identify. Measurements of body length are taken from 23. Cnemogonini
the anterior margin of the eyes to the apex of the elytra; the snout 24. Hypurini
is not included. On the elytra, intervals are numbered with the 25. Mononychini
sutural interval being interval 1. Tarsal articles are numbered from 26. Phytobiini
1 through 5, with 5 being the terminal or apical article bearing the 27. Scleropterini
claws; article 4 is very small and recessed between the lobes of VIII. Conoderinae
article 3. I use the term ventrite to apply to the visible abdominal 28. Lechriopini
sternites and they are numbered from 1 through 5, the latter 29. Zygopini
being terminal. 30. Tachygonini
In older literature the terms ‘uncus’ and ‘mucro’ are used to IX. Cossoninae
describe the structure of the apical tooth on the hind tibia. Fol- 31. Cossonini
lowing Thompson (1992), I have chosen not to use these terms 32. Acamptini
as comparative study shows them to refer to the same structure, 33. Dryotribini
 Family 131. Curculionidae · 725

34. Onycholipini 82. Petalochilini

35. Pentarthrini 83. Piazorhinini
36. Proecini 84. Pissodini
37. Rhyncolini 85. Sternechini
X. Cryptorhynchinae 86. Thalasselephantini
38. Cryptorhynchini 87. Trypetidini
39. Gasterocercini XVII. Scolytinae
XI. Cyclominae 88. Hylesinini
40. Rhythirrinini 89. Scolytini
XII. Entiminae XVIII. Platypodinae
41. Agraphini 90. Platypodini
42. Alophini
43. Anypotactini KEY TO THE NEARCTIC SUBFAMILIES OF CURCULIONIDAE
44. Brachyderini
45. Cneorhinini 1. Pregular sutures present; pregular sclerite distinct,
located between median gular suture and labial
46. Cyphicerini
articulation; head with rostrum virtually absent;
47. Eudiagogini at least one pair of tibiae with denticles or stout
48. Eustylini socketed setae along the dorsal (outer) margin
49. Geonemini ......................................................................... 2
— Pregular sutures absent; pregular sclerite not evi-
50. Hormorini
dent; head with rostrum variable from very long
51. Naupactini and cylindrical to short and broad, or (rarely)
52. Omiini nearly absent; tibiae lacking denticles or stout
53. Ophryastini socketed setae along the dorsal (outer) margin
......................................................................... 3
54. Otiorhynchini
55. Peritelini 2(1). Tarsus with article 1 as long as articles 2-5 com-
56. Phyllobiini bined; head as wide as pronotum; pronotum usu-
57. Polydrusini ally with lateral constriction near middle; anten-
nal club without sutures; lateral denticles on front
58. Sciaphilini
tibia not socketed .... XVIII. Platypodinae (p. 805)
59. Sitonini — Tarsus with article 1 not longer than articles 2 or 3;
60. Tanymecini head narrower than pronotum, often concealed
61. Thecesternini by pronotum when viewed dorsally; pronotum
not constricted laterally; antennal club with su-
62. Trachyphloeini
tures; lateral denticles on front tibia socketed or
63. Tropiphorini (rarely) not .................... XVII. Scolytinae (p. 792)
XIII. Hyperinae
64. Hyperini 3(1). Tarsus of 4 subequal articles (Fig. 12); eyes absent
(Fig. 11); body size small (<5mm); body color gen-
XIV. Lixinae
erally pale orange-red or pale brown; tibia at in-
65. Lixini ner apical angle with small tooth much shorter
66. Cleonini than a tarsal claw ................................................
67. Rhinocyllini .......................... III. Raymondionyminae (p. 732)
— Tarsus of 5 articles, but with article 4 very small
XV. Mesoptiliinae
and difficult to see between lobes of article 3
68. Laemosaccini (Fig. 88); eyes absent or present, well-developed,
69. Magdalidini or reduced in size and represented by only from
XVI. Molytinae 1 to a few facets (Fig. 51); body size various;
body color various; tibia at apex various but if
70. Molytini
eyes are lacking or almost so, tibia with large tooth
71. Trachodini arising from outer apical angle ......................... 4
72. Anchonini
73. Camarotini 4(3).1 Tarsus with claws widely separated by dermal lobes
extended between them from both dorsal and
74. Cholini
ventral surfaces at apex of article 5; mouthparts
75. Cleogonini with prementum withdrawn into oral cavity, palpi
76. Conotrachelini mostly or entirely concealed; antenna inserted
77. Cycloterini near base of rostrum, with scape long, projected
some distance beyond the hind margin of the
78. Erodiscini
eye and not fitting into antennal scrobe (Fig. 5)
79. Hylobiini (exceptions; Dryophthorus [Fig. 4], Orthognathus
80. Lepyrini [Fig. 7], Yuccaborus [Fig. 6] have a more distal
81. Lymantini insertion of the antennae, possess a scrobe and
726 · Family 131. Curculionidae

the scape does not pass, or only slightly passes, the apical face of the tibia; apical comb of setae
beyond hind margin of eye); antenna with club of present or absent, if present, oriented either trans-
two basic parts, with basal glabrous and glossy versely, obliquely or subparallel to the length of
portion, and apical uniformly pilose portion (Figs. the tibia ............................................................ 7
4-7); funicle with 4, 5 or 6 articles; body surface — Legs with apex of front, middle and hind tibiae with
lacking broad flat scales; pygydium formed of tooth, if present, small to moderately large (usu-
tergite 7 in male ........ I. Dryophthorinae (p. 728) ally larger on front or middle tibiae), usually smaller
— Tarsus with claws single, connate at base or sepa- than tarsal claw, arising from inner apical angle
rate, but with dorsal and ventral surfaces at apex and with outer curved face distinctly separated
of article 5 not extended between bases of tarsal from, and not continuous with, outer tibial margin
claws; mouthparts with prementum visible, not or with carina traversing the apical face of the
withdrawn, palpi mostly visible; antenna inserted tibia; apical comb of setae oriented transversely
variously along length of rostrum, usually some to length of tibia ............................................. 18
distance from base, with scape short or long, and
fitting into antennal scrobe, but at most only 7(6). Mesepimeron strongly ascended, truncated by
slightly projected beyond the hind margin of the elytral humeri and visible (or nearly so) in dorsal
eye (Figs. 8, 13, 27, 70); antenna with club vari- view between pronotum and elytra (Figs. 23-26);
ous, but mostly with three articles, each pilose tarsus with 1 (rarely) or 2 claws .........................
to some extent, basal article not or rarely glossy, ................................ VI. Baridinae (most) (p. 740)
subequal in length to other articles or rarely vari- — Mesepimeron not ascended, not visible in dorsal
ously longer than other 2 articles combined, su- view between pronotum and elytra (exception;
tures evident between all articles; funicle with 5, Laemosaccus [Fig. 92] recognized by short,
6 or 7 articles; body surface mostly with some straight rostrum, basal margin of elytra extended
broad flat scales or fine hair-like scales; pygydium over base of pronotum, exposed pygydium, and
formed of tergite 8 in male ............................... 5 small, acute tooth on the inner margin of the front
femur); tarsus with 2 claws ............................... 8
5(4). Male with aedeagus with tectum and pedon sepa-
rate (dissection necessary), tegmen as long as or 8(7). Rostrum in repose received into ventral channel
longer than aedeagus (including the apodemes); which may be limited to prosternum or extended
species associated with freshwater aquatic habi- beyond into meso- or metasternum (Figs. 21, 59)
tats, many with dense varnish-like coating over ......................................................................... 9
scales or with dense hydrofuge scales ............. — Rostrum in repose not received into ventral chan-
.......................................... II. Erirhininae (p. 730) nel, but may rest between front, middle and/or
— Male with aedeagus with tectum and pedon fused hind coxae ..................................................... 14
(dissection necessary), tegmen shorter than
aedeagus (including the apodemes); species as- 9(8). Eyes large, elongate-oval, subcontiguous (or nearly
sociated with various habitats, most with scales so) dorsally, frons very narrow (Fig. 45); eyes situ-
present, various in density, but lacking varnish- ated towards top front of head, in lateral view
like coating (exception; Bagous recognized by with lower margin of eye clearly situated above
presence of prosternal channel) or with scales level of dorsum of base of rostrum (Fig. 46) ......
lacking entirely ................................................ 6 ................................... VIII. Conoderinae (p. 754)
— Eyes small to moderate in size, more or less rounded,
6(5).2 Legs with well-developed, usually large hook-like more widely separated dorsally, frons broad; eyes
tooth at apex of front, middle and hind tibiae: situated towards sides of head, in lateral view
tooth arising from one of, a) outer apical angle with lower margin of eye situated near or below
(Fig. 57), b) from middle of apical margin (Fig. 93), level of dorsum of base of rostrum ................ 10
or c) at inner apical angle, but if at inner apical
angle, tooth on hind tibia is more or less as long 10(9). Rostrum very short, not much longer than wide,
as or longer than tarsal claw (Fig. 89) and outer broad and flat dorsally, subquadrate in form (Fig.
curved face of tooth is continuous with apex of 82); dorsal vestiture of pronotum and elytra in
outer tibial margin or is connected to it by a dis- part bifid (Bangasternus) ....................................
tinct, continuous sharp carina which traverses ................................... XIV. Lixinae (part) (p. 783)
1
— Rostrum moderately long, many times longer than
In small specimens it may be difficult to see the states of the tarsal wide, elongate and narrow; dorsal vestiture, if
claws and the mouthparts. There are only two genera of small-sized present, simple .............................................. 11
Dryophthorinae included here. Dryophthorus (Fig. 4) can be recognized
by an antennal funicle of 4 articles in combination with the antennal 11(10). Ventral channel extended beyond prosternum into
club character, whereas Sitophilus may be recognized by the form of meso- or metasternum (Fig. 59) ..........................
the apex of the hind tibia which has a small preapical tooth on the ............................. X. Cryptorhynchinae (p. 761)
inner margin in addition to the larger hook-like tooth at the inner — Ventral channel limited to prosternum (Fig. 21); even
though rostrum in repose may overlie meso-,
apical angle (the tibiae appearing “pincer-like”), in combination with
metasternum and some abdominal ventrites) ....
the antennal club. ....................................................................... 12
2
This is often a difficult character to see clearly and to assess. Some
groups (e.g., many Baridinae and some Curculioninae) are equivocal 12(11). Hind tibia with outer face at apex lacking apical
and are thus considered in both halves of this couplet. In general, comb of setae lateral to base of apical tooth (as in
taxa associated with woody plants tend to develop a larger and curved Fig. 57); body with distinct and dense suberect
apical tooth whereas those associated with herbaceous plants have a or erect broad scales, body of some specimens
less developed tooth or apical spine, or none at all.
 Family 131. Curculionidae · 727

with crustose coating (Acamptini, Acamptus) ... face with many fine scales and/or setae, man-
.............................. IX. Cossoninae (part) (p. 756) dibles generally robust and thick; rostrum short
— Hind tibia with outer face at apex with apical comb and broad, usually quadrate or subquadrate in
of setae lateral to base of apical tooth (Fig. 99); form, often expanded laterally towards apex, not
body vestiture various but surface not with crus- different in males and females in length or form
tose coating ................................................... 13 (Figs. 70-77) ............ XII. Entiminae (most) (p. 766)
— Mandible lacking scar and therefore lacking decidu-
13(12). Body lacking distinct vestiture, with smooth var- ous process, either glabrous or with a few small
nish-like coating over scales; elytra tuberculate setae on outer apical face, mandibles generally
or not; legs elongate, slender; commonly associ- less robust, smaller and thinner; rostrum more elon-
ated with aquatic habitats ... V. Bagoinae (p. 740) gate and cylindrical, usually as long as or longer
— Body with vestiture of appressed scales or suberect than pronotum, or (rarely) shorter than pronotum,
or erect hair-like scales, lacking smooth varnish- different in males and females in length and/or
like coating over scales, or obvious vestiture lack- form or not (Figs. 13, 17, 19) .......................... 19
ing; elytra tuberculate or not; legs more robust;
rarely associated with aquatic habitats ............. 19(18). Rostrum in repose received into distinct ventral
........................................ XVI. Molytinae (p. 786) channel in prosternum (rarely into mesosternum)
....................................................................... 20
14(8). Mouthparts with labial palpi of 3 articles but short, — Rostrum in repose not received into ventral chan-
globular, telescoping and appearing composed nel, but may rest between front, middle and/or
of 1 article, ventrally situated at apex of large hind coxae ..................................................... 23
prementum (Fig. 90); female with large paired sym-
biont sacs attached to vagina near base of 20(19). Rostrum very broad, more or less triangular in dor-
gonocoxites; body size mostly medium to large sal view, fitting into large, deep emargination in
(>5 mm) (exception; Microlarinus) ...................... front of front coxae; emargination limited poste-
............................................ XIV. Lixinae (p. 783) riorly by small, triangular prosternum (Fig. 69)
— Mouthparts with labial palpi of 3 distinct articles (Thecesternini, Thecesternus) ............................
but elongate, not telescoping, dorsally situated ................................ XII. Entiminae (part) (p. 766)
at apex of variously sized prementum; female lack- — Rostrum more elongate and cylindrical in form, the
ing large paired symbiont sacs attached to va- prosternal channel extended behind the front
gina near base of gonocoxites (dissection nec- coxae (rarely onto mesosternum) and the rostrum
essary); body size mostly small to medium (<10 (when in repose) extended between and/or be-
mm) ................................................................. 15 yond front coxae ........................................... 21

15(14). One or more of mesepisternum, mesepimeron, 21(20). Antenna with funicle with 5 articles; prothorax lack-
metepisternum and metepimeron with vestiture in ing postocular lobes; claws free, simple; dorsum
form of dense plumose (pectinate) hairs (Fig. 15), covered with fine, erect hair-like vestiture
rarely hairs may be sparse, fine and at most bifid (Mecinini, Cleopomiarus) ....................................
only in anterior portion of metepisternum ....... 16 .......................... IV. Curculioninae (part) (p. 732)
— Mesepisternum, mesepimeron, metepisternum and — Antenna with funicle with 6 or 7 articles; other char-
metepimeron with vestiture, if present, simple not acters various ................................................. 22
plumose or bifid ............................................. 17
22(21). Pygydium covered by elytra; rostrum longer than
16(15). Tooth at apex of tibia, large and hook-like, larger pronotum, straight and slender, abruptly attenu-
than tarsal claw (Fig. 93); pronotum only slightly ate immediately beyond antennal insertion (Fig.
narrower than base of elytra in dorsal view (Fig. 28); antenna with article 2 of funicle long, more
91); elytra with basal margin at intervals 2-4 ex- or less one-half length of scape (Madarini,
tended anteriorly overlapping base of pronotum Zygobaridina, Amercedes) .................................
(Fig. 91) ........................ XV. Mesoptilinae (p. 786) .................................. VI. Baridinae (part) (p. 740)
— Tooth at apex of tibia, small, at most subequal in — Pygydium not covered by elytra; rostrum various in
length to tarsal claw; pronotum distinctly nar- length, straight or slightly curved, more or less
rower than base of elytra in dorsal view (Fig. 14); of uniform width throughout length, not abruptly
elytra with basal margin at intervals 2-4 straight, attenuate (Fig. 34); antenna with article 2 of fu-
not overlapping base of pronotum (Fig. 14) nicle short, much less than one-half length of
(Otidocephalini) .................................................. scape .......... VII. Ceutorhynchinae (part) (p. 747)
.......................... IV. Curculioninae (part) (p. 732)
23(19). Mesepimeron strongly ascended, truncated by
17(15). Hind tibia with outer face at apex with apical comb elytral humeri and visible in dorsal view between
of setae lateral to base of apical tooth, oriented pronotum and elytra (Figs. 31-33); pygydium not
either transversely, obliquely or subparallel to covered by elytra (Figs. 31-33) ..........................
the length of the tibia (Figs. 99-101) .................. ................... VII. Ceutorhynchinae (part) (p. 747)
............................. XVI. Molytinae (most) (p. 786) — Mesepimeron not ascended, not visible in dorsal
— Hind tibia with outer face at apex lacking apical view between pronotum and elytra; pygydium
comb of setae lateral to base of apical tooth (Fig. mostly covered by elytra ............................... 24
57) ....................... IX. Cossoninae (most) (p. 756)
24(23). Tarsus with claws separate, each with basal pro-
18(6). Mandible with prominent scar on outer apical face c e s s ................... IV. Curculioninae (part) (p. 732)
indicating point of attachment of deciduous pro- — Tarsus with claws separate, simple ................... 25
cess (Fig. 68), or else clothed on outer apical
728 · Family 131. Curculionidae

25(24). Eyes rounded, rostrum mostly very elongate, slen- KEY TO THE NEARCTIC GENERA OF DRYOPHTHORINAE
der and cylindrical in cross section (Figs. 13, 15-
19); antenna with scape not or just reaching an-
1. Antenna with funicle of 4 articles (Fig. 4); tarsus
terior margin of eye (Figs. 13) ............................
with 5 distinct articles; body usually covered with
.......................... IV. Curculioninae (part) (p. 732)
a crusty deposit; size small, less than 4.0 mm in
— Eyes more or less elongate-oval, rostrum shorter,
body length ................................. Dryophthorus
more robust and subquadrate in cross section
— Antenna with funicle of 6 articles (Figs. 5-7); tarsus
(Figs. 64, 81); antenna with scape just reaching
with 5 articles but with article 4 small and diffi-
or passing anterior margin of eye (Fig. 64) .... 26
cult to see at base of article 5; body lacking sur-
face deposit; size various ............................... 2
26(25). Pronotum with anterolateral margin with distinct pos-
tocular lobe present (Fig. 64) .............................
2(1). Pygydium covered by apex of elytra; antenna with
...................................... XI. Cyclominae (p. 765)
scape not reaching anterior margin of eye (Figs.
— Pronotum with anterolateral margin straight, simple
6-7); metepimeron not visible .......................... 3
or postocular lobe at most very slightly devel-
— Pygydium exposed at apex of elytra; antenna with
oped (Fig. 81) .................................................. 27
scape projected at least past anterior margin of
eye (Fig. 5); metepimeron visible (obscure in
27(26). Vestiture with at least some bifid scales (limited on
Sitophilus) ......................................................... 4
some specimens to thoracic sterna), if bifid scales
appear absent, humeri obviously quadrate; hu-
3(2). Front coxae contiguous; hind tibia expanded
meri quadrate to subquadrate, rarely rounded, if
apically and with broad wide apical bevel;
humeri rounded, bifid scales are distinct on dor-
pronotum with postocular lobes; mandible large,
sum ................................. XIII. Hyperinae (p. 782)
lacking teeth on exterior face .... Orthognathus
— Vestiture simple, lacking bifid scales; humeri
— Front coxae separated by prosternum; hind tibia
rounded ................... XII. Entiminae (part) (p. 766)
linear, not expanded apically and with narrow api-
cal bevel; pronotum lacking postocular lobes;
mandible small, with 3 teeth on exterior face ...
CLASSIFICATION OF THE NEARCTIC CURCULIONIDAE ......................................................... Yuccaborus

4(2). Size small, total body length less than 5 mm; tibiae
I. Dryophthorinae Schoenherr 1825 (especially front) with distinct subapical tooth on
inner margin in addition to larger apical tooth ...
by Robert S. Anderson ............................................................ Sitophilus
— Size moderate to large, total body length greater
than 5 mm; tibia with at most a rounded subapical
This group of weevils is characterized by the form of the anten- swelling on inner margin in addition to larger api-
nal club with the basal article glabrous and glossy, the presence of cal tooth ........................................................... 5
what Zimmerman (1993) called ‘dermal lobes’ extended between
the tarsal claws from both dorsal and ventral surfaces of the apex
of tarsal article 5, the antenna (usually) with the scape long and
extended far beyond the posterior margin of the eye, and male 4 5
genitalia with a distinct lateral line dividing the aedeagus into
upper (tectum) and lower (pedon) parts. This primitive form of
genitalia is shared with Raymondionyminae and Erirhininae and
is the basis for some authors removing these three subfamilies
from Curculionidae and giving them each separate family status.
By removing these three groups, the hypothesis of monophyly
of Curculionidae is strengthened based on their unique derived
form of genitalia not shared with other Curculionoidea.
Dryophthorinae are a tropical group, and few species occur in
North America. Except for the diverse genus Sphenophorus, of the
North American genera each is represented by but one or a few 6 7
species. Most dryophthorines are associated with monocots, in-
cluding Poaceae, Cyperaceae, Liliaceae and Arecaceae. Some species
are serious pests of bananas, bromeliads, corn, turfgrass and
stored products. Larvae generally mine stems or roots, some in
semiaquatic habitats. The odd genus Dryophthorus is associated
with moist dead wood.

FIGURES 4.131-7.131. Dryophthorinae, lateral view of head. 4.

Dryophthorus americanus Bedel; 5. Sphenophorus zeae Walsh; 6. Yuccaborus
frontalis (LeConte); 7. Orthognathus subparallelus (Chevrolat).
 Family 131. Curculionidae · 729

5(4). Metepisternum very broad, length more or less 2 CLASSIFICATION OF THE NEARCTIC DRYOPHTHORINAE
times width; antenna with club transverse, wider
than long, lateral margins at base widely diver-
gent, shape sub-triangular; body size very large, 1. Dryophthorini Schoenherr 1825
total body length greater than 25 mm ...............
.................................................. Rhynchophorus Dryophthorus Germar 1824, 1 sp., D. americanus Bedel 1885, gener-
— Metepisternum narrow, length 3 or more times width;
ally distributed in eastern North America. Adults are found un-
antenna with club elongate, longer than wide,
lateral margins at base sub-parallel to slightly di- der bark, in association with old rotten logs or in forest litter.
vergent, shape sub-quadrate or sub-oval; body Bulbifer Dejean 1821
size moderate to large, total body length greater Dryophora Berthold 1827
than 5 mm but less than 25 mm ........................ 6
Tetratemnus Wollaston 1873
6(5). Scutellum (exposed portion) widest at or near Tetraspartus Pascoe 1885
middle, shape rhomboidal or sub-circular; more
or less as long as wide .................. Cosmopolites 2. Orthognathini Lacordaire 1866
— Scutellum (exposed portion) widest at or near base,
shape triangular or sub-triangular; generally
longer than wide .............................................. 7 Orthognathina Lacordaire 1866

7(6). Tarsus with article 3 with ventral pilosity long, con- Orthognathus Schoenherr 1838, 1 sp., O. subparallelus (Chevrolat
fined to apical margin as a continuous fringe,
1880), Arizona. Adults have been collected at lights.
ventral surface otherwise glabrous; antenna with
club obliquely truncate at apex with apical pi- Sphenognathus Schoenherr 1840
lose part very short, appearing recessed within
glabrous part, visible only as a narrow line in lat- Rhinostomina Kuschel 1995
eral view ....................................... Scyphophorus
— Tarsus with article 3 with ventral pilosity long or
short, uniformly covering 1/3 or more of ventral Yuccaborus LeConte 1876, 1 sp., Y. frontalis (LeConte 1876), gener-
surface, or with pilosity sparse and confined to ally distributed in southwestern United States. Two subspecies
anterolateral angle or lateral margins, ventral sur- are recognized. Adults and larvae are associated with Yucca
face otherwise glabrous; antenna with apex
(Liliaceae); adults come to lights.
evenly rounded or truncate, with apical pilose
part long, distinctly visible as more than a narrow
line in lateral view ............................................ 8 3. Rhynchophorini Schoenherr 1833
8(7). Tarsus with article 5 ventrally excavated and
Rhynchophorina Schoenherr 1833
bilamellate at middle of apex; rostrum hump-like
at base, directed posteroventrally; associated
with Asteraceae, Asclepiadaceae ...................... Rhynchophorus Herbst 1795, 2 spp., R. palmarum (Linnaeus 1758)
...................................................... Rhodobaenus and R. cruentatus (Fabricius 1775). Extreme southeastern United
— Tarsus with article 5 ventrally evenly rounded at
States, Texas and California. Adults and larvae are associated with
middle of apex; rostrum straight (few) or evenly
rounded at base (many), directed anteroventrally; various species of palms (Arecaceae). See Wattanapongsiri (1966)
associated with monocotyledons .................... 9 to separate the species. (Volume 1, Color Fig. 14)
Cordyle Thunberg 1797
9(8). Tarsus with article 3 with ventral pilosity restricted
to anterolateral areas, median area largely gla-
brous, article 3 narrow, subequal in width to ar- Litosomina Lacordaire 1866
ticle 2 (many) or broad, wider than article 2 (few)
..................................................... Sphenophorus Sitophilus Schoenherr 1838, 5 spp., generally distributed; adven-
— Tarsus with article 3 with ventral pilosity extensive
tive. Three species, S. granarius (Linnaeus 1758), S. zeamais
covering nearly all of ventral surface except near
base at middle, article 3 broad, wider than article Motschulsky 1855, and S. oryzae (Linnaeus 1763) are serious pests
2 ..................................................................... 10 of stored grain products. See Kuschel (1961) for a partial key to
species.
10(9). Front coxae widely separated by width of antennal
club; middle coxae widely separated by width of
a coxa; prementum toothed ventrally or slightly Sphenophorina Lacordaire 1866
emarginate at apex; Florida; on Arececeae,
Bromeliaceae ................................... Metamasius Cactophagus LeConte 1876, 1 sp., C. spinolae (Gyllenhal 1838), Ari-
— Front coxae narrowly separated by one-half width
zona and California, adults and larvae are associated with Carnegiea
of antennal club; middle coxae narrowly sepa-
rated by one-half width of a coxa; prementum gigantea (Engelm.) and other cacti (Cactaceae) (Anderson 1948).
broadly sulcate throughout length; Arizona, Cali- Cactophagus graphipterus (Champion 1910) has been found in or-
fornia; on Cactaceae .................... Cactophagus chid houses in Connecticut, Washington DC, and New Jersey
(Barber 1917). It is not known if this species is established there.
See Vaurie (1967) to separate the species.
730 · Family 131. Curculionidae

Eucactophagus Champion 1910 Raymondionyminae, they possess male genitalia that are primi-
Phyllerythrurus Chevrolat 1885 tive in structure with the aedeagus with separate tectum and pedon,
and the tegmen as long as or longer than the aedeagus. Most
Cosmopolites Chevrolat 1885, 1 sp., C. sordidus (Germar 1824), species are associated with aquatic or semi-aquatic habitats and
Florida, adventive. This species is associated with banana trees the members of the subtribe Stenopelmina possess a dense,
(Musa sapientum L.); larvae mine stem and corm (Woodruff 1969). varnish-like coating over the scales or have dense hydrofuge scales.
Many species are active swimmers.
Metamasius Horn 1873, 3 spp., M. hemipterus (Linnaeus 1758) and Most species mine the stems or other parts of aquatic mac-
M. callizona (Chevrolat 1883), adventive; M. mosieri Barber 1920, rophytes. Species in the genera Cyrtobagous, Neochetina and
native; Florida. Metamasius hemipterus is associated with palms, Neohydronomus have been introduced for biological control of
sugar cane, and bananas (Woodruff and Baranowski 1985), aquatic weeds, mainly in Florida. Grypus equiseti (Fabricius 1775)
whereas, M. callizona is a serious pest in Tillandsia (O'Brien and is associated with primitive horsetails of the genus Equisetum.
Thomas 1990, Frank and Thomas 2000, Larson and Frank 2000);
M. mosieri is also associated with bromeliads (Larson et al. 2001). KEY TO THE NEARCTIC GENERA OF ERIRHININAE
See Vaurie (1966) to separate the species.
Odontorhynchus Chevrolat 1880 1. Antenna with funicle of 6 articles ....................... 2
— Antenna with funicle of 7 articles ..................... 11
Odontorrhynchus Kirby 1881
Metmasiopsis Champion 1910 2(1). Tarsus with single claw ................... Brachybamus
Subphyllerythrurus Voss 1954 — Tarsus with two claws ......................................... 3

3(2). Antenna with club with basal article glabrous and

Rhodobaenus LeConte 1876, 2 spp., R. tredecimpunctatus (Illiger 1794)
glossy and almost as long as rest of club (Fig. 8);
and R. quinquepunctatus (Say 1824), generally distributed in United tarsus with article 3 not emarginate, usually not
States and southeastern Canada. Species are associated with vari- wider than article 2 .......................................... 4
ous Asteraceae and Asclepiadaceae; larvae are in stems (Vaurie — Antenna with club uniformly pubescent (Fig. 9); tar-
sus with article 3 various ................................. 6
1981). See Vaurie (1981) to separate the species.
Homalostylus Chevrolat 1885 4(3). Pronotum with anterolateral margin straight,
postocular lobe absent; tarsus with article 5 longer
Scyphophorus Schoenherr 1838, 2 spp., S. acupunctatus Gyllenhal than four other articles combined; dorsal vestiture
of only isolated appressed, rounded scales, no
1838 and S. yuccae Horn 1873, generally distributed in extreme
obvious varnish-like coating overlying scales ..
southern United States. Species are associated with Agave and ....................................................... Cyrtobagous
Yucca (Liliaceae); larvae mine the roots and stems. See Vaurie — Pronotum with anterolateral margin with well-devel-
(1971) to separate the species. oped postocular lobe; tarsus with article 5 shorter
than four other articles combined; dorsal vestiture
of dense appressed scales, with varnish-like coat-
Sphenophorus Schoenherr 1838, 65 spp., generally distributed. Spe- ing overlying scales ......................................... 5
cies are associated with various monocots including grasses
(Poaceae) and sedges (Cyperaceae) (Vaurie 1951). Some species are 5(4). Rostrum short, stout, nearly straight (Fig. 8); middle
tibia flattened, with outer margin evenly curved,
pests of turfgrass or corn. See Vaurie (1951) to separate the spe-
and with both inner and outer margins with nu-
cies. (Volume 2, Color Fig. 30) merous long, dense, fine hairs ..... Lissorhoptrus
Sitonobia Gistel 1856 — Rostrum slender, elongate, evenly curved; middle
Merothricus Chevrolat 1885 tibia not flattened, with outer margin more or less
Trichischius LeConte 1876
Nesorthognathus Voss 1943

Diocalandrina Zimmerman 1993

[Diocalandra Faust 1894, 3 spp., intercepted in quarantine; British

Columbia, Washington, California and Arizona. Not established
in North America.]

II. Erirhininae Schoenherr 1825

8 9 10
by Robert S. Anderson

This group of weevils is unfortunately very difficult to character- FIGURES 8.131-10.131. Erirhininae. 8-9. Lateral view of head. 8.
ize based solely on external characters. Like Dryophthorinae and Lissorhoptrus oryzophilus Kuschel; 9. Stenopelmus rufinasus Gyllenhal;
10. Notiodes setosus (LeConte), tarsus, dorsal view.
 Family 131. Curculionidae · 731

straight, and with both inner and outer margins species, G. equiseti (Fabricius 1775), is associated with Equisetum
with short, stout scales and at most a few scat-
(Equisetaceae) in wetlands. See Cawthra (1957) to separate the
tered, fine longer hairs .................. Neobagoidus
species.
6(3). Tarsus with article 3 very broad, apex of article 5 Aplopus Dejean 1821
not or very slightly projected beyond lobes of Grypidius Schoenherr 1826
article 3 (Fig. 10) ............................................... 7
— Tarsus with article 3 emarginate or bilobed, apex of
article 5 distinctly projected beyond lobes of ar- Notaris Germar 1817, 2 spp., N. puncticollis (LeConte 1876) and N.
ticle 3 by at least one-half length article 5 ...... 9 aethiops (Fabricius 1792), generally distributed in Canada and
northern United States. Notaris aethiops is associated with
7(6). Body size less than 1.5 mm; frons about half as
Sparganium ramosum Curt. (Sparganiaceae) in Europe and Typha
wide as rostrum in dorsal view at point of anten-
nal insertion; pronotum lacking postocular lobes (Typhaceae) in wetlands in North America (Anderson 1997). See
...................................................... Tanysphyrus Buchanan (1927) to separate the species.
— Body size distinctly greater than 1.5 mm; frons wider Pilumnus Dejean 1821
than rostrum in dorsal view at point of antennal
Erirhinus Schoenherr 1825
insertion; pronotum with postocular lobes present,
slightly to well developed ............................... 8 Erycus Tournier 1874

8(7). Tarsus with article 5 very slightly projected beyond Procas Stephens 1831, 1 sp., P. lecontei Bedel 1879, Michigan,
apices of lobes of article 3 .............. Neochetina
Ontario, Quebec and Yukon Territory.
— Tarsus with article 5 not projected beyond apices
of lobes of article 3 .............................. Notiodes Apachiscelus Desbrochers 1875
Notodermus Desbrochers 1875
9(6). Rostrum very short, subequal in length to scape Pseudypera Voss 1936
(Fig. 9); pronotum with anterolateral margin
straight, postocular lobe absent (Fig. 9) .............
....................................................... Stenopelmus Tournotaris Alonso-Zarazaga and Lyal 1999, 2 spp., generally dis-
— Rostrum more elongate, from 1.5 to 2.0 times length tributed in Canada, Alaska, and northern United States south
of scape; pronotum with anterolateral margin with into Nevada and California. At least one species, T. bimaculata
postocular lobe present, slightly to well devel-
(Fabricius 1787), is associated with Typha (Typhaceae) in wetlands
oped ............................................................... 10
(Anderson 1997). See Buchanan (1927) to separate some of the
10(9). Rostrum straight, robust; eyes large, narrowly sepa- species.
rated ventrally by less than the width of rostrum;
pronotum with anterolateral margin with postocu-
Stenopelmina LeConte 1876
lar lobe slightly developed ...... Neohydronomus
— Rostrum evenly curved, slender; eyes moderate,
separated ventrally by about the width of ros- Brachybamus Germar 1835, 1 sp., B. electus Germar 1835, generally
trum; pronotum with anterolateral margin with pos- distributed in eastern North America. Adults have been associ-
tocular lobe well developed .............. Onychylis
ated with Eleocharis (Cyperaceae) in wetlands.
11(1). Each tibiae with small spur(s) in addition to small
tooth at inner apical angle ............................. 12 Cyrtobagous Hustache 1929, 1 sp., C. salviniae Calder and Sands
— Tibiae all lacking spurs ...................................... 13 1985, Florida. This species has been introduced for biological
control of Salvinia molesta Mitchell (Salviniaceae) (O’Brien 1995).
12(11). Each tibia with 2 spurs ................................ Procas
— Front tibia with 1 spur, middle and hind tibiae each
with 2 spurs ............................................ Notaris Lissorhoptrus LeConte 1876, 6 spp., generally distributed. Species
are associated with wetlands; L. oryzophilus Kuschel 1952 is a pest
13(11). Antenna with funicle with fine pubescence; elytra
of cultivated rice; larvae feed externally on roots (Anderson 1993a).
with stria 10 not margined along last interval; body
densely covered with broad scales ....... Grypus See Kuschel (1952) to separate the species.
— Antenna with funicle with distinct setae; elytra with Lissocordylus Kuschel 1952
stria 10 finely margined along last interval; body
with fine setae or elongate-linear scales ...........
Neobagoidus O’Brien 1990, 1 sp., N. carlsoni O’Brien 1990, Florida.
......................................................... Tournotaris
This species is associated with Lachnanthes caroliniana (Lamarck)
CLASSIFICATION OF THE NEARCTIC ERIRHININAE Dandy (Haemodoraceae) in wetlands (O’Brien 1990).

4. Erirhinini Schoenherr 1825 Neochetina Hustache 1926, 2 spp., N. bruchi Hustache 1926 and N.
eichhorniae Warner 1970, Florida, Louisiana and Texas. These spe-
Erirhinina Schoenherr 1825 cies have been introduced for control of Eichhornia crassipes (Mart.)
Solms. (water hyacinth; Pontederiaceae) (O’Brien 1995). See
Grypus Germar 1917, 3 spp., generally distributed in Canada and O’Brien (1976) or DeLoach (1975) to separate the species.
northern United States, south in West to Colorado. At least one
732 · Family 131. Curculionidae

Neohydronomus Hustache 1926, 1 sp., N. affinis Hustache, Florida.

This species has been introduced for control of Pistia stratiotes L.
(water lettuce; Araceae) (O’Brien 1995). 12

Notiodes Schoenherr 1838, 12 spp., generally distributed. Associ- 11

ated with wetlands. At least three species of Notiodes have been
associated with Cyperaceae but Notiodes celatus (Burke 1961) is FIGURES 11.131-12.131. Raymondionyminae, Alaocybites californica
associated with the fern Marsilea mucronata A. Br. (Marsileaceae) Gilbert, 11. Lateral habitus; 12. Tarsus, dorsal view.
(Burke 1971). See Tanner (1943) and Burke (1961a, 1965) to sepa-
rate the species. KEY TO THE NEARCTIC GENERA OF RAYMONDIONYMINAE
Notiophilus Schoenherr 1835; not Duméril 1805
Endalus Laporte 1840 1. Front coxae not separated by prosternum;
prosternum lacking lateral ridges in front of coxae;
Notionomus Erichson 1842 abdomen with ventrite 4 separated from 5 by a
deep suture similar to suture between ventrites
Onychylis LeConte 1876, 6 spp., generally distributed in eastern 3 and 4; antenna with funicle with 7 articles .....
North America. Species are associated with Pontederia cordata L. ......................................................... Alaocybites
— Front coxae narrowly separated by prosternum;
(Pontederiaceae) and Nuphar luteum (L.) Sibhorn and Smith prosternum with lateral ridge in front of each coxa
(Nymphaeaceae) in wetlands (Burke 1961b, Anderson 1993a). slightly to well developed; abdomen with ventrite
See Burke (1961b) to separate the species. This genus is compos- 4 separated from 5 by a shallow suture; antenna
ite and is being subdivided by Charles O’Brien and Guillermo with funicle with 5 or 7 articles ....................... 2
Wibmer. 2(1). Antenna with funicle with 5 articles; hind tibia lin-
ear or triangular in form; prosternum with ridges
Stenopelmus Schoenherr 1835, 1 sp., S. rufinasus Gyllenhal 1836, in front of coxae low ......................... Gilbertiola
generally distributed in the United States and southern Canada. — Antenna with funicle with 7 articles; hind tibia mark-
edly expanded towards apex, subtriangular in
This species is associated with Azolla (Salviniaceae) in wetland form; prosternum with ridges in front of coxae
habitats (Scherf 1964). well developed ............................ Schizomicrus
Panscopus Schoenherr 1843; not Schoenherr 1842
Monius Schoenherr 1845 CLASSIFICATION OF THE NEARCTIC RAYMONDIONYMINAE
Degorsia Bedel 1902
5. Raymondionymini Reitter 1913
Tanysphyrina Gistel 1856
Alaocybites Gilbert 1956, 2 spp., California. Adults have been col-
Tanysphyrus Germar 1817, 2 spp., generally distributed in the east- lected in coniferous leaf litter. See Gilbert (1956) to separate the
ern United States and Canada west across the north to British species.
Columbia and south to Utah. Tanysphyrus lemnae (Fabricius 1792)
is a widespread Holarctic species associated with Lemna (duck- Gilbertiola Osella 1982, 2 spp., California and Oregon. Adults
weed; Lemnaceae) whereas T. ater Blatchley 1928 is associated have been collected in redwood leaf litter. See Gilbert (1956) to
with Ricciocarpus natans (L.) Corda (Bryophyta: Ricciaceae); larvae separate the species.
mine the leaves. Gilbertia Osella 1977; not Cossman 1889; not Jordan and
Tanysphyroides Egorov 1996 (valid subgenus) Eigenmann 1890; not Walsingham 1891

III. Raymondionyminae Reitter 1913 Schizomicrus Casey 1905, 1 sp., S. caecus (Casey 1892), California.
Adults have been collected in leaf litter.
by Robert S. Anderson Schizonotus Casey 1892; not Ratzeburg 1852; not Thorell
1888; not Reuter 1892
This is a small group of three genera of eyeless weevils found in
North America only in California and adjacent Oregon. They are IV. Curculioninae Latreille 1802
easily recognized by their eyeless condition (Fig. 11) but also by
the tarsi, which have only 4 articles (Fig. 12). Like Dryophthorinae By Robert S. Anderson
and Erirhininae they possess primitive male genitalia and have
recently been given family status by Thompson (1992) and Traditionally this subfamily has been restricted to members of
Alonso-Zarazaga and Lyal (1999). Adults are collected in various the genus Curculio and some close relatives but it is now a large
kinds of leaf litter. Nothing is known of larval biology. conglomerate of taxa of questionable relationships. Members
have a small or no tooth on the inner angle at the apex of the
hind tibia, eyes are rounded, the rostrum mostly elongate to very
 Family 131. Curculionidae · 733

elongate and cylindrical in cross section, and the antenna with the 8(7). Front femur with ventral margin simple, lacking tooth
......................................................................... 9
scape not or just reaching the anterior margin of the eye. They
— Front femur with ventral margin with slightly to well-
may be confused with Baridinae or Ceutorhynchinae but mem- developed tooth ............................................ 12
bers of these latter two subfamilies have the mesepimeron strongly
ascended, truncated by elytral humeri and visible in dorsal view 9(8). Pronotum with anterolateral margin with postocular
lobe present; hind femur with ventral margin with
between the pronotum and elytra. Sexual dimorphism in rostral
large broad tooth ........................... Pachytychius
form in Curculioninae is extreme in some taxa; generally, the — Pronotum with anterolateral margin straight,
female rostrum in longer and finer and the antennae are inserted postocular lobe absent; hind femur with ventral
more basally than in males. This dimorphism appears to be re- margin simple, lacking tooth .......................... 10
lated to oviposition and may be a key adaptation in explaining
10(9). Pronotum with distinct lateral margin defined by low
weevil diversity (Anderson 1995). carina, apically with carina slightly produced lat-
Curculionines tend to be associated with many herbaceous erally, denticulate or serrate ............ Elaeidobius
as well as some woody plants. Most have larvae that develop in — Pronotum with lateral margin rounded, not defined
by carina, no lateral protrusions, denticulations
reproductive structures such as fruits, seeds or flower buds; some
or serrations ................................................... 11
also mine stems. Many plant families serve as hosts and knowl-
edge of the host plant can facilitate identifications. Larvae of 11(10). Abdomen with suture between ventrites 2 and 3
Rhamphini are leaf miners. Most species in Cionini and Mecinini straight laterally; rostrum longer than pronotum;
antenna with funicle from article 2 to apex, long
are adventive. Anthonomini are the most diverse group, espe-
and slender, about as long as club .... Acalyptus
cially the genus Anthonomus. An excellent review of the natural — Abdomen with suture between ventrites 2 and 3
history of Anthonomini is by Burke (1976). angulate posteriorly at lateral margin; rostrum
shorter than pronotum; antenna with funicle from
article 2 to apex, very short and stout, shorter
KEY TO THE NEARCTIC GENERA OF CURCULIONINAE
than length of club ............................ Phyllotrox

1. Rostrum in repose received into distinct ventral 12(8). Body greater than 2.3 mm in length; tarsal claws
channel in prosternum; antennae with 5 funicle widely divergent, tooth on claw extended from
articles .......................................... Cleopomiarus underside of claw ............................ Dorytomus
— Rostrum in repose not received into ventral chan- — Body less than 2.3 mm in length; tarsal claws not
nel, but may rest between front, middle and hind widely divergent, tooth on claw extended from
coxae; antennae with 5-7 funicle articles ....... 2 inside face of claw ......................................... 13

2(1). Tarsus with claws connate at base ..................... 3 13(12). Elytra nearly glabrous except for group of white
— Tarsus with claws free at base, simple or with basal scales near middle of interval 4; scutellum with
process ............................................................ 7 dense white scales; middle coxae separated by
distance nearly equal to width of a coxa ..........
3(2). Antenna with funicle with 5 articles .................... 4 ............................................................. Ephelops
— Antenna with funicle with 6 or 7 articles ............. 6 — Elytra with more or less uniformly distributed scales
or vestiture; middle coxae separated by distance
4(3). Pygydium covered by elytral apices .......... Cionus distinctly less than width of a coxa ............... 14
— Pygydium exposed beyond elytral apices .......... 5
14(13). Body with sparse fine pubescence; hind tibia with
5(4). Body oval, length less than twice greatest width; apical tooth minute; body color light brown .....
pronotum with lateral margins markedly arcuate .......................................................... Dietzianus
from base to apex ............................ Gymnetron — Body with sparse to dense scales; hind tibia with
— Body elongate and cylindrical, length more than apical tooth about half as long as tarsal claw; body
twice greatest width; pronotum with lateral mar- color darker reddish brown to black .............. 15
gins more or less subparallel in basal half .........
.............................................................. Mecinus 15(14). Rostrum with lateral groove defined to anterior mar-
gin of eye, with at most a few scattered scales
6(3). Tarsus with article 5 shorter than articles 1 to 3 adjacent to eye; body elongate-oval; scales gen-
combined .......................................... Smicronyx erally of one color; apical third of elytra in lateral
— Tarsus with article 5 about as long as articles 1 to 3 view markedly rounded to apex; associated with
combined .................................. Promecotarsus Asteraceae ........................................ Epimechus
— Rostrum with lateral groove not defined immediately
7(2). Tarsus with claw simple, lacking basal process or anterior to eye, obliterated by dense scales adja-
tooth ................................................................. 8 cent to eye; body stout; scales ornate, of more
— Tarsus with claw with basal tooth or process3 .. 16 than one color; apical half of elytra in lateral view
sloped gradually to apex; associated with Solan-
aceae ............................................ Brachyogmus
3
Nanops has a minute tooth that is difficult to see at high magnification 16(7). Abdomen with suture between ventrites 2 and 3
and it may appear absent. It can be recognized by its small size (1.4- markedly extended posteriorly towards lateral
1.5 mm) and front femur lacking a ventral tooth. Species are associated margins, extended to or beyond suture between
with Hypericum (Hypericaceae). ventrites 3 and 4 (Fig. 16) .............................. 17
734 · Family 131. Curculionidae

13 15 16

14

19
17
18

FIGURES 13.131-19.131. Curculioninae. 13. Curculio monticola (Casey), head, lateral view; 14. Myrmex arizonicus (Schaeffer), dorsal habitus.
15-19. Lateral habitus, 15. Myrmex arizonicus (Schaeffer); 16. Tychius tectus LeConte; 17. Notolomus bicolor LeConte; 18. Tachyerges ephippiatus
(Say); 19. Anthonomus fulvus LeConte.

— Abdomen with suture between ventrites 2 and 3 2.0-3.0 mm; associated with Salicaceae ...........
more or less straight, if extended posteriorly, not ............................................................ Archarius
extended to suture between ventrites 3 and 4
(Fig. 19) ........................................................... 18 21(19). Rostrum longer than head and pronotum combined;
elytra black with sparse, recumbent vestiture;
17(16). Pygydium covered by elytra; antenna with funicle pronotum slightly constricted toward apex;
with 6 or 7 articles; associated with subfamily southern Rocky Mountain United States; associ-
Papilionoideae (Fabaceae) ..................... Tychius ated with Geraniaceae ................... Hypoleschus
— Pygydium exposed beyond elytra apex (especially — Rostrum shorter than head and pronotum combined;
so in male); antenna with funicle with 5 or 6 ar- elytra yellowish or light reddish brown, lacking
ticles; associated with subfamily Mimosoideae obvious vestiture; pronotum rather markedly con-
(Fabaceae) ................................................ Sibinia stricted toward apex; southeastern United States
west into Texas; associated with Arecaceae ....
18(16). Front coxae positioned much closer to posterior .......................................................... Notolomus
margin of prosternum than to anterior margin, dis-
tance to anterior margin greater than twice dis- 22(18). Pronotum longer than wide, distinctly constricted
tance to posterior margin (Fig. 17) ................. 19 at base such that width at midlength much greater
— Front coxae positioned near middle of prosternum, than at base (Fig. 14); black, or black and red,
coxae more or less equidistant from anterior and glossy and ant-like in form ............................. 23
posterior margins of prosternum (Figs. 15, 18-19) — Pronotum wider than long, base not distinctly con-
....................................................................... 22 stricted such that width at midlength is at most
slightly greater than at base; form various .... 25
19(18). Eye partly covered by anterior margin of pronotum
(Fig. 13); mandible prominent, slender, triangular 23(22). Head with supraocular sulcus present and angulate
in outline, inner face simple, not dentate; ros- dorsolaterally; front femur lacking tooth on ven-
trum very long and slender (Fig. 13) .............. 20 tral margin; extreme southern Florida ................
— Eye distant from anterior margin of pronotum (Fig. 17); ..................................................... Micromyrmex
mandible not prominent, inner face dentate; ros- — Head with supraocular sulcus lacking, no obvious
trum moderately long and slender (Fig. 17) ...... 21 sulcus or impression above eye; front femur with
tooth on ventral margin, tooth may be obsolete in
20(19). Antenna with club longer than wide; tarsus with some specimens; widespread ........................ 24
claw with distinct and long basal tooth; body with
more or less uniform vestiture of brown or grey 24(23). Elytra oval, humeri rounded, flight wings absent;
appressed scales; body size 4.2-13.0 mm; asso- eyes slightly reduced in size and number of fac-
ciated with Fagaceae, Juglandaceae and ets .................................................... Oopterinus
Betulaceae ............................................. Curculio — Elytra elongate-oval, humeri quadrate (Fig. 14), flight
— Antenna with club as wide as long; tarsus with claw wings present; eyes well-developed (Fig. 15) ...
with short and fine basal tooth; body with vestiture .............................................................. Myrmex
of scattered white appressed scales; body size
 Family 131. Curculionidae · 735

25(22). Hind tibia with apical comb of setae oblique, set at — Front femur lacking tooth or with at most a short
an angle to long axis of tibia; hind tibia narrowed tooth, not longer than tarsal claw .................. 33
apically; hind femur stouter than middle femur,
slightly so in some specimens; jumping forms (Fig. 33(32). Tarsus with claw with short, broad, blunt basal pro-
18) .................................................................. 26 c e s s ................................................................ 34
— Hind tibia with apical comb of setae transverse, per- — Tarsus with claw with long, fine, acute basal tooth
pendicular to long axis of tibia; hind tibia not nar- ....................................................................... 35
rowed apically; hind femur not distinctly stouter
than middle femur .......................................... 28 34(33). Front femur simple, lacking tooth; rostrum shorter
than pronotum ........................................ Ellescus
26(25). Antenna with funicle of 7 articles; eyes — Front femur with minute tooth on ventral margin;
subcontiguous to contiguous in anterior view; rostrum longer than pronotum ........... Proctorus
elytra with or without distinct pattern of contrast-
ing pale vestiture ............................. Tachyerges 35(33). Rostrum with scrobe descended, antenna with
— Antenna with funicle of 6 articles; eyes distinctly scape rested below lateral rostral groove and
separated at point of closest approach by a dis- below ventral margin of rostrum; antenna with
tance greater than 0.10 X width of an eye in ante- funicle with 6 or 7 articles; associated with
rior view; elytra without distinct pattern of con- Rubiaceae ............................................. Plocetes
trasting pale vestiture .................................... 27 — Rostrum with scrobe not descended, rostrum with-
out lateral grooves, antenna with scape parallel
27(26). Metasternum (lateral portion), mesepisternum and to long axis of rostrum; antenna with funicle with
metepisternum with short, dense, plumose white 7 articles; associated with Oleaceae .................
scales which contrast markedly with the rest of ........................................................... Lignyodes
body vestiture; hind femur slightly expanded,
length greater than 3.10 X maximum width, ven- 36(31). Front femur with ventral margin simple, lacking
tral margin simple; body size small, 1.0-1.8 mm . tooth; tooth on tarsal claw minute (may appear
............................................................. Isochnus absent); body size small, 1.4-1.5 mm ..... Nanops
— Metasternum, mesepisternum and metepisternum — Front femur with ventral margin with tooth; tooth on
with vestiture as on rest of body, mesepisternum tarsal claw distinct; body size various, most
and metepisternum in some speicmens with broad greater than 1.5 mm ....................................... 37
bifurcate (but not plumose) scales; hind femur
slightly to markedly expanded, length less than 37(36). Front femur with large, broad, triangular tooth,
3.20 X maximum width, ventral margin with vari- middle and hind femora simple, lacking tooth; as-
ous spines and setae set in denticles; body size sociated with Malvaceae ............. Macrorhoptus
moderate, 1.6-2.5 mm ........................ Orchestes — Front, middle and hind femora each with tooth; as-
sociated with various plants (including
28(25). Front coxae distinctly separated by process of Malvaceae) ..................................................... 38
prosternum; middle coxae widely separated by
distance nearly equal to width of a coxa; body 38(37). Antenna with funicle with 6 articles; antenna with
size 1.1-1.4 mm; extreme southern Florida ........ club with basal article glossy, almost glabrous,
................................................................. Huaca remaining articles densely pubescent; dorsal
— Front coxae contiguous; middle coxae separated margin of eye elevated above level of interocular
by distance less than width of a coxa; body size area ........................................... Anthonomopsis
greater than 1.3 mm; widespread ................... 29 — Antenna with funicle with 7 articles; antenna with
club various; eyes various ............................. 39
29(28). Hind tibia with distinct apical tooth, tooth large and
curved, subequal in size to tarsal claw ......... 30 39(38). Elytra with surface even, not tuberculate; pygydium
— Hind tibia with at most only small, usually straight exposed beyond apices of elytra; scales of elytra
apical tooth, tooth much smaller than tarsal claw, evenly distributed, without contrasting pattern;
or tibial apex simple, lacking tooth ................ 40 body stout ...................................... Chelonychus
— Elytra with serrate tubercle at base of interval 3;
30(29). Antenna with funicle with coarse, elongate, erect pygydium covered by elytra; elytra with broad,
scales; antenna with club compact, glossy and conspicuous band of white scales across elytra
nearly glabrous; tarsus with claws usually with a near base; body more elongate ...... Smicraulax
long, slender tooth extended on inside of claw
well distad of base ......................... Magdalinops 40(29). Rostrum with lateral groove short, apex of groove
— Antenna with funicle with very fine setae; antenna not extended to anterior margin of eye (short by
with club various, usually less compact and with distance at least equal to diameter of eye); asso-
distinct pubescence; tarsus with claws various ciated with Viscaceae (mistletoe) ... Cionomimus
....................................................................... 31 — Rostrum with lateral groove long, apex of groove
extended to anterior margin of eye (if short, by
31(30). Abdomen with sutures between ventrites angled much less than diameter of eye); associated with
posteriorly at lateral margins .......................... 32 various plants ................................................. 41
— Abdomen with sutures between ventrites straight,
not angled posteriorly at lateral margins ....... 36 41(40). Antenna with funicle of 5 articles; antenna with club
with basal article glossy, almost glabrous; front
32(31). Front femur with large, broad, triangular tooth, tooth coxae of some slightly separated; middle coxae
longer than tarsal claw ................... Ochyromera widely separated; femora simple, lacking tooth
736 · Family 131. Curculionidae

on ventral margin; body size 1.3-1.5 mm .......... CLASSIFICATION OF THE NEARCTIC CURCULIONINAE
......................................................... Neomastix
— Antenna with funicle of 6 or 7 articles; antenna with
club various; front coxae various; middle coxae 6. Curculionini Latreille 1802
various; femora with tooth or simple, lacking tooth Curculionina Latreille 1802
on ventral margin; body size greater than 1.3 mm,
most greater than 1.5 mm ............................... 42
Archarius Gistel 1856, 1 sp., A. salicivorus (Paykull 1792), Quebec;
42(41). Rostrum with lateral groove descended, directed adventive. This species is associated with galls on Salix (Salicaceae).
slightly to well below middle of eye; elytra with Recently confirmed as established in Quebec by Sylvain Côté (pers.
base of interval 3 elevated; antenna with funicle comm.).
of 6 articles ........................... Pseudanthonomus
Archarias Villa and Villa 1833; not Dejean 1821
— Rostrum with lateral groove not descended, di-
rected to middle of eye; elytra with base of inter- Balanobius Jekel 1861
val 3 various, flat to elevated; antenna with fu- Longifistulia Hong and Wang 1987
nicle of 6 or 7 articles .................................... 43 Toptaria Kwon and Lee 1990 (valid subgenus)
43(42). Rostrum with dense scales throughout almost en-
tire length, scales obscuring underlying cuticle; Curculio Linnaeus 1758, 27 spp., generally distributed. Species are
head constricted behind eyes, hind margin of associated with various Fagaceae, Juglandaceae and Betulaceae.
eye markedly produced; associated with See Gibson (1969) to separate the species.
Bernardia (Euphorbiaceae); Texas .... Narberdia
Balaninus Germar 1817
— Rostrum with scales if present, not dense and lim-
ited to basal half of length, scales not obscuring Pelecinus Wiedemann 1823; not Latreille 1800
underlying cuticle; head not constricted behind Tropibalaninus Heller 1927 (valid subgenus)
eyes, eye produced but hind margin flat against Carponinophilus Voss 1962 (valid subgenus)
cuticle; associated with various plants; widely
distributed ...................................................... 44
7. Acalyptini Thomson 1859
44(43). Front femur markedly expanded, width about twice
that of middle or hind femur, with large biserrate Acalyptus Schoenherr 1833, 1 sp., A. carpini (Herbst 1795), gener-
tooth on ventral margin; head subconical; asso-
ally distributed in Alaska, Canada and northern United States.
ciated with Serjania (Sapindaceae); southern
Texas .................................................. Cionopsis This species is associated with Salix (Salicaceae) (Anderson 1997).
— Front femur at most slightly expanded, width less Orsophagus Roelofs 1874
than twice that of middle or hind femur, tooth on
ventral margin various; head subconical or not, if
8. Anthonomini Thomson 1859
subconical, then front femur not expanded; as-
sociated with various plants; widely distributed
....................................................................... 45 Anthonomopsis Dietz 1891, 1 sp., A. mixta (LeConte 1876), gener-
ally distributed in eastern and central United States and Canada.
45(44). Ventrite 5 of male very short at middle, deeply and
This species is associated with Prunus (Rosaceae) (Ahmad and
broadly emarginate; pronotum with low median
carina in basal third to one-half; rostrum short, Burke 1972).
subequal in length to pronotum or slightly longer,
and straight; abdomen with ventrites flat .......... Anthonomus Germar 1817, 110 spp., generally distributed. Species
.......................................................... Coccotorus
are associated with various families of plants including Asteraceae,
— Ventrite 5 of male longer at middle, at most slightly
and shallowly emarginate; pronotum lacking me- Caprifoliaceae, Cistaceae, Cupressaceae, Euphorbiaceae, Fabaceae,
dian carina; rostrum moderate to long, slightly to Juglandaceae, Krameriaceae, Malpighiaceae, Malvaceae, Myrtaceae,
distinctly longer than pronotum, and slightly to Rosaceae, Rutaceae, Salicaceae, Solanaceae, and Vitaceae; larvae
markedly curved; abdomen with ventrites con-
mostly develop in reproductive structures or in galls (Burke 1976).
v e x ................................................................. 46
See Dietz (1891), Hatch (1971), Blatchley and Leng (1916) to
46(45). Front tibia moderately curved, with apical half of separate some of the species. The genus presently is being revised
inner margin expanded and carinate; elytra with in the New World by Wayne Clark and Horace Burke; some of
interval 2 descended lateral to scutellum, inter-
their papers include North American species (Clark 1987a, b, 1988,
val 3 with prominent swelling at base; mesoster-
num markedly declivious; body size 4.0 - 5.8 mm; 1990, 1991a, b; Clark and Burke 1985, 1986, 1996).
southern Florida ........................... Atractomerus Pallene Dejean 1821
— Front tibia only slightly curved, with apical half of Furcipus Desbrochers 1868 (valid subgenus)
inner margin simple; elytra with interval 2 flat lat-
Toplithus Gozis 1882
eral to scutellum, interval 3 with slight to promi-
nent swelling at base; mesosternum at most Anthomorphus Weise 1883 (valid subgenus)
slightly declivious; body size various, most less Furcipes Bedel 1884
than 4.0 mm; widely distributed ..... Anthonomus Toplethus Bedel 1884
Anthonomochaeta Dietz 1891 (valid subgenus)
Anthonomocyllus Dietz 1891 (valid subgenus)
 Family 131. Curculionidae · 737

Anthonomorphus Dietz 1891 (valid subgenus) Huaca Clark 1993, 2 spp., southern Florida. Huaca apian Clark
Cnemocyllus Dietz 1891 (valid subgenus) 1993 has been associated with Zanthoxylum flavum Vahl. (Rutaceae)
Paranthonomus Dietz 1891 (as Anthonomini new genus 1, new species 1; Anderson 1993a).
Tachypterus Dietz 1891; not Guérin-Méneville 1838 See Clark (1993a) to separate the species.
Trichobaropsis Dietz 1891
Listrorrhynchus Champion 1903 Magdalinops Dietz 1891, 4 spp., generally distributed in western
Tachypterellus Fall and Cockerell 1907 (valid subgenus) United States and Canada. Species are associated with Asteraceae.
Anthonomidius Reitter 1915 (valid subgenus) See Clark and Burke (in press b) to separate the species.
Sexarthrus Blatchley 1916
Pterochalybs Ter-Minasian 1936 (valid subgenus) Nanops Dietz 1891, 1 sp., N. schwarzii Dietz 1891, southeastern
Persexarthrus Voss 1944 (valid subgenus) United States. This species is associated with Hypericum
Parafurcipes Voss 1956 (valid subgenus) (Hypericaceae).
Exanthonomus Voss 1960
Neobradybatus Hoffmann 1963 Narberdia Burke 1976, 1 sp., N. aridulus Burke 1976, Texas. This
species is associated with Bernardia myricaefolia (Scheele) Wats.
Atractomerus Duponchel and Chevrolat 1842, 1 sp., A. punctipennis (Euphorbiaceae); larvae in fruits (Burke and Rector 1976).
(Gyllenhal 1836), southern Florida. This species is associated with
Eugenia (Myrtaceae) (Anderson 1993a). Neomastix Dietz 1891, 1 sp., N. solidaginis Dietz 1891, southeast-
Leptarthrus Dietz 1891; not Stephens 1829 ern United States. Adults have been associated with various plants
Cissoanthonomus Hustache 1939 (Clark 1993b).
Arthleptrus Burke 1982
Pseudanthonomus Dietz 1891, 7 spp., generally distributed in eastern
Brachyogmus Linell 1897, 1 sp., B. ornatus Linell 1897, southwest- and central United States and Canada extending as far west as Ari-
ern United States. This species is associated with Lycium (Solan- zona and Colorado, and as far north as Yukon Territory. Species are
aceae) (Burke 1968). associated with various Rosaceae, Ericaceae, Betulaceae, Saxifragaceae,
Hamamelidaceae and Krameraceae; larvae in flower buds and fruits
Chelonychus Dietz 1891, 2 spp., generally distributed in Western (Clark 1987c). See Clark (1987c) to separate the species.
United States and Canada. See Clark and Burke (in press b) to
separate the species. Smicraulax Pierce 1908, 2 spp., Arizona and Texas. Species are
associated with Phoradendron (mistletoe; Viscaceae); larvae mine
Cionomimus Marshall 1939, 2 spp., southwestern and western stems. See Burke (1975) to separate the species.
United States. Species are associated with Phoradendron (mistletoe;
Viscaceae) (Burke 1981). See Burke (1981) or Anderson (1994) to 9. Cionini Schoenherr 1825
separate the species.
Cionistes Dietz 1891; not Wright 1861 Cionus Clairville 1798, 1 sp., C. scrophulariae (Linnaeus 1758), New
York; adventive. This species is associated with Scrophularia and
Cionopsis Champion 1903, 2 spp., southern Texas. Species are Verbascum (Scrophulariaceae); larvae feed externally on the leaves
associated with Serjania (Sapindaceae); larvae in fruits (Anderson and pupate in round translucent cocoons among flowers and
and Burke 1990). See Burke (1982) to separate the species. seed-capsules. Recently confirmed as established by Hoebeke (pers.
comm.).
Coccotorus LeConte 1876, 4 spp., generally distributed in eastern and
central United States and Canada. Species are associated with Prunus 10. Derelomini Lacordaire 1866
(Rosaceae) (Brown 1966a). See Brown (1966a) to separate the species.
Elaeidobius Kuschel 1952, 1 sp., E. subvittatus (Faust 1898), Florida;
Dietzianus Sleeper 1953, 2 spp., generally distributed in eastern United adventive. This species is associated with the male flowers of
States. See Blatchley and Leng (1916) to separate the species. Elaeis guineensis Jacquin (African oil palm; Arecaceae) (O’Brien
Xanthus Dietz 1891; not Gistl 1834; not Agassiz 1843 and Woodruff 1986).

Ephelops Dietz 1891, 1 sp., E. triguttatus Dietz 1891, southern Hypoleschus Fall 1907, 1 sp., H. atratus Fall 1907, New Mexico and
Florida. This species may be associated with Piscidia (Fabaceae) Colorado. This species is associated with Geranium sp. (cranesbill;
(Anderson 1993a). Geraniaceae) (C.W. O’Brien, pers. comm.).

Epimechus Dietz 1891, 11 spp., generally distributed in western Notolomus LeConte 1876, 2 spp., southeastern United States west
United States. Species are associated with various Asteraceae. See to southern Texas. Species are associated with flowers of Serenoa
Clark and Burke (in press a) to separate the species. repens (Bartr.) Small and Sabal palmetto (Walt.) Lodd (saw pal-
738 · Family 131. Curculionidae

metto and cabbage palm; Arecaceae); larvae develop in male flow- Gymnetron Schoenherr 1825, 4 spp., generally distributed; adven-
ers (Anderson 1993a). See Blatchley and Leng (1916) to separate tive. Species are associated with Verbascum thapsis Linnaeus, Linaria
the species. vulgaris Miller (both Scrophulariaceae) and Plantago lanceolata
Linnaeus (Plantaginaceae); larvae in seed capsules (Anderson 1973).
Phyllotrox Schoenherr 1843, 7 spp., generally distributed in the See Buchanan (1937) to separate three of the four species; Sleeper
United States. Phyllotrox canyonacerensis Warner 1976 is associated (1954a) presents notes on the fourth. Downie and Arnett (1996)
with fruits of Acer grandidentatum Nutt. (maple; Aceraceae) (Warner provide a brief key to the four species.
1976). The genus needs revision. Gymnetrum Agassiz 1846
Euclyptus Dietz 1891 Carpolinus Gistel 1848
Aprinus Desbrochers 1893
11. Ellescini Thomson 1859 Eutemnoscelus Desbrochers 1893 (valid subgenus)

Ellescina Thomson 1859 Mecinus Germar 1821, 2 spp., M. pyraster (Herbst 1795) and M.
janthinus (Germar 1817), eastern and western United States and
Ellescus Dejean 1821, 4 spp., generally distributed. Species are Canada (disjunct); adventive. Mecinus pyraster is associated with
associated with Salix and Populus (willow, poplar and aspen; Plantago lanceolata Linnaeus (Plantaginaceae); larvae are in seed
Salicaceae); larvae mine the central axis of female catkins (Scherf capsules (Anderson 1973). Mecinus janthinus has been introduced
1964). The genus needs revision. into Montana, Wyoming, Washington, British Columbia, Alberta
Sarapus Villa and Villa 1833; not Fischer von Waldheim 1821 and Nova Scotia (Harris et al. 2001; DeClerk-Floate and Harris in
Elleschus Schoenherr 1838 press) for the biological control of Linaria vulgaris Miller (yellow
Alyca LeConte 1876 toad-flax) and L. dalmatica (L.) Miller (Dalmation toad-flax)
Anisarctus Desbrochers 1907 (Scrophularaceae). There is no key to separate the two species in
North America.
Proctorus LeConte 1876, 2 spp., generally distributed in northern Hexaphyllus Dejean 1821
United States, Canada and Alaska. Associated with Salix (willow; Macipus Fischer de Waldheim 1829
Salicaceae). See LeConte and Horn (1876) to separate the species. Mecinopsis Escalera 1914
Encalus LeConte 1876
13. Otidocephalini Lacordaire 1863
Dorytomina Bedel 1886
Micromyrmex Sleeper 1953, 2 spp., M. cavirostris (Casey 1892) and
Dorytomus Germar 1817, 21 spp., generally distributed. Species M. poeyi (Chevrolat 1832), southern Florida. See Blatchley and
are associated with Salix and Populus (willow, poplar and aspen; Leng (1916; as Otidocephalus) to separate the species.
Salicaceae); larvae feed in catkins and one develops in sawfly galls
in the stems of Salix. See O’Brien (1970a) to separate the species Myrmex Sturm 1826, 31 spp., generally distributed in the United
but note subsequent synonymy as summarized in O’Brien and States and southeastern Canada; most species in southwestern
Wibmer (1982). United States. Species are associated mainly with various Asteraceae,
Solenorhinus Motschulsky 1860 also Fagaceae, Ulmaceae, Arecaceae, Smilacaceae, Viscaceae and
Doratotomus Gistel 1886 Sapotaceae (Anderson 1993b). Larvae mostly mine stems. The
Eteophilus Bedel 1886 genus needs revision. See Horn (1873) and Schaeffer (1907) to
Alycodes Dietz 1891 separate most of the species.
Euolamus Reitter 1916 (valid subgenus) Otidocephalus Chevrolat 1832
Olamus Reitter 1916 (valid subgenus) Cycotida Pascoe 1872
Praeolamus Zumpt 1932
Paradorytomus Zumpt 1932 Oopterinus Casey 1892, 2 spp., eastern United States. Larvae of O.
Chaetodorytomus Iablokov-Khnzorian 1970 (valid subgenus) perforatus develop in cynipid galls on the roots of Quercus. See
O’Brien (1985) to separate the species.
12. Mecinini Gistel 1856
14. Rhamphini Rafinesque 1815
Cleopomiarus Pierce 1919, 1 sp., C. hispidulus (LeConte 1876), gen-
erally distributed in eastern United States. This species is associ- Rhamphina Rafinesque 1815
ated with Lobelia (Campanulaceae); larvae in seed capsules (Ander-
son 1973). Isochnus Thomson 1859, 5 spp., generally distributed in North
Miaromimus Solari 1947 America, including far northern Canada and Alaska; not in south-
Hemimiarus Franz 1947 eastern United States. Species are associated with Salix and Populus
 Family 131. Curculionidae · 739

(willow, poplar and aspen; Salicaceae); larvae mine leaves (Ander- family Mimosoideae); larvae in reproductive structures (Clark
son 1989a). See Anderson (1989a) to separate the species. 1978). See Clark (1978) to separate the species.
Sibynes Schoenherr 1825
Orchestes Illiger 1798, 5 spp., generally distributed. Species are Campipterus Motschulsky 1845
associated with Betulaceae, Rosaceae and Ulmaceae; larvae mine Campopterus Agassiz 1846
leaves (Anderson 1989a). See Anderson (1989a) to separate the Sibynia Agassiz 1846
species. Aocnus Schoenherr 1859
Salius Schrank 1798 (valid subgenus) Sibynia Wollaston 1865; not Agassiz 1846
Alyctus Thomson 1859 Paragoges LeConte 1876
Threcticus Thomson 1859 Dichotychius Bedel 1885 (valid subgenus)
Euthoron Thomson 1859 Mecynopyga Pierce 1908
Nomizo Morimoto 1984 (valid subgenus) Microtychius Casey 1910 (valid subgenus)
Teratonychus Bondar 1949
Tachyerges Schoenherr 1825, 3 spp., generally distributed. Species Itychus Kissinger 1962
are associated with Salix and Populus (willow, poplar and aspen;
Salicaceae); larvae mine leaves (Anderson 1989a). See Anderson Tychius Germar 1817, 16 spp., generally distributed; four species
(1989a) to separate the species. adventive (Anderson and Howden 1994). Species are associated
with various native and adventive Fabaceae (subfamily
15. Smicronychini Seidlitz 1891 Papilionoideae); larvae in reproductive structures (Clark 1971; Clark
and Burke 1977). See Clark (1971, 1977) and Anderson and
Promecotarsus Casey 1892, 3 spp., generally distributed in western Howden (1994) to separate the species.
United States and Canada. See Casey (1892) to separate the spe- Miccotrogus Schoenherr 1825
cies. Apeltarius Desbrochers 1873 (valid subgenus)
Ectatotychius Tournier 1874
Smicronyx Schoenherr 1843, 70 spp., generally distributed. Species Hypactus Marseul 1888
are associated with various plants, mostly Asteraceae and Henonia Pic 1897
Convolvulaceae (Cuscuta; dodder); larvae are in seeds or may cause Xenotychius Reitter 1897
galls (Anderson 1962). See Anderson (1962) to separate the spe- Pseudolignyodes Pic 1899
cies. Paratychius Casey 1910
Micronyx Schoenherr 1835; not Boisduval 1835 Aoromius Desbrochers 1907
Desmoris LeConte 1876 (valid subgenus) Lepidotychius Penecke 1922
Pachyphanes Dietz 1894 (valid subgenus) Elleschidius Penecke 1938
Pseudromicronyx Dietz 1894 (valid subgenus) Heliotychius Franz 1943
Synertha Dietz 1894 Neotychius Hustache 1945
Chalybodontus Desbrochers 1897 (valid subgenus) Mongolotychius Korotyaev 1990
Oligocaricis Lea 1926
Lignyodina Bedel 1884
16. Storeini Lacordaire 1863
Lignyodes Dejean 1835, 17 spp., generally distributed. Species are
Pachytychius Jekel 1861, 1 sp., P. haematocephalus (Gyllenhal 1836), associated with Oleaceae; larvae are in reproductive structures (Clark
New York; adventive. This species is associated in Europe with 1980a, 1980b, 1981). Subgenus Lignyodes are associated with
Lotus corniculatus L. (Fabaceae) (Hoffmann 1958). Fraxinus (ash), subgenus Chionanthobius with Chionanthus,
Styphlotychius Jekel 1861 Forestiera and Osmanthus, and subgenus Neotylopterus with Forestiera.
Barytychius Jekel 1861 See Clark (1980a, 1980b, 1981) to separate the species.
Scyphotychius Desbrochers 1875 Lignyodes Schoenherr 1835; not Dejean 1835
Rabdotorhinus Desbrochers 1894 Stenorhynchus Villa and Villa 1835; not Lamarck 1818; not
Fogatianus Caldara 1978 Hemprich 1820; not Berthold 1827
Rhaestes Gistel 1856
17. Tychiini Thomson 1859 Thysanocnemis LeConte 1876
Tylopterus LeConte 1876; not Capiomont 1868
Tychiina Thomson 1859 Chionanthobius Pierce 1912 (valid subgenus)
Lignyodius Dieckmann 1970
Sibinia Germar 1817, 22 spp., generally distributed in western Neotylopterus Clark, Whitehead and Warner 1977 (valid sub-
United States. Species are associated with various Fabaceae (sub- genus)
740 · Family 131. Curculionidae

subtribe do not possess a sternal channel for reception of the

rostrum and have different male genitalia.

KEY TO THE NEARCTIC GENERA OF BAGOINAE

1. Pronotum slightly constricted behind apex (Fig. 20)

................................................................ Bagous
20 — Pronotum markedly constricted behind apex .......
............................................................. Pnigodes

21
CLASSIFICATION OF THE NEARCTIC BAGOINAE
FIGURES 20.131-21.131. Bagoinae, Bagous americanus LeConte, 20.
Lateral habitus; 21. Thoracic sterna, ventral view. Bagous Germar 1817, 33 spp., generally distributed. Species are
associated with various wetland plants such as Limnobium spongia
Plocetes LeConte 1876, 4 spp., generally distributed in southeast- (Bosc) Steud. (Hydrocharitaceae), Brasenia schreberi Gmel. and
ern United States west to southern Texas (two species are re- Nymphaea (Nymphaeaceae), Eleocharis and Carex (Cyperaceae), and
stricted to extreme southern Florida; one to extreme southern Potamogeton (Potamogetonaceae) (O’Brien and Marshall 1979).
Texas). Plocetes ulmi LeConte 1876 is widespread in the southeast- Bagous pictus Blatchley 1920 is associated with Sesuvium
ern United States and is associated with Cephalanthus occidentalis portulacastrum (L.) L. (Aizoaceae). See Tanner (1943) to separate
L. Species are all associated with Rubiaceae; larvae likely in repro- the species. The genus Pnigodes is questionably distinct from
ductive structures (Clark 1982; Anderson 1991). See Clark (1982) Bagous. The genus is being revised by Charles O’Brien.
and Anderson (1991) to separate the species. Macropelmus Dejean 1821
Dietzia Champion 1903 Hydronomus Schoenherr 1825
Hamaba Casey 1910 Cyprus Schoenherr 1825
Rosella Whitehead 1977 Lyprus Schoenherr 1826
Dicranthus Motschulsky 1845
Ochyromerina Voss 1935 Ephimeropus Hochhuth 1847
Elmidomorphus Cussac 1851
Ochyromera Pascoe 1874, 1 sp., O. ligustri Warner 1961, southeast- Bagoas Gistel 1856
ern United States; adventive. This species is associated with Anactodes Brisout 1863
Ligustrum (adventive; privet; Oleaceae) (Warner 1961). Helminthimorphus Bedel 1884
Exochyromera Voss 1937 Bagoimorphus Desbrochers 1884
Parabagous Schilsky 1907
Incertae sedis (Curculioninae) Abagous Sharp 1916
Parabagous Sharp 1916; not Schilsky 1907
Macrorhoptus LeConte 1876, 6 spp., generally distributed in cen- Probagous Sharp 1916
tral and western United States and Canada. Species are associated Heterobagous Solari 1930
with Sphaeralcea, Sidalcea and Callirhoe (Malvaceae); larvae are in Himeniphades Kôno 1934
reproductive structures (Burke 1973). The genus needs revision. Memptorrhynchus Iablokov-Khnzorian 1960
See Sleeper (1957a) to separate the species. Fontenelleus Hoffmann 1962
Paraceratopus Brèthes 1910
Pnigodes LeConte 1876, 1 sp., P. setosus LeConte 1876, generally
distributed in central and southwestern United States. This ge-
V. Bagoinae Thomson 1859 nus is questionably distinct from Bagous.

by Robert S. Anderson VI. Baridinae Schoenherr 1836

Only the genera Bagous and Pnigodes constitute Bagoinae in North by Robert S. Anderson
America and the status of the latter as distinct is questionable.
Most are found in aquatic or semi-aquatic habitats where larvae Among all weevils, those in the Baridinae are in need of the most
are associated with a variety of plant families. Members are easily study. The group as a whole is difficult to characterize and generic
recognized by the median prosternal channel (Fig. 21), the smooth concepts and definitions need much refinement. There are many
varnish-like coating over the scales, the mostly tuberculate elytra, genera and some are of questionable validity. Some genera (e.g.,
and the elongate and slender legs. They are very similar in appear- Baris, Pseudobaris, Onychobaris, Sibariops, etc.) have numerous in-
ance to the Stenopelmina (Erirhininae) but the members of that cluded species but these have not been studied since their original
descriptions and many of them are still known only from type
 Family 131. Curculionidae · 741

series and localities. Thomas Lincoln Casey was the last person to of funicle, basal article of club about a third as
long as club ...................................................... 7
seriously study this subfamily and is responsible for most of the
generic and species concepts and names in use today. His types are 7(6). Prosternum with apical excavation but lacking sul-
all located at the Smithsonian Institution in Washington D.C. cus immediately anterior to coxae; elytra with
and a critical study of this collection is central to resolving the intervals flat; body nearly glabrous; femora not
toothed ........................................ Ampeloglypter
state of taxonomy in this subfamily.
— Prosternum with median sulcus extended from
Most baridines are glossy and black, with few (usually white) coxae to near apex; elytra with intervals various;
or no scales on the body, and are most readily recognized by an elytra often with scattered white scales or with a
ascended mesepisternum that is visible between the hind angle patch of white scales at base of interval 3; femora
with or without tooth ........................................ 8
of the pronotum and the elytral humerus. They share this latter
feature with Ceutorhynchinae but the latter have an exposed 8(7). Elytra with intervals rather wide and flat on disk
pygydium (shared with some Baridinae) and have a very small or (Fig. 25); body color black or dark piceous; elytra
no apical tooth on the hind tibia. This tooth is generally well- often with scattered white scales or with a patch
of white scales at base of interval 3; femora with
developed in Baridinae or the outer curved face of the tooth is
or without tooth .............................. Pseudobaris
continuous with the apex of the outer tibial margin or is con- — Elytra with intervals narrow and convex on disk (Fig.
nected to it by a distinct, continuous sharp carina that traverses 24); body color pale reddish brown; elytra nearly
the apical face of the tibia. glabrous; femora without tooth ... Desmoglyptus
The natural history of baridines is poorly known. Some
9(5). Antenna with club about as long as preceding six
species are associated with monocots such as various grasses, articles of funicle .......................... Hesperobaris
sedges, and palms. Larvae mostly mine stems. Some species in — Antenna with club shorter than preceding five ar-
the genera Buchananius and Plocamus appear to be associated with ticles of funicle .............................................. 10
fungi on dead wood. Adults, especially of the tribe Madopterini,
10(9). Rostrum distinctly separated from head by marked
frequently visit flowers. Many baridines can also be found in constriction at base of rostrum; body with dense
semi-aquatic habitats. scales; size greater than 3.0 mm ..... Trichobaris
— Rostrum at most slightly separated from head by
slight constriction at base of rostrum; body nearly
KEY TO THE NEARCTIC GENERA OF BARIDINAE
glabrous; size less than 2.0 mm ....... Microbaris
(slightly modified from Kissinger 1964)
11(4). Prosternum not sulcate in front of coxae and/or
1. Tarsus with a single claw .................................... 2 coxae separated by distance greater than width
— Tarsus with two claws (may be connate at base) .. of a coxa ........................................................ 12
......................................................................... 3 — Prosternum with deep, narrow sulcus in front of
coxae and/or coxae separated by distance less
2(1). Body subcylindrical in form, elytra about twice as than diameter of a coxa .................................. 14
long as wide; middle coxae separated by a dis-
tance less than the width of a coxa .. Barilepton 12(11). Rostrum short and stout, shorter than pronotum;
— Body elongate-oval in form, elytra about 1.5 times body nearly glabrous, with sparse, minute hair-
as long as wide; middle coxae separated by a like scales, elytra blue in color, intervals nearly
distance about equal to the diameter of a coxa impunctate ..................................... Zygobarinus
............................................................... Eisonyx — Rostrum long and slender, longer than pronotum;
body with some sparse, broad scales, elytra black
3(1). Tarsus with claws connate at base ..................... 4 or piceous in color, intervals with deep, coarse
— Tarsus with claws separate at base ................... 20 punctures ....................................................... 13

4(3). Pygydium not covered by elytra, broadly exposed, 13(12). Elytra with striae narrow, punctures wider than
punctate, nearly vertical ................................. 5 striae; prosternum in front of coxae with a pair of
— Pygydium more or less covered by elytra, mostly low ridges which are divergent posteriorly; south-
smooth, lacking obvious punctures, oblique .... ern Florida .......................................... Zygobaris
....................................................................... 11 — Elytra with striae broad, punctures not as wide as
striae; prosternum in front of coxae with ridges
5(4). Front coxae widely separated by a distance greater developed only near apex; southern Texas .....
than the width of a coxa .................................. 6 ........................................................ Zygobarella
— Front coxae narrowly separated by a distance less
than the width of a coxa .................................. 9 14(11). Prosternum behind coxae with deep narrow sulcus;
rostrum longer than pronotum, abruptly attenu-
6(5). Prosternum unimpressed in front of coxae; antenna ate immediately beyond antennal insertion, an-
with club more or less subcylindrical in shape, tennal insertion sub-basal (Fig. 28); antenna with
about as wide as article 7 of funicle, basal article article 2 of funicle long, more or less one-half
of club about half as long as club ...................... length of scape ............................... Amercedes
..................................................... Orchidophilus — Prosternum behind coxae lacking sulcus; rostrum
— Prosternum with median sulcus or apical impres- various in length, of more or less subequal width
sion in front of coxae; antenna with club more or throughout length, not attenuate beyond anten-
less oval in shape, distinctly wider than article 7 nal insertion, antennal insertion near or in front
742 · Family 131. Curculionidae

29

22

30

27 28
23 24 25 26

FIGURES 22.131-30.131. Baridinae. 22. Plocamus echidna (LeConte), lateral habitus. 23-26. Dorsal habitus, 23. Geraeus patagoniensis (Sleeper);
24. Desmoglyptus arizonicus Casey 1920; 25. Pseudobaris nigrina (Say); 26. Glyptobaris lecontei Champion. 27-28. Lateral view of head, 27. Baris sp.;
28. Amercedes subulirostris Casey. 29-30. Dorsal view of apex of rostrum and mandibles, 29. Odontocorynus salebrosus (Casey); 30. Haplostethops sp.

of middle of rostrum; antenna with article 2 of — Prosternum with median impression distinctly de-
funicle short, much less than length of scape .. fined and ridged laterally, wider anteriorly ........
....................................................................... 15 ...................................................... Catapastinus

15(14). Prosternum lacking sulcus medially in front of coxae; 20(3). Pygydium more or less completely exposed beyond
rostrum about as long as pronotum, slender, cy- elytral apex, punctate, nearly vertical ........... 21
lindrical, slightly curved ................ Strongylotes — Pygydium covered by elytra, mostly smooth, lack-
— Prosternum with median sulcus in front of coxae; ing obvious punctures, oblique ..................... 36
rostrum shorter than pronotum, stout, distinctly
c u r v e d ............................................................ 16 21(20). Hind tibia lacking tooth at apical margin or with tooth
or process shorter than tarsal claw ............... 22
16(15). Body nearly glabrous; prosternum in front of coxae — Hind tibia with tooth at apical margin about as long
with glabrous, median, broad sulcus with acute as tarsal claw .................................................. 25
lateral margins; pronotum with fine, shallow punc-
tures ...................................... Stethobaris (part) 22(21). Mandible prominent, triangular, inner face smooth
— Body with narrow or broad scales; prosternum in and straight ............ Pseudocentrinus (part; male)
front of coxae with median suclus with scales — Mandible with distinct teeth on inner face ....... 23
and with low, rounded lateral margins; pronotum
with deep punctures ...................................... 17 23(22). Antenna with club shorter than articles 2-7 of fu-
nicle, article 2 of funicle longer than 3; abdomi-
17(16). Mandibles prominent, not overlapped when closed; nal ventrite 5 distinctly longer than 3 and 4 com-
rostrum abruptly separated from head by deep bined .......................... Centrinogyna (part; male)
constriction; hind tarsus with article 5 longer than — Antenna with club longer than articles 2-7 of fu-
articles 1 and 2 combined .............. Acentrinops nicle, articles 2 and 3 of funicle subequal in
— Mandibles small, not prominent, overlapped when length; abdominal ventrite 5 shorter than 3 and 4
closed; rostrum only slightly separated from head combined ....................................................... 24
by slight constriction; hind tarsus with article 5
subequal in length to shorter than articles 1 and 24(23). Prosternum with shallow median sulcus in front of
2 combined .................................................... 18 coxae; rostrum in lateral view distinctly separated
from head by a marked transverse impression at
18(17). Antenna with basal article of funicle elongate, slen- base; scutellum quadrate ...................... Orthoris
der, longer than articles 2 to 5 combined; elytra — Prosternum flat in front of coxae, lacking median
with broad scales arranged in groups; body length sulcus; rostrum in lateral view at most slightly
greater than 3.0 mm; body form subcylindrical; separated from head by a slight transverse im-
tarsal claws connate to near apex ......... Barinus pression at base; scutellum triangular ...............
— Antenna with basal article of funicle stout, shorter ........................................................ Rhoptobaris
than articles 2 to 4 combined; elytra with some
solitary white scales; body length less than 2.5 25(21). Front coxae narrowly separated by a distance much
mm; body form oval; tarsal claws connate only at less than the width of a coxa ......................... 26
base ................................................................ 19 — Front coxae widely separated by a distance greater
than the width of a coxa ................................ 34
19(18). Prosternum with median impression indistinctly de-
fined laterally, wider posteriorly ...... Catapastus 26(25). Prosternum with deep, narrow median sulcus in front
of coxae ......................................................... 27
 Family 131. Curculionidae · 743

— Prosternum flat in front of coxae or with at most a — Surface of pronotum punctate; elytra glabrous or
slight median longitudinal impression ........... 28 nearly so, with at most fine short, hair-like scales;
prosternum produced posteriorly over mesoster-
27(26). Antenna with article 2 of funicle more than twice as num; mandible with inner face deeply notched
long as wide, as long as articles 3 and 4 com- ....................................................................... 35
bined; body form elliptical in dorsal view ..........
............................................................ Aulobaris 35(34). Femora with ventral margin with tooth; pronotum
— Antenna with article 2 of funicle less than twice as and elytra with fine, sparse punctures; body
long as wide, shorter than articles 3 and 4 com- nearly glabrous; elytra with surface uneven .....
bined; body form elongate, subparallel in dorsal .......................................................... Madarellus
view ................................................... Trepobaris — Femora with ventral margin simple, lacking tooth;
pronotum and elytra with deep, uniform punc-
28(26). Elytra with striae 1and 2 deeply linearly punctate in tures; body with fine short, suberect hair-like
basal one-third, striae deeply continuously im- scales; elytra with surface more or less even ...
pressed in apical two-thirds only; male with ros- ....................................................... Onychobaris
trum with ventral surface with long dense pilos-
ity ......................................................... Myctides 36(20). Body with sparse, erect, coarse setae (Fig. 22); body
— Elytra with striae 1and 2 deeply continuously im- size small, less than 3.0 mm ........................... 37
pressed throughout entire length; male with ros- — Body with at most fine setae or appressed scales;
trum with ventral surface glabrous or with a few body size various .......................................... 38
short setae ..................................................... 29
37(36). Front coxae widely separated by distance much
29(28). Rostrum in lateral view continuous with head, not greater than width of a coxa; prosternum flat in
separated from head by transverse impression; front of coxae; body densely covered with broad,
eyes large, extended onto dorsal surface of head; appressed scales in addition to erect setae (Fig.
frons about one-half as wide as rostrum at apex; 22) ....................................................... Plocamus
body subcylindrical in form .............. Stenobaris — Front coxae narrowly separated by distance less
— Rostrum in lateral view distinctly separated from than width of a coxa; prosternum medially shal-
head by transverse impression; eyes smaller, lat- lowly, broadly impressed in front of coxae; body
eral, not extended onto dorsum of head; frons with at most some fine hair-like scales in addition
about as wide as rostrum at apex; body various in to erect setae ................................. Buchananius
form ................................................................ 30
38(36). Mandibles prominent, triangular in form when
30(29). Pronotum with sides covered with broad, round viewed dorsally, not or only slightly overlapped
scales; body with white and tan colored scales or crossed when closed (Fig. 29) ................... 39
intermixed ....................................... Cosmobaris — Mandibles not prominent, less obviously triangular
— Pronotum with sides lacking broad, round scales; in form when viewed dorsally, overlapped or
body either subglabrous or with only white scales crossed when closed (Fig. 30) ....................... 52
....................................................................... 31
39(38). Mandible with inner face smooth (Fig. 29), not den-
31(30). Rostrum in lateral view separated from head by shal- tate or emarginate, usually straight but divergent
low, broad impression (Fig. 27) ....................... 32 in some specimens ........................................ 40
— Rostrum in lateral view separated from head by a — Mandible with inner face dentate or crenulate,
deep groove or dorsal constriction of the base of straight ........................................................... 45
the rostrum ..................................................... 33
40(39). Abdomen with ventrite 5 tumid medially, more than
32(31). Elytra subglabrous, vestiture fine, minute, of uni- three times as long as ventrites 3 and 4 com-
form length and form; antenna with basal article bined; prosternum deeply, narrowly sulcate in
of club less pubescent and more glossy than front of coxae ..... Pseudocentrinus (part; female)
other articles of club; body size 2.8-6.0 mm .... — Abdomen with ventrite 5 flat medially, at most
................................................................... Baris slightly longer than ventrites 3 and 4 combined;
— Elytra fine minute hair-like vestiture as well as broad, prosternum various ........................................ 41
white scales which form a spot at the base of
elytral interval 3 (other spots may also be 41(40). Antenna with distal articles of funicle obliquely trun-
present); antenna with club uniformly pubescent; cate, club with basal article with glabrous area
body size 1.8-3.0 mm ........................ Plesiobaris with blunt or dentiform process on inner face ..
................................ Odontocorynus (part; male)
33(31). Pronotum broadly constricted at apex, not tubulate; — Antenna with distal articles of funicle and basal ar-
vestiture of fine scales, recumbent .................. ticle of club simple, not modified .................. 42
......................................................... Pycnobaris
— Pronotum sharply constricted at apex, tubulate; 42(41). Male with prosternum flat in front of front coxae ..
vestiture of fine scales and setae intermixed, ....................................................................... 43
suberect ........................................... Stictobaris — Male with prosternum deeply excavated anterior to
front coxae ..................................................... 44
34(25). Surface of pronotum distinctly rugose (Fig. 26);
elytra with coarse elongate yellow scales and 43(42)4. Pronotum with anterior tubulate portion with a longi-
fine setae; prosternum produced posteriorly only tudinal fold on each side; prosternum with vestiture
slightly over mesosternum; mandible with inner on median line not radiating from a central point,
face nearly smooth ......................... Glyptobaris but directed backwards (Fig. 23) ............. Geraeus
744 · Family 131. Curculionidae

— Pronotum with anterior tubulate portion lacking a 51(50). Pronotum constricted apically, markedly tubulate
longitudinal fold on each side; prosternum with ......................................................... Centrinites
vestiture on median line radiating from a central — Pronotum not constricted or tubulate at apex ......
point, lying either behind, on, or before the pos- ............................................................ Nicentrus
terior line of the tubulate portion ... Linogeraeus
52(38). Lateral profile with dorsal surface markedly, evenly
44(42). Prosternum of male with large, deep pit between convex; elytra with striae obsolete, indicated
prosternal spines ......................... Pachygeraeus only by rows on punctures ............................ 53
— Prosternum of male flat between spines but with — Lateral profile with dorsal surface flattened near
transverse, deep, median anterior fossa ........... middle, not evenly convex; elytra with striae dis-
..................................................... Pycnogeraeus tinct, moderately deep ................................... 54

45(39). Front coxae widely separated by distance equal to 53(52). Prosternum with median longitudinal impression in
width of a coxa; hind tibia with large tooth at api- front of coxae; pronotum markedly tubulate at
cal margin at least as long as a tarsal claw .... 46 apex .............................................. Oomorphidius
— Front coxae narrowly separated by distance obvi- — Prosternum flat in front of coxae; pronotum not
ously less than width of a coxa; hind tibia lacking tubulate at apex ............................ Cholinobaris
tooth at apical margin or tooth obviously shorter
than a tarsal claw ............................................ 48 54(52). Shortest distance between middle and hind coxae
less than, or equal to, one-half length of
46(45). Rostrum with point of antennal insertion medial; metepisternum ............................................... 55
prosternum flat, lacking sulcus; body elongate — Shortest distance between middle and hind coxae
oval in form ...................................... Calandrinus usually greater than one-half length of
— Rostrum with point of antennal insertion in basal metepisternum ............................................... 58
one-half; prosternum with median sulcus in front
of coxae; body broadly oval in form .............. 47 55(54). Antenna with article 2 of funicle about as wide as
long, stout; antennal club large, elongate, about
47(46). Elytra, except at base, nearly glabrous; pronotum as long as funicle; body nearly glabrous, with
with basal margin distinctly produced posteriorly minute hair-like scales .................... Stethobaris
and emarginate immediately in front of scutel- — Antenna with article 2 of funicle distinctly longer
lum; front tibia simple, not excavated to receive than wide, slender; antennal club moderately long,
base of tarsus .................................. Pachybaris not as long as funicle; body with fine, but dis-
— Elytra with scattered broad white scales; pronotum tinct, long hair-like scales .............................. 56
with basal margin distinctly produced posteriorly
but not emarginate immediately in front of scutel- 56(55). Pronotum constricted apically but not tubulate ...
lum; front tibia with outer surface at apex deeply ......................................................... Oligolochus
excavated to receive base of tarsus ................. — Pronotum tubulate at apex ................................ 57
.................................................... Diorymeropsis
57(56). Prosternum deeply sulcate in front of coxae; body
48(45). Rostrum with point of antennal insertion behind lacking dense white scales on venter ..............
midlength; prosternum of female with median lon- ......................................................... Idiostethus
gitudinal sulcus in front of coxae, in male, with — Prosternum shallowly sulcate in front of coxae; lat-
two erect processes in front of which is an elon- eral margins of thoracic sterna with dense, white,
gate-oval impression ....................... Centrinopus overlapped scales ...................... Haplostethops
— Rostrum with point of antennal insertion at or in
front of midlength; prosternum of male unarmed 58(54). Prosternum of male with a pair of erect, slender pro-
in front of coxae ............................................ 49 cesses in front of coxae or with low transverse
carina ............................................................. 59
49(48). Metasternum short, middle and hind coxae sepa- — Prosternum of male unarmed in front of coxae .....
rated by less than length of abdominal ventrite 1 ....................................................................... 60
behind hind coxa; body broadly oval in dorsal
form .................................................. Microcholus 59(58). Body form in dorsal view elongate-oval; prosternum
— Metasternum longer, middle and hind coxae sepa- of male with short, slender spines ...... Sibariops
rated by a distance at least equal to length of — Body form in dorsal view slender, subcylindrical;
abdominal ventrite 1 behind hind coxa; body elon- prosternum of male with long, slender spines and
gate in dorsal form ......................................... 50 a deep rounded fossa anterior to spines ...........
.......................................................... Cylindridia
50(49). Front coxae separated by more than one-half width
of a coxa; mandible with outer face denticulate 60(58). Front coxae narrowly separated by distance dis-
..................................................... Centrinogyna tinctly less than width of a coxa ................... 61
— Front coxae narrowly separated by distance less — Front coxae widely separated by distance greater
than one-half width of a coxa (exception some than width of a coxa ....................................... 62
Nicentrus); mandible with outer margin not den-
ticulate ........................................................... 51 61(60). Elytra with fine, inconspicuous hair-like scales;
pronotum strongly constricted at apex and
tubulate ................................................. Dirabius
4
The genus Centrinus with only the species C. pistor (Germar 1824) — Elytra with moderately coarse, elongate, white con-
should key here (not seen by me). Its relationship to Geraeus and spicuous scales; pronotum not constricted or
Linogeraeus needs to be reassessed (see text). tubulate at apex ......................... Trichodirabius
 Family 131. Curculionidae · 745

62(60). Body with fine, elongate scales; antenna with fu- Pycnobaris Casey 1892, 4 spp., Kansas, Colorado, Texas and Cali-
nicle with very fine, long setae ............ Apinocis
fornia. See Casey (1892, 1920) to separate the species.
— Body with coarse, elongate scales; antenna with
coarse, elongate scales ...................... Barilepsis
Rhoptobaris LeConte 1876, 1 sp., R. canescens LeConte 1876, Kan-
sas, Colorado, Texas and Oregon.
CLASSIFICATION OF THE NEARCTIC BARIDINAE
Stenobaris Linell 1897, 1 sp., S. avicenniae Linell 1897, Florida. This
18. Baridini Schoenherr 1836 species is associated with Avicennia germinans (L.) L. (black man-
grove; Avicenniaceae) (Linell 1897).
Baridina Schoenherr 1836
Trepobaris Casey 1892, 1 sp., T. elongata Casey 1892, Arizona and
Aulobaris LeConte 1876, 10 spp., generally distributed in eastern Texas.
United States and Canada, also California. Some species are asso-
ciated with wetlands. See Casey (1892, 1920) to separate the spe- Trichobaris LeConte 1876, 9 spp., generally distributed in eastern and
cies. The genus needs revision. southern United States and southern Canada. Species are associated
with various Solanaceae; larvae are in stems (Barber 1935; Cuda and
Baris Germar 1817, 92 spp., generally distributed. Species are as- Burke 1985). See Barber (1935) to separate the species.
sociated with various plants, mostly Asteraceae. See Casey (1892)
and Gilbert (1964) to separate some of the species. The genus 19. Madarini Jekel 1865
needs revision; many species are of questionable validity.
Baridius Schoenherr 1825 Madarina Jekel 1865
Cyphirhinus Schoenherr 1826
Aegyptobaris Pic 1889 (valid subgenus) Ampeloglypter LeConte 1876, 3 spp., generally distributed in east-
Turkmenobaris Zaslavskij 1956 (valid subgenus) ern United States and southern Canada. Species are associated
with Vitis (grape; Vitaceae); larvae make galls on stems (Kissinger
Cosmobaris Casey 1920, 1 sp., C. americana Casey 1920, generally 1964). See Blatchley and Leng (1916) to separate the species.
distributed. This species is associated with Chenopodium
(Chenopodiaceae); larvae mine stems (Kissinger 1964). Glyptobaris Casey 1892, 1 sp., G. lecontei Champion 1909, generally
distributed in the eastern United States.
Desmoglyptus Casey 1892, 2 spp., Maryland, Pennsylvania, Vir-
ginia, District of Columbia, and Arizona. Species are associated Madarellus Casey 1892, 5 spp., generally distributed in the eastern
with Vitis (grape; Vitaceae) (Kissinger 1964). See Casey (1920) to United States and Canada west to Texas. Species are associated
separate the species. with Vitis (Vitaceae) (Blatchley and Leng 1916). See Casey (1892,
1920) and Blatchley and Leng (1916) to separate the species. The
Hesperobaris Casey 1892, 2 spp., Missouri, Kansas and Texas. See genus needs revision.
Casey (1892) to separate the species. Willinkia Bondar 1949

Microbaris Casey 1892, 1 sp., M. galvestonica Casey 1892, Texas. Onychobaris LeConte 1876, 33 spp., generally distributed in the
United States, but especially in the southwest including Califor-
Orthoris LeConte 1876, 7 spp., generally distributed in the western nia. See Casey (1892, 1920) to separate some of the species. The
United States and Canada. Species are associated with Mentzelia genus needs revision; many species are of questionable validity.
(Loasaceae); larvae are in pods, stems and roots (Pierce 1907). See
Casey (1892, 1920) to separate the species. The genus needs revision. Orchidophilus Buchanan 1935, 3 spp., New Jersey, District of Co-
lumbia, and California; adventive in orchid houses, likely not
Plesiobaris Casey 1892, 6 spp., generally distributed in eastern United established in the wild. See Buchanan (1935) to separate the spe-
States. Species are associated with Hypericum (Hypericaceae) in cies.
wetlands. See Casey (1892, 1920) to separate the species.
Stictobaris Casey 1892, 4 spp., southwestern and central United
Pseudobaris LeConte 1876, 31 spp., generally distributed in eastern States. See Sleeper (1957b) to separate the species.
United States and Canada, west to California, Colorado and Utah.
At least one species is associated with Lycopus (Labiatae) (Kissinger Tonesiina Alonso-Zarazaga and Lyal 1999
1963). See Casey (1892) to separate the species. The genus needs
revision; many species are of questionable validity. Myctides Pascoe 1874, 1 sp., M. imberbis Lea 1906, Florida; adven-
Pseudobaridia Casey 1920 (valid subgenus) tive. This species is associated with Syzygium jambos Alston
(Myrtaceae); larvae in seeds (Woodruff 1977; Anderson 1993a).
746 · Family 131. Curculionidae

20. Madopterini Lacordaire 1866 Centrinites Casey 1892, 2 spp., generally distributed in the eastern
United States. Species are associated with Melanthium virginicum L.
Torcina Bondar 1943 (Liliaceae) (Blatchley and Leng 1916). See Casey (1920) to separate
the species.
Sibariops Casey 1920, 41 spp., generally distributed in eastern United Leptosaldius Casey 1922
States and Canada. Species are associated with sedges in wetlands.
See Casey (1920) to separate the species. The genus needs revi- Centrinogyna Casey 1892, 5 spp., generally distributed in the west-
sion; many species are of questionable validity. ern United States and central Canada. See Casey (1892, 1920) to
separate the species. The genus needs revision.
Zygobaridina Pierce 1907
Centrinopus Casey 1892, 6 spp., generally distributed in the eastern
Acentrinops Casey 1920, 1 sp., A. brevicollis Casey 1920, New Mexico United States. Adults are found on flowers of Asteraceae
and Texas. (Kissinger 1964). See Casey (1920) to separate the species. The
genus needs revision.
Amercedes Casey 1894, 1 sp., A. subulirostris Casey 1893, Louisiana
and Texas. This species is associated with Zanthoxylum (Rutaceae) Centrinus Schoenherr 1825, 1 sp., C. pistor (Germar 1824), Ken-
(Pierce 1907). tucky. This species is of uncertain affinity. It has not been seen by
Zygobaroides Pierce 1907 me and its relationships to Geraeus and Linogeraeus need to be
reassessed. Champion (1908: 261-261) limited Centrinus to large
Apinocis Lea 1927, 15 spp., generally distributed. Larvae of at least South American species with the mandibles strongly denticulate
one species, A. saccharinus (Marshall 1952), have been associated along the inner margins.
with grasses (Poacaeae) (Woodruff 1972). See Buchanan (1932) Toxeres Germar 1829
to separate most of the species. The genus needs revision. Toxeres Schoenherr 1833; not Germar 1829
Prosaldius Ogloblin 1930 Telephus Gistel 1848
Anacentrinus Buchanan 1932
Cholinobaris Casey 1920, 1 sp., C. rhomboidea Casey 1920, North
Barilepis Casey 1920, 3 spp., generally distributed in the eastern Carolina.
United States, Arizona and Texas. See Casey (1920) to separate
the species. Cylindridia Casey 1920, 4 spp., generally distributed in the eastern
United States and southern Canada, west to Texas and Colorado.
Barilepton LeConte 1876, 4 spp., generally distributed in eastern Species are associated with sedges (Cyperaceae). See Casey (1920)
and southern United States. Species are associated with wetlands. to separate the species.
See Casey (1892) to separate the species.
Diorymeropsis Champion 1908, 1 sp., D. xanthoxyli (Linell 1897),
Barinus Casey 1892, 14 spp., generally distributed in eastern United Texas. This species is associated with Zanthoxylum (Rutaceae).
States, also California. Species are associated with sedges in wet- Pseudogarnia Casey 1920
lands. See Sleeper (1956a) to separate the species.
Dirabius Casey 1920, 9 spp., generally distributed in the eastern
Buchananius Kissinger 1957, 2 spp., generally distributed in east- United States and southern Canada, also one species in Califor-
ern United States. Species are associated with various dead tree nia. One species has been associated with Scirpus cyperinus L.
limbs on the ground (Kissinger 1964). See Kissinger (1958) or (Cyperaceae); larvae are in the stems (Blatchley and Leng 1916).
Blatchley and Leng (1916) to separate the species. Limnobaropsis Casey 1920 (valid subgenus)
Zaglyptus LeConte 1876; not Foerster 1868
Eisonyx LeConte 1880, 3 spp., Texas, New Mexico, Kansas, Okla-
Calandrinus LeConte 1876, 2 spp., New Mexico, Texas, Colorado homa, Missouri, Iowa and Tennessee. Species are associated with
and Alberta. See Casey (1892, 1920) to separate the species. Senecio and Hymenoxys (Asteraceae); larvae are in stems, crowns
and roots (Pakaluk and Carlow 1994). See Pakaluk and Carlow
Catapastinus Champion 1908, 1 sp., C. caseyi Champion 1908, (1994) to separate the species.
southern Texas. This species is associated with Zanthoxylum fagara Eumononycha Casey 1893 (valid subgenus)
(L.) Sarg. (Rutaceae).
Geraeus Pascoe 1889, 21 spp., generally distributed in the eastern
Catapastus Casey 1892, 6 spp., generally distributed in the eastern and southwestern United States and southeastern Canada. At
United States, Florida and Texas. Species are associated with vari- least some species are associated with grasses (Poaceae) (Kissinger
ous species of Zanthoxylum (Rutaceae). See Casey (1892, 1920) to 1964); adults frequently visit flowers. See Casey (1892, 1920) and
separate the species. Blatchley and Leng (1916) to separate some of the species; con-
 Family 131. Curculionidae · 747

sult O’Brien and Wibmer (1984) for a listing of species included Pachybaris LeConte 1876, 1 sp., P. porosa LeConte 1876, Florida
in the genus. The genus needs revision; many species are of and Louisiana. Adults have been associated with flowers of Ser-
questionable validity. Generic definitions in this part of the enoa repens (Bartr.) Small and Sabal palmetto (Walt.) Lodd (saw
Baridinae need much study. palmetto and cabbage palm; Arecaceae) (Anderson 1993a).
Centrinaspis Casey 1920
Pachygeraeus Casey 1920, 3 spp., generally distributed in the central
Haplostethops Casey 1920, 7 spp., central United States, four spe- United States. See Casey (1920) to separate the species.
cies known only from Missouri. Species are associated with wet-
lands. See Casey (1920) to separate the species. The genus needs Plocamus LeConte 1876, 2 spp., generally distributed in the east-
revision; some species are of questionable validity. ern United States and southern Canada. Adults have been associ-
ated with hickory, beech and maple (Blatchley and Leng 1916). See
Idiostethus Casey 1892, 16 spp., generally distributed in the eastern Blatchley and Leng (1916) to separate the species.
United States and southern Canada. Adults have been associated Euchaetes LeConte 1876; not Harris 1841; not Philippi 1843;
with various flowers (Blatchley and Leng 1916); larvae may be asso- not Sclater 1858
ciated with orchids. See Casey (1892, 1920) to separate the species. Eunyssobia Casey 1892
The genus needs revision; many species are of questionable validity. Epeuchaetes Lyman 1902

Linogeraeus Casey 1920, 15 spp., generally distributed in the south- Pseudocentrinus Champion 1908, 1 sp., P. ochraceus (Boheman
eastern and southwestern United States. Adults frequently visit flow- 1844), Texas.
ers. See Casey (1920) to separate some of the species; consult O’Brien
and Wibmer (1984) for listing of species included in the genus. The Pycnogeraeus Casey 1920, 3 spp., Pennsylvania, Florida, Texas, New
genus needs revision; many species are of questionable validity. Mexico and Arizona. See Casey (1892) to separate the species.
Stereogeraeus Casey 1920
Conocentrinus Casey 1920 Stethobaris LeConte 1876, 11 spp., generally distributed in the eastern
Glyptogeraeus Casey 1920 United States and southern Canada, west to Texas and Arizona. A
Brachygeraeus Casey 1920 number of the species are associated with orchids (Brown 1966a;
Centrinaspidia Casey 1920 Hull Sieg and O’Brien 1993; Howden 1995). See Casey (1892) to
Lepidobaris Lea 1927; not Champion 1909 separate some of the species. The genus needs revision.
Diorymerellus Champion 1908
Microcholus LeConte 1876, 2 spp., New Jersey, Georgia and Florida.
Species are associated with wetlands. See Casey (1892) and Blatchley Trichodirabius Casey 1920, 2 spp., Florida, Louisiana and Texas.
and Leng (1916) to separate the species. See Casey (1920) to separate the species.

Nicentrus Casey 1892, 20 spp., generally distributed in the eastern Zygobarella Casey 1920, 1 sp., Z. xanthoxyli (Pierce 1907), Texas.
United States west to Texas and Arizona. Adults are frequently This species is associated with Zanthoxylum (Rutaceae); larvae de-
found on flowers in various habitats. See Casey (1892, 1920) and velop in fruits (Pierce 1907).
Blatchley and Leng (1916) to separate some of the species. The
genus needs revision. Zygobarinus Pierce 1907, 1 sp., Z. coelestinus (Linell 1897), Florida.
Nicentrites Casey 1922
Eunicentrus Casey 1922 Zygobaris LeConte 1876, 1 sp., Z. nitens LeConte 1876, Florida.
Adults have been collected on Zanthoxylum flavum Vahl. (Rutaceae)
Odontocorynus Schoenherr 1844, 51 spp., generally distributed in (Anderson 1993a).
the eastern United States and southern Canada, west to Texas
and Colorado. Adults are frequently found on flowers (mostly 21. Nertinini Voss 1954
Asteraceae) in various habitats. See Casey (1920) and Blatchley
and Leng (1916) to separate some of the species. The genus Strongylotes Schoenherr 1836, 1 sp., S. parallelus Champion 1907,
needs revision; many species are of questionable validity. Texas and Arizona.

Oligolochus Casey 1892, 7 spp., generally distributed in the eastern

United States, Arizona and California. See Buchanan (1932) to VII. Ceutorhynchinae Gistel 1856
separate the species.
Anacentrus Casey 1920 by Boris A. Korotyaev and Robert S. Anderson

Oomorphidius Casey 1892, 2 spp., southeastern and central United Ceutorhynchinae are a relatively well-known group of small wee-
States. See Casey (1892) to separate the species. vils found in both terrestrial and freshwater aquatic habitats
748 · Family 131. Curculionidae

throughout North America. They are readily recognized by the — Rostrum much longer than pronotum, more than 3
times as long as wide; if short and thick, then
ascended mesepimeron (as in Baridinae), an exposed pygydium,
femora simple, unarmed, and hind femur weakly
and presence of a small or no tooth at the apex of the hind tibia. widened, less than 1.3 times as wide as middle
Many possess a prosternal channel for the reception of the ros- one; associated with various plants, not
trum and have pronotal postocular lobes that cover the eyes when Portulaceae ...................................................... 3
the rostrum is in repose, but some do not. Adults of some
3(2). Rostrum no more than 3 times as long as wide (Fig.
species have expanded hind femora and are good jumpers. 35), wider than front femur; femora unarmed; an-
Species of Ceutorhynchinae are associated with a variety of terior margin of pronotum not raised, often with 2
plant families. In terrestrial habitats the Cruciferae are a common more or less acute denticles or with emargination
limited by angular prominences ....................... 4
host, whereas in aquatic habitats the most common host would
— Rostrum more than 3 times as long as wide (Fig. 34);
appear to be Polygonaceae or semi-aquatic Cruciferae. Larvae of if less than 3 times, then femora dentate, anterior
terrestrial species usually mine the stems or crowns of the plants margin of pronotum strongly raised and size over
but some aquatic taxa in the Phytobiini such as Phytobius have 3.5 mm (Phrydiuchus), or metasternum between
middle coxae depressed ............................... 12
larvae that live and feed externally on plant reproductive struc-
tures. Some species in the genus Ceutorhynchus are adventive and 4(3). Antenna with funicle of 7 articles; tarsal claws den-
serious pests of cultivated Cruciferae (especially rapeseed or canola) tate; anterior margin of pronotum simple, without
in western North America. Species in the genera Phrydiuchus, sharp denticles; prosternum anterior to front
coxae long, with high keels; distance between
Microplontus, Mogulones and Trichosirocalus have been deliberately
front coxae usually not less than one-half width
introduced for the biological control of pest weeds. of rostrum ........................................... Rhinoncus
— Antenna with funicle of 6 articles; tarsal claws den-
KEY TO THE NEARCTIC GENERA OF CEUTORHYNCHINAE tate or simple; anterior margin of pronotum often
with 2 sharp denticles; prosternum anterior to
front coxae sometimes short, with low, sometimes
1. Tarsus with single claw; body size 3.6-5.5 mm; as-
obsolete keels; distance between front coxae
sociated with Iris versicolor L. (Iridaceae) .........
less than one-half width of rostrum ................. 5
....................................................... Mononychus
— Tarsus with 2 claws; body size 1.5-5.0 mm (most
5(4). Tarsal claw with well-developed tooth at base;
less than 3.5 mm); associated with various plants,
prosternum in front of coxae deeply excavated
not Iridaceae .................................................... 2
with its anterior margin deeply angularly emar-
ginate, the emargination extending behind the
2(1). Rostrum shorter than pronotum, thick, not more than
level of anterior margins of coxal cavities; inner,
3 times as long as wide, weakly and unevenly
usually also posterior margins of eyes sharply
curved; femora with tooth on ventral margin; hind
raised ............................................................... 6
femur rather strongly widened, 1.5-2.0 times as
— Tarsal claw simple, if dentate (in Neophytobius), then
wide as middle femur; associated with Portulaca
front coxae separated at most by width of anten-
oleracea L. (Portulaceae) ....................... Hypurus
nal club and apical margin of pronotum narrowly

34 35
32 33
31
36

37 38
FIGURES 31.131-38.131. Ceutorhynchinae. 31-33. Dorsal habitus, 31. Allosirocalus sp.; 32. Glocianus punctiger (Gyllenhal); 33. Homorosoma
sulcipennis (LeConte). 34-35. Lateral view of head, 34. Ceutorhynchus rapae Gyllenhal; 35. Rhinoncus pericarpius (Linnaeus). 36. Perigasteromimus
tetracanthus (Champion), head and rostrum, dorsal view. 37-38. Front tibia, 37. Acanthoscelis acephalus (Say); 38. Glocianus punctiger (Gyllenhal).
 Family 131. Curculionidae · 749

excised; prosternum in front of coxae very ming hairs; middle coxae separated by about their
weakly depressed and shallowly arcuately emar- width .................................................... Parenthis
ginate, the emargination not reaching the level
of anterior margin of coxal cavities; inner mar- 11(8). Tarsus with claws dentate; apical margin of
gins of eyes slightly, if at all, raised ................ 8 pronotum at middle produced anteriorly and not
raised, with shallow emargination narrower than
6(5). Rostrum about 3 times as long as wide, scarcely base of rostrum, margin lateral to emargination
widened apically; antennal scrobe well devel- finely serrate; elytra with alternate intervals rather
oped, the dorsal margin reaching eye; elytra with strongly convex, with rows of large sharp gran-
alternate intervals moderately to rather strongly ules ............................................... Neophytobius
convex, more strongly so on apical prominences — Tarsus with claws simple; apical margin of pronotum
............................................................. Dietzella in the middle not produced anteriorly, with
— Rostrum usually not more than twice as long as wide, straight part limited by 2 sharp tubercles, distance
strongly widened apically; antennal scrobe fovei- between them not less than width of rostrum,
form or very weakly developed, vanishing at most margin lateral to tubercles smooth, not serrate;
halfway to eye; elytra with alternate intervals not elytra with alternate intervals not conspicuously
conspicuously convex .................................... 7 c o n v e x .............................................. Pelenomus

7(6). Rostrum less than twice as long as wide; antennae 12(3). Prosternum anterior to front coxae short, without
inserted on dorsal surface of rostrum, scape very traces of keels; distance between front coxae
short, shorter than two basal articles of funicle equal to width of antennal funicle; antenna with
combined (Fig. 36); Florida ... Perigasteromimus funicle with 6 articles ............................ Amalus
— Rostrum twice or more as long as wide, antennae — Prosternum anterior to front coxae longer, with more
inserted laterally; scape longer than two basal or less developed keels; front coxae usually
articles of funicle combined; widespread ......... widely separated by distance greater than width
........................................................... Perigaster of antennal funicle; antenna with funicle with 7
articles, if of 6 articles, then claws simple, or
8(5). Body with very dense vestiture of short, recum- femora dentate, or rostrum wider than front femur
bent or subrecumbent matte hydrophobous ....................................................................... 13
scales; narrow metallic-glossy scales may be
present only on apical part of rostrum; tarsi nar- 13(12). Rostrum usually wider than width of front femur; if
row, 3rd article often scarcely wider than article about as wide, then at least mesosternum dis-
2; if (in Parenthis) 1.5 times as wide as the latter, tinctly depressed (flat in Phrydiuchus, recognized
then anterior margin of pronotum without den- by large body size of 4.0-5.0 mm); antenna with
ticles and inconspicuously emarginate medially; scape with elongate lamelliform translucent pro-
tarsal claw simple ............................................. 9 jection and/or 1-3 setae at apex ................... 14
— Body with sparse covering of scales composed — Rostrum narrower than width of front femur; meso-
partly (at least on rostrum) or mostly from metal- sternum mostly flat (only in Nedyus deeply de-
lic-glossy narrow scales; tarsi wider, 3 rd article pressed); antenna with scape lacking apical pro-
1.5 to (mostly) 2.0 times as wide as article 2; tar- jections or setae ............................................ 27
sal claw simple or dentate ............................. 11
14(13). Body globose, elytra slightly longer than wide, with
9(8). Body size larger, 2.6-3.0 mm long; tarsus very long completely rounded shoulders and 7 th stria al-
and narrow, 3rd tarsal article slightly wider than most reaching basal margin of elytron; meso- and
article 2; dorsal and lateral surface of tarsal ar- metasterna very short, not more than half length
ticles lacking long swimming hairs; pronotum with of respective coxae ............................ Acallodes
acute lateral tubercles; its rounded apical margin — Body less convex and rounded, elytra usually
with shallow median emargination limited by acute rounded-triangular or with staightened sides;
angulations; elytra with interval 5 keel-shaped meso- and metasterna longer than one-half of re-
and finely muricate in basal one-third ............... spective coxae .............................................. 15
............................................................ Phytobius
— Body size smaller, 2.2-2.7 mm long; 3rd tarsal article 15(14). Meso- and metasterna flat; body size large, 4.0-5.0
1.6-1.7 times as wide as article 2; if scarcely wider, mm; associated with Salvia (Labiatae) ................
then tarsus dorsally with a few long and very fine ........................................................ Phrydiuchus
swimming hairs; pronotum with obsolete obtuse — Meso- and often metasternum more or less deeply
lateral tubercles and apical margin lacking any depressed for reception of rostrum; body size
trace of median emargination or acute angulations; smaller, less than 3.5 mm; associated with vari-
elytra with interval 5 not carinate in basal part . ous plants, not Labiatae ................................. 16
....................................................................... 10
16(15). Mesosternum more or less deeply depressed but
10(9). Tarsus with article 3 less than 1.4 times as wide and depression not limited by keels at sides; if rostral
about one-half as long as article 2; dorsal and sulcus extends onto metasternum, its sides gen-
lateral surface of tarsal articles covered with long tly sloping; femora with ventral tooth ............ 17
very fine semi-erect swimming hairs; middle — Mesosternum with depression limited by keels at
coxae separated by about one-half their width sides, often extended onto metasternum and
..................................................... Euhrychiopsis walls very steep or abrupt; femora simple, lack-
— Tarsus with 3rd article 1.6-1.7 times as wide and al- ing ventral tooth ............................................ 19
most as long as article 2; tarsi lacking long swim-
750 · Family 131. Curculionidae

17(16). Antenna with funicle of 7 articles; meso- and forming acute divergent ridges; elytra sparsely
metasterna very shallowly depressed; elytra clothed with narrow, lanceolate scales arranged
rounded-triangular, with moderately large sharp in narrow broken transverse bands; body size
granules densely arranged along intervals and 2.3-3.6 mm .......................................... Craponius
provided with short scale-like subrecumbent seta — Outer margin of middle and hind tibiae sometimes
apically (Fig. 33); elytral disc lacking scale pat- more or less distinctly grooved but not conspicu-
tern other than short postscutellar spot on su- ously emarginate; pronotum with 2 acute ridge-
tural interval (Fig. 33); body size larger, 2.1-3.2 shaped discal prominences divergent anteriorly;
mm ................................................. Homorosoma elytra with moderately dense vestiture partly
— Antenna with funicle of 6 articles; meso- and formed by lanceolate to oval scales; body size
metasterna moderately deeply depressed; elytra 2.1-2.3 mm ................................ Orchestomerus
either with straightened sides in basal one-half
and vague transverse band of white scales im- 22(20). Outer margin of front and middle tibiae very deeply
mediately behind middle, or with very large acute emarginate and compressed, almost blade-
granules on intervals and entire body with very shaped, lacking tarsal grooves, emarginations lim-
long erect hairs; body size smaller, 1.9-2.4 mm ited by large acute dentiform prominences .....
....................................................................... 18 ....................................................... Cnemogonus
— Outer margin of all tibiae straight or inconspicu-
18(17). Elytra with sides in basal one-half parallel or weakly ously emarginate, sometimes with more or less
rounded; body lacking erect hairy pubescence; developed tarsal grooves .............................. 23
elytral intervals with small rounded granules; as-
sociated with Salicaceae ................ Rutidosoma 23(22). Front tibia with outer margin with acute dentiform
— Elytra with sides rounded evenly from base; body prominence apically (Fig. 37); body size large,
with long and fine, erect pubescence; elytral in- 2.6-3.7 mm ............................... Acanthoscelidius
tervals with rows of sparsely arranged, very large, — Front tibia with outer apical angle not produced
acute piliferous granules; associated with into acute prominence (Fig. 38); if weakly pro-
Heuchera richardsoni (Saxifragiaceae) .............. duced (in Auleutes isolatus Sleeper) or with 2
....................................................... Asperosoma spines larger than other setae in apical comb (in
Auleutes asper LeConte), then dorsal surface with
19(16). Rostrum dilated or, rarely, parallel-sided in female moderately dense vestiture of white scales and
(in Auleutes donaldi Colonnelli) in apical part; ros- more or less distinct scutellar spot, elytral inter-
tral sulcus ending between middle coxae on vals with one row of sharp granules each; body
metasternum, very deep, its walls abrupt; antenna size usually smaller, 2.1-3.5 mm ..................... 24
with club with dense, short, very fine erect pu-
bescence (may be absent on large basal seg- 24(23). Pronotum with a pair of rather high discal promi-
ment); associated with Rubiaceae (A. whiteheadi nences and sharp lateral tubercles; elytra with
Colonnelli, A. tachygonoides Dietz, A. dense semi-erect vestiture of varied brown to
subfasciatus Dietz) ...................... Auleutes (part) black very broadly lanceolate scales; long scutel-
— Rostrum somewhat tapered apically, not conspicu- lar spot on 1st interval black, velvety ................
ously dilated in apical part; rostral sulcus usually ....................................................... Pelenosomus
becoming less deep posteriorly; antenna with — Pronotum without discal prominences; elytra with
club lacking dense short, erect pubescence; as- recumbent, usually sparse to moderately dense
sociated with Vitaceae and Onagraceae ....... 20 pubescence formed mostly by white or metallic-
glossy narrow scales ..................................... 25
20(19). Pronotum and elytra with basal margins straight, no-
ticeably raised at junction and crenulate; 25(24). Dorsal surface with sparse or moderately dense
pronotum usually with a pair of discal promi- vestiture of white scales, lacking metallic-glossy
nences, lateral tubercles acute, well developed, scales, elytra with more or less distinct scutellar
located close to pronotal base; rostral sulcus spot; rostral sulcus extended onto metasternum,
extended onto metasternum (which is sometimes on mesosternum sulcus deep, with abrupt walls;
strongly convex longitudinally), often 1st ventrite elytra with intervals often with sharp granules
is also deeply depressed medially; rostrum mod- (antennal funicle then may be 6-segmented) (A.
erately to strongly curved; associated with epilobii Paykull and several species from Canada
Vitaceae ......................................................... 21 and the U.S. except extreme southwestern and
— Pronotum and elytra with basal margins neither southeastern coastal regions) ...........................
raised nor crenulate; pronotum lacking discal ..................................................... Auleutes (part)
prominences (but present in Pelenosomus, then — Dorsal surface either with a mixture of narrow dark
rostral sulcus limited to mesosternum, very shal- scales with metallic sheen or with narrow broken
low, margined by fine low keels); rostral sulcus bands of narrow white scales (then body chest-
often limited to mesosternum, not extended onto nut-brown), scutellar spot poorly defined; rostral
1st abdominal ventrite; metasternum not strongly sulcus not extended onto metasternum, on me-
convex longitudinally; rostrum moderately sosternum sulcus shallow, its sides gentle, or the
curved to straight; associated with Onagraceae sulcus limited by keels projected behind middle
....................................................................... 22 coxae; elytra with intervals lacking conspicuous
granules ......................................................... 26
21(20). Outer margin of middle and hind (in C. inaequalis
LeConte, also of front) tibiae emarginate, with 26(25). Depression on mesosternum shallow, broad, its mar-
well-developed tarsal grooves; pronotum with gins very finely keel-shaped raised; suture be-
obtuse rounded or elongate prominences, not tween meso- and metasterna not raised; body
 Family 131. Curculionidae · 751

black, legs often brown; elytra with more or less if this is more or less distinctly T-shaped and ob-
distinct scutellar spot and transverse bands, with lique bands on 6-8(9)th intervals also present,
moderately dense dark, narrow, metallic-glossy then bands touch humeral prominence; associ-
scales between bands; rostrum at least weakly ated with Fumariaceae and Papaveraceae .........
curved (A. nebulosus LeConte) .... Auleutes (part) ......................................................... Sirocalodes
— Depression on mesosternum margined by high
lamelliform keels projecting behind middle coxae; 33(30). Elytra with pale pattern, consisting of scutellar spot
body dark chestnut-brown; elytra with narrow with lateral branches oblique or perpendicular to
broken bands of narrow white scales, lacking the suture, and lateral bands on 6-8th intervals,
metallic glossy scales in between; rostrum nearly running to sides of elytra behind humeral promi-
straight (A. inspersus Champion) ....................... n e n c e s ........................................................... 34
..................................................... Auleutes (part) — Elytra without lateral bands, scutellar spot, if dis-
tinct, limited to sutural interval ...................... 35
27(13). Body narrow, strongly elongate, elytra 1.5-1.6 times
as long as wide; legs long, slender; tarsus with 34(33). Scutellar spot with oblique extensions from its
claws simple ...................................... Poophagus transverse part, directed to lateral bands and ei-
— Body less elongate-narrow, elytra less than 1.5 ther connected with them or separated by dark
times as long as wide; legs various, but not long area; lateral bands running to humeral promi-
and slender; tarsus with claws various .......... 28 nences or somewhat behind them; femora with
tooth simple, medium-sized; body size small 2.2-
28(27). Mesosternum shallowly, metasternum deeply de- 2.8 mm; associated with Matricaria perforata
pressed between coxae for reception of rostrum, Mérat and Chrysanthemum leucanthemum L.
the depression slightly extending beyond mar- (Asteraceae) ................................... Microplontus
gins of middle coxae; associated with Urtica dioica — Elytra with wide white cruciform scutellar spot and
L. (Urticaceae) ....................................... Nedyus transverse, more or less oblique bands in middle
— Meso- and metasterna lacking sulcus for reception of 6-9th intervals almost always separated from
of rostrum, not at all depressed medially; associ- the scutellar spot by at least one dark interval;
ated with plants other than Urticaceae ......... 29 femora with tooth on front and middle femora with
truncate apical (facing apex of femur) slope; body
29(28). Antenna with funicle of 6 articles; body size large, size large, 3.7-4.6 mm; associated with
3.5-4.5 mm long, reddish brown with darker un- Cynoglossum officinale L. (Boraginaceae) ..........
derside and rostrum; dorsal surface glossy, with ........................................................... Mogulones
erect or semi-erect long parallel-sided white and
brown scales; basal margins of pronotum and 35(33). Base of pronotum straight or slightly angularly pro-
elytra raised at junction and coarsely crenulate duced posteriorly in the middle, not distinctly
................................................... Trichosirocalus bisinuate; apical margin of pygidium, at least in
— Antennal funicle mostly of 7 articles; if of 6 articles, male, deeply excised in the middle; associated
then body size smaller (less than 3.5 mm), basal with Taraxacum (Asteraceae) or (possibly)
margins of pronotum and elytra not raised at junc- Liliaceae ......................................................... 36
tion and not crenulate, body lacking coarse erect — Base of pronotum more or less distinctly bisinuate;
vestiture, not glossy chestnut-brown ........... 30 apical margin of pygidium entire; if sulcate and
with combs of yellow hairs on sides of excision,
30(29). Tarsus with claw simple ..................................... 31 then male hind tibia with usual small pointed api-
— Tarsus with claw dentate .................................. 33 cal tooth, female middle tibia with small fine api-
cal tooth; associated with Cruciferae ............ 37
31(30). Antenna with funicle of 7 articles; femora with large
tooth, all tibiae in male with large apical tooth, 36(35). Pronotum wide, with strongly rounded sides; ante-
female tibiae simple; associated with Asteraceae, rior margin strongly bent; disc convex, without
mostly Cirsium and Carduus ........ Hadroplontus median sulcus, but with deep prescutellar fovea
— Antenna with funicle of 6 articles; femora with (Fig. 32); lateral tubercles fold-shaped; puncta-
slightly defined tooth or angular prominence; tion dense, uniform and rather fine; elytra with
front tibia of male simple; associated with short scutellar spot of dense white or yellowish
Liliaceae, Scrophulariaceae, Fumariaceae or scales on 1st interval, without oval white scales
Papaveraceae ................................................. 32 on base of other intervals and small pale spot at
the end of basal third of 6th interval (Fig. 32); asso-
32(31). All tibiae in both sexes simple; base of pronotum ciated with Taraxacum officinale Weber
distinctly bisinuate; elytra with pattern consist- (Asteraceae); southeastern Canada and northeast-
ing of scutellar spot with lateral arms stretching ern United States ................................ Glocianus
obliquely back from sutural strip and separated — Pronotum wide at base, but with more or less con-
by dark 5th interval from oblique bands on 6-9th cave sides and sharp, but not fold-shaped lateral
intervals running anteriorly to sides behind hu- tubercles; elytra with small, sometimes indistinct
meral prominences and not touching them (Fig. spot at the end of basal third of 6th interval; asso-
31), sometimes pale pattern reduced to incon- ciated (possibly) with Liliaceae; Yukon Territory
spicuous macula in basal third of 6th interval on .............................................................. Prisistus
dull dark brown background; associated with
Liliaceae and Scrophulariaceae .... Allosirocalus 37(35). Antenna with funicle of 6 articles; anterior margin
— Middle and hind tibiae in male with small apical tooth; of pronotum not raised, sides without any trace
base of pronotum straight; elytra with pattern con- of tubercles, in basal one-half weakly rounded;
sisting only of scutellar spot on sutural interval; base distinctly bisinuate, disk without medial sul-
752 · Family 131. Curculionidae

cus or shortly and narrowly sulcate only at base, Ceuthorrhynchus Gemminger and Harold 1871
strongly convex, densely and finely punctate,
Ceuthorrhynchidius Gemminger and Harold 1871
intervals between punctures with matte reticu-
late microsculpture; body and legs black; associ- Sirocalus Heyden 1906
ated with Nasturtium officinale R. Br. (Cruciferae) Dionorenus Reitter 1913
................................................ Amalorrhynchus Marklissus Reitter 1916
— Antenna with funicle of 6 or 7 articles; if 6-seg-
Heterosirocalus Wagner 1944
mented, then anterior margin of pronotum notice-
ably raised, sides usually with small but distinct Neosirocalus Wagner 1944
tubercles, moderately rounded in basal one-half, Persirocalus Wagner 1944
disk with median sulcus, glossy, more or less Ceuthamiocolus Colonnelli 1983
coarsely punctate; elytral striae rather wide, legs
Nipporhynchus Korotyaev 1996, not Chandler 1934
often pale; associated with various Cruciferae .
....................................................................... 38 Heorhynchus Korotyaev 1999

38(37). Mesosternum not depressed; scales not conceal- Glocianus Reitter 1916, 1 sp., G. punctiger (Sahlberg 1835), generally
ing integument completely, not imbricate (except
distributed; adventive. This species is associated with Taraxacum
in C. opertus Brown) ................... Ceutorhynchus
— Mesosternum between middle coxae moderately officinale Weber (Asteraceae); larvae feed on seeds in flower heads
deeply depressed, sides of the depression (McAvoy et al. 1983).
gentle; scales brownish-grey and imbricate, con- Prenesdus Reitter 1916
cealing integument completely; semi-erect
coarse setae or narrow scales also present ......
....................................................... Rileyonymus Hadroplontus Thomson 1859, 1 sp., H. litura (Fabricius 1775),
Nova Scotia, Ontario, Saskatchewan, Alberta, British Columbia,
CLASSIFICATION OF THE NEARCTIC CEUTORHYNCHINAE South Dakota, Montana, Idaho, Oregon and Washington. In-
troduced for biological control of Carduus and Cirsium thistles
22. Ceutorhynchini Gistel 1856 (Asteraceae) (Peschken and Wilkinson 1981); larvae mine the stems
and crown (Hoffmann 1954).
Allosirocalus Colonnelli 1983, 5 spp., generally distributed in the
central and western United States and southern Canada. Species Microplontus Wagner 1944, 2 spp., M. edentulus (Schultze 1896),
may be associated with Allium (wild onion; Liliaceae) in Texas and Alberta, Saskatchewan, and M. campestris (Gyllenhal 1837), Ontario;
Mimulus and Pedicularis (Scrophulariaceae) in the western United both exotic. Microplontus edentulus has been released locally at two
States. See Hatch (1971) to separate some of the species. sites for biological control of scentless chamomile, Matricaria
perforata Mérat (Asteraceae) (A. S. McClay, pers. comm.).
Amalorrhynchus Reitter 1913, 1 sp., A. melanarius (Stephens 1831), Microplontus campestris has only recently been documented as
Quebec, Connecticut, Massachusetts and West Virginia; adven- present in North America; it is associated with Chrysanthemum
tive. This species is associated with Nasturtium officinale R. Br. leucanthemum L. (Asteraceae).
(watercress; Cruciferae).
Mogulones Reitter 1916, 1 sp., M. cruciger (Herbst 1784), British
Amalus Schoenherr 1825, 1 sp., A. scortillum (Herbst 1795), gener- Columbia and Alberta. This species has been introduced for the
ally distributed in the northern United States and Canada; adven- biological control of Cynoglossum officinale L. (hound’s-tongue;
tive. This species is associated with Polygonum (Polygonaceae); Boraginaceae) (DeClerk-Floate and Schwarzländer in press).
larvae feed in the crown (Hoffmann 1954). Boraginobius Wagner 1944
Leptocaryurgus Gistel 1856
Nedyus Schoenherr 1825, 2 spp., generally distributed in the east-
Ceutorhynchus Germar 1824, 68 spp., generally distributed; some ern United States west to Texas and Canada west to Alberta.
adventive and of pest status. Species are associated with Cruciferae; Species are associated with Urtica dioica L. (nettle; Urticaceae)
larvae often mine in collars of roots or stems (Anderson 1993b). (Blatchley and Leng 1916). See Blatchley and Leng (1916) to sepa-
See Dietz (1896), Blatchley and Leng (1916) and Hatch (1971) to rate the species.
separate some of the species. An unpublished 1963 Ph.D. thesis Cidnorhinus Thomson 1859
by Rudolph Scheibner from Michigan State University allows for
the identification of most species. The genus needs revision. Phrydiuchus Gozis 1885, 2 spp., P. tau Warner and P. spilmani Warner
Calosirus Thompson is given distinct generic status by Wibmer (latter may not be established), California, Oregon, Washington;
and O’Brien (1989) and Alonso-Zarazaga and Lyal (1999); 8 spe- adventive. Introduced for the biological control of Salvia aethiops
cies could be placed provisionally in this taxon in North America. L. (Mediterranean sage; Labiatae) (Warner 1969). See Warner (1969)
Falciger Dejean 1821 to separate the species.
Ceuthorhynchus Schoenherr 1837
Ceuthorhynchidius Jacquelin du Val 1855
Calosirus Thompson 1859
 Family 131. Curculionidae · 753

Poophagus Schoenherr 1837, 1 sp., P. sisymbrii (Fabricius 1776), Dietzella Champion 1907, 2 spp., D. zimmermanni (Gyllenhal 1837),
Quebec; adventive. This species is associated with Nasturtium generally distributed in the eastern United States, southern Canada
(Cruciferae); larvae mine stems and roots (Hoffmann 1954). and western United States, and D. sextuberculata (Boheman 1845),
Poephagus Gistel 1856 Colorado. Species are associated with Epilobium (Onagraceae).
Acnemiscelis Desbrochers 1896
Orchestomerus Dietz 1896, 3 spp., generally distributed in the east-
Prisistus Reitter 1916, 1 sp., P. olgae Korotyaev 1988, Yukon Terri- ern United States, Texas and Arizona. At least one species, O.
tory. This species may be associated with Liliaceae. whiteheadi Colonnelli 1991, is associated with Vitis (wild grape;
Austroceutorhynchus Korotyaev 1980 (valid subgenus) Vitaceae) in Arizona.
Ranunculiphilus Dieckmann 1970 (valid subgenus) Platymeristes Dietz 1896
Svetlaniolus Korotyaev 1997 (valid subgenus)
Pelenosomus Dietz 1896, 1 sp., P. cristatus Dietz 1896, southeastern
Rileyonymus Dietz 1896, 1 sp., R. relictus Dietz 1896, Arizona, United States.
California. This genus is questionably distinct from Ceutorhynchus.
Perigaster Dietz 1896, 4 spp., generally distributed in the eastern
Sirocalodes Voss 1958, 3 spp., S. tescorum (Fall 1907), S. sericans United States, southern Canada and western United States. Spe-
(LeConte 1876), and S. siculus (Dietz 1896), generally distributed cies are associated with Ludwigia (Onagraceae) (Anderson 1993a).
in the western and southern United States and Manitoba. See Buchanan (1931) to separate the species.
Sirocalodes tescorum (and S. wickhami (Champion 1907) from
Mexico) have been associated with Argemone (Papaveraceae) and Perigasteromimus Colonnelli 1999, 1 sp., P. tetracanthus (Champion
S. siculus with Corydalis (Fumariaceae). 1907), Florida. This species is associated with Ludwigia spp.
(Onagraceae) (C. W. O’Brien, pers. comm.).
Trichosirocalus Colonnelli 1979, 1 sp., T. horridus (Panzer 1801),
Virginia. Introduced for the biological control of Carduus (thistles; 24. Hypurini Schultze 1902
Asteraceae); larvae mine in the crown and stem (Trumble and
Kok 1979). Hypurus Rey 1882, 1 sp., H. bertrandi (Perris 1852), California and
Florida; adventive. This species is associated with Portulaca oleracea
23. Cnemogonini Colonnelli 1979 L. (Portulacaceae) in Europe but other Portulacaceae are suitable
hosts (Zimmerman 1957; Anderson 1993a).
Acanthoscelidius Hustache 1930, 13 spp., generally distributed.
Species are associated with Oenothera, Gaura and perhaps other 25. Mononychini LeConte 1876
Onagraceae (Anderson 1993b). See Dietz (1896) to separate most
of the species. The genus needs revision and its relationships Mononychus Germar 1824, 1 sp., M. vulpeculus (Fabricius 1801),
with Auleutes reassessed. generally distributed in the eastern United States and southern
Acanthoscelis Dietz 1896; not Dejean 1825 Canada. This species is associated with Iris versicolor L. (Iridaceae);
Acantharthrus Marshall 1939 larvae are in seed pods (Blatchley and Leng 1916).

Auleutes Dietz 1896, 12 spp., generally distributed. Species are 26. Phytobiini Gistel 1848
associated with Ludwigia, Calylophus, Oenothera, Gaura and per-
haps other Onagraceae (Blatchley and Leng 1916; Anderson [Eubrychius Thomson 1859, 1 sp., E. velutus (Beck 1817). This
1993b). Auleutes donaldi Colonnelli 1991 has been associated with species has been recorded from the eastern United States west
Bouvardia glaberrima Engelm. (Rubiaceae). See Dietz (1896) to sepa- through the north to the western United States and British Co-
rate most of the species. The genus needs revision and its rela- lumbia. It does not occur in North America; all records are
tionships with Acanthoscelidius and neotropical taxa placed as misidentifications of Euhrychiopsis lecontei (Dietz 1896), see
Auleutes reassessed. The key presented here recognizes three dis- Tamayo et al. 1999.]
tinct groups of Auleutes likely warranting separate generic status.
Euhrychiopsis Dietz 1896, 1 sp., E. lecontei (Dietz 1896), eastern to
Cnemogonus LeConte 1876, 1 sp., C. lecontei Dietz 1896, generally central United States and western Canada. This species is associ-
distributed in the northern United States and Canada. This spe- ated with Potamogeton (Potamogetonaceae) and Myriophyllum
cies may be associated with Onagraceae. (Haloragaceae) (Kissinger 1964; Hatch 1971). See Tamayo et al.
(1999) for information about this species. All records of Eubrychius
Craponius LeConte 1876, 1 sp., C. inaequalis (Say 1831), generally velutus (Beck 1817) are misidentifications of E. lecontei (Tamayo et
distributed in the eastern and central United States and southern al. 1999).
Canada. This species is associated with Vitis (grapes; Vitaceae);
larvae feed within fruits on seeds (Blatchley and Leng 1916).
754 · Family 131. Curculionidae

Neophytobius Wagner 1936, 1 sp., N. cavifrons (LeConte 1876), gen- and Alaska. This species is associated with Populus (Salicaceae)
erally distributed in the western United States and Canada. This (Anderson 1997).
species is associated with Polygonum (Polygonaceae). Rhytidosomus Schoenherr 1837
Nemophytobius Voss 1952 Rhytidosoma Agassiz 1846
Oligodites Gistel 1856
Parenthis Dietz, 1896, 1 sp., P. vestitus Dietz 1896, southeastern Rhytidosomus Gemminger and Harold 1871; not Schoenherr
United States. This genus is considered as a junior synonym of 1837
Phytobius Schoenherr by Alonso-Zarazaga and Lyal (1999). Scleropteridius Otto 1897 (valid subgenus)
Prorutidosoma Korotyaev 1999 (valid subgenus)
Pelenomus Thomson 1859, 13 spp., generally distributed in the Victorinus Korotyaev 1999 (valid subgenus)
more northerly United States and Canada. Most Palearctic species
are associated with Polygonum (Polygonaceae) (Hoffmann 1954). VIII. Conoderinae Schoenherr 1833
See Dietz (1896), Blatchley and Leng (1916) and Hatch (1971) to
separate some of the species. The genus needs revision. by Henry A. Hespenheide
Pachyrhinus Stephens 1829; not Schoenherr 1825
Phytobius Dejean 1835; not Schoenherr 1833 The Conoderinae have been defined by the combination of a
Mecopeltus Dietz 1896 prosternal channel for the reception of the rostrum, large ap-
Paraphytobius Wagner 1936 proximate eyes, and the absence of postocular lobes on the ante-
rolateral margin of the pronotum. They are usually placed be-
Phytobius Schoenherr 1833, 1 sp., P. leucogaster (Marsham 1802), tween the Cryptorhynchinae and the Ceutorhynchinae and are
generally distributed in the eastern United States west through probably more closely related to the former. Adults are typically
the north to the western United States. This species is associated diurnal and very wary and active fliers. There is considerable struc-
with Myriophyllum (Haloragaceae); larvae feed externally on flow- tural diversity within the subfamily, even among the North Ameri-
ers (Buckingham and Bennett 1981). can forms, but the group is much more diverse in the Neotropi-
Hydaticus Schoenherr 1825; not Leach 1817 cal Region.
Litodactylus Redtenbacher 1849 Most conoderine larvae are borers of wood or herbaceous
stems although a few feed on seeds. The genus Tachygonus, pro-
Rhinoncus Schoenherr 1825, 7 spp., generally distributed; three visionally placed here as a highly derived subgroup, has larvae
species adventive. Species are associated with Polygonum which mine leaves. A few North American species such as
(Polygonaceae) (Hoebeke and Whitehead 1980). See Hoebeke and Cylindrocopturus adspersus (LeConte 1876), the sunflower stem
Whitehead (1980) to separate six of the species. Rhinoncus borer, are economically significant pests.
perpendicularis (Reiche 1797) recently has been collected in Ontario.
Cryptorhis Billberg 1820 KEY TO THE NEARCTIC GENERA OF CONODERINAE
Campylirhynchus Dejean 1821
Camplirhynchus Gistl 1834 1. Antenna straight, not geniculate; hind coxae widely
separated by a distance 4 or 5 times greater than
Campylorhynchus Agassiz 1846; not Spix 1824
width of a coxa; hind femur long (much longer
than front or middle femora), spinose ventrally
(Fig. 47); body form broadly ovate, flattened; with
27. Scleropterini Schultze 1902 appressed scales and tufts of erect setae (Fig.
47) ................................................... Tachygonus
— Antenna geniculate; hind coxae narrowly separated
Acallodes LeConte 1876, 3 spp., generally distributed in the east- by a distance less than twice width of a coxa;
ern United States and southern Canada. One species is associated hind femur short (subequal in length to front or
with Lysimachia terrestris L. (Primulaceae) (Blatchley and Leng 1916). middle femora), simple or with single ventral tooth;
body form elongate ovate, subtriangular in lat-
See Blatchley and Leng (1916) to separate the species.
eral form, not flattened (Figs. 39-44); vestiture
various, mostly with only appressed scales or
Asperosoma Korotyaev 1999, 1 sp., A. echinatum (Fall 1917), scattered setae ................................................ 2
Manitoba. This odd species is associated with Heuchera richardsoni
2(1). Pygydium exposed dorsally (Fig. 41); body form
R. Br. (Saxifragiaceae) (Fall 1917).
more or less flattened dorsally and ventrally (Fig.
42); body length greater than 6.0 mm ...............
Homorosoma Frivaldszky 1894, 1 sp., H. sulcipenne (LeConte 1876), ......................................................... Peltophorus
generally distributed. This species is associated with Polygonum — Pygydium covered by elytra (Figs. 39, 43); body
form with ventral or dorsal surface convex (Figs.
(Polygonaceae).
40, 44); body length smaller than 6.0 mm ........ 3

Rutidosoma Stephens 1831, 1 sp., R. decipiens (LeConte 1876), gen-

erally distributed in the eastern and western United States, Canada
 Family 131. Curculionidae · 755

39 40 41 42 45

46
47
43 44
FIGURES 39.131-47.131. Conoderinae 39-40. Psomus armatus (Dietz), habitus, 39. Dorsal; 40. Lateral. 41-42. Peltophorus polymitus seminiveus
(LeConte), habitus, 41. Dorsal; 42. Lateral. 43-44. Cylindrocopturus adspersus (LeConte), habitus, 43. Dorsal; 44. Lateral. 45-46. Cylindrocopturus
adspersus (LeConte), head, 45. Anterior view; 46. Lateral view. 47. Tachygonus lecontei Gyllenhal (Conoderinae), dorsal habitus (left side show
scale pattern, right side shows color).

3(2). Abdomen with ventrites in lateral view in same — Antenna with article 2 of funicle about twice length
plane, horizontal; elytra in lateral view more or of article 1; femora with large ventral tooth ......
less continuously convex (Fig. 40) ................. 4 ............................................................. Copturus
— Abdomen with ventrites in lateral view ascended;
elytra in lateral view flattened at base (Fig. 44) . 7(5). Femora carinate on outer face and with ventral tooth
......................................................................... 5 ............................................................ Lechriops
— Femora not carinate on outer face, lacking ventral
4(3). Elytra with distinct setae; femora each with ventral tooth ................................................................. 8
tooth; body length greater than 3.0 mm ...........
.............................................................. Acoptus 8(7). Metasternum with anterior margin excavated for re-
— Elytra lacking distinct setae; femora simple, lacking ception of rostrum; body length less than 3.0 mm
ventral tooth; body length less than 2.0 mm ..... ......................................................... Eulechriops
............................................................... Psomus — Metasternum with anterior margin simple, not modi-
fied for reception of rostrum; body length greater
5(3). Mesosternum simple, unmodified, apex of rostrum than 3.0 mm ......................... Cylindrocopturinus
free ................................................................... 6
— Mesosternum excavated, with lateral margins cari-
CLASSIFICATION OF THE NEARCTIC CONODERINAE
nate; anterior margin of metasternum excavated
to receive the tip of the rostrum or not ........... 7
28. Lechriopini Lacordaire 1866
6(5). Antenna with articles 1 and 2 of funicle subequal in
length; femora simple, lacking ventral tooth .....
Acoptus LeConte 1876, 1 sp., A. suturalis LeConte 1876, generally
................................................ Cylindrocopturus
distributed in eastern United States and southern Canada. This
756 · Family 131. Curculionidae

species is associated with dead wood of beech trees, also with Tachygonus Guérin-Méneville 1833, 5 spp., generally distributed
hop-hornbeam and hickory (Blatchley and Leng 1916). in eastern United States and southeastern Canada, west to Texas,
Arizona, New Mexico and Colorado. Species are associated with
Copturus Schoenherr 1825, 1 sp., C. floridanus (Fall 1906), Florida. Quercus (Fagaceae), Ulmus (Ulmaceae), Robinia (Fabaceae), Coursetia
This species is associated with Swietenia mahagoni (L.) Jacq. (Fabaceae)) and Berchemia (Rhamnaceae); larvae mine leaves
(Meliaceae); larvae bore under bark of living branches (Anderson (Hespenheide 1992). See Hespenheide (1992) to separate the spe-
1993a). cies.
Zurus Heller 1895; not Amyot 1846 Tachygonus Schoenherr 1833; not Guérin-Méneville 1833
Neozurus O’Brien and Wibmer 1982 Tachyopus Zimmermann 1840

Cylindrocopturinus Sleeper 1963, 1 sp., C. pictus (Schaeffer 1908), IX. Cossoninae Schoenherr 1825
Arizona. This species is associated with Phoradendron (mistletoe;
Viscaceae) (Anderson 1994). by Robert S. Anderson

Eulechriops Faust 1896, 2 spp., generally distributed in the eastern Cossonine weevils are easily recognized by the large, hook-like
United States and Arizona. Species are associated with Quercus tooth at the apex of the hind tibia and the lack of an apical comb
(oak; Fagaceae). of setae. They are usually black or brown, lack scales but have
Zygomicrus Casey 1897 appressed or erect hairs, and are generally long, slender and dor-
soventrally compressed. Most species are associated with dead
Lechriops Schoenherr 1825, 4 spp., generally distributed. See plant material of some sort, usually of woody angiosperms (where
Blatchley and Leng (1916) to separate some of the species. they live under bark), but some are also found in dead fern fronds,
Gelus Casey 1897 palm fronds, agave leaves, yucca stalks, etc. A number of taxa are
found on sandy beaches in association with driftwood. Species
Psomus Casey 1892, 1 sp., P. armatus (Dietz 1891), northeastern of Acamptus and at least some Pseudopentarthrum are found in tree
United States and southern Canada. This species is associated holes or rotten hollowed out trees. Most of the genera in North
with Fraxinus americanus L. (white ash; Oleacae) (Blatchley and America are represented by only one or a few species. Some spe-
Leng 1916). cies, such as those in the genus Hexarthrum, are difficult to sepa-
rate from Scolytinae.
29. Zygopini Lacordaire 1866
KEY TO THE NEARCTIC GENERA OF COSSONINAE
Cylindrocopturus Heller 1895, 29 spp., generally distributed. Many
species are associated with various Asteraceae and some with 1. Prosternum with distinct channel for reception of
rostrum when in repose; pronotum and elytra with
Pinaceae. See Casey (1897), Fall (1906) and Hatch (1971) to sepa-
dense, erect or suberect, broad scales ............
rate some of the species. The genus needs revision. ........................................................... Acamptus
Paratimorus Heller 1895 — Prosternum simple, smooth, lacking channel for re-
Gyrotus Casey 1897 ception of rostrum; pronotum and elytra with
vestiture various, if erect or suberect, composed
Copturodes Casey 1897
of finer, hair-like scales or sparse, scattered broad
scales ............................................................... 2
Peltophorus Schoenherr 1845, 3 spp., Arizona, New Mexico and
Texas. Species are associated with Agave (Amaryllidaceae); larvae 2(1). Eyes obviously located on base of rostrum, head
distinctly constricted and globular behind eyes
mine the stalks or in the seeds (Kissinger 1964). See Sleeper (1963)
(Fig. 51); Florida ................................................ 3
to separate the species. — Eyes located on head or at junction of head and
Apatorhynchus Desbrochers 1891 rostrum, head not distinctly constricted dorsally
Opalocetus Desbrochers 1910 behind eyes (Fig. 52); various locations .......... 4

3(2). Eye reduced in size to less than 10 facets (Fig. 51);

30. Tachygonini Lacordaire 1866 antenna with basal article of club subglabrous,
glossy; rostrum in lateral view with ventral mar-
Alonso-Zarazaga and Lyal (1999) place Tachygonina as a subtribe gin straight (Fig. 51) ............................. Paralicus
— Eye with more than 10 facets; antenna with basal
within Curculioninae; Rhamphini. This distictive genus is placed
article of club setose, not glossy; rostrum in lat-
here as a tribe within Conoderinae based upon the form of the eral view with ventral margin curved ventrally
unci at the apex of the tibiae, large eyes and form of vestiture. towards apex .................................... Dryotribus
Similarity in the pectinate form of the scales suggests a relation-
4(2). Eyes markedly reduced in size to 3 or 4 facets ....
ship with the Neotropical genera Philinna Champion 1906 and
..................................................... Amaurorhinus
Philides Champion 1906. — Eye with more than 10 facets .............................. 5
 Family 131. Curculionidae · 757

50 52 53
48 49 51

58 56 55 54
57
FIGURES 48.131-58.131. Cossoninae. 48-50. Dorsal habitus, 48. Cossonus piniphilus Boheman; 49. Macroscytalus chisosensis (O’Brien); 50.
Stenotrupis acicula Wollaston. 51-52. Lateral view of head, 51. Paralicus minyops O’Brien; 52. Micromimus minimus (Boheman). 53-56. Dorsal view
of head, 53. Aphanommata tenuis (Casey); 54. Tomolips quercicola (Boheman); 55. Pseudopentarthrum robustum Casey; 56. Rhyncolus brunneus
Mannerheim. 57-58. Hind tibia, 57. Cossonus piniphilus Boheman; 58. Elassoptes marinus Horn.

5(4). Elytra with intervals with obvious vestiture of elon- 10(9). Elytra three to four times as long as pronotum (Fig.
gate-narrow, recurved hair-like scales, alternate 50); pronotum about as wide as long (Fig. 50);
intervals also with row of suberect, broad, trun- antenna with scape extended slightly beyond
cate scales; eyes situated low on head, in lateral hind margin of eye; Florida .............. Stenotrupis
view with the ventral margin of rostrum directed — Elytra two to three times as long as pronotum (Fig.
to middle of eye ................................ Himatium 49); pronotum longer than wide (Fig. 49); antenna
— Elytra with intervals with obvious vestiture absent, with scape extended to about middle of eye;
or with obvious vestiture of elongate, fine, hair- southwestern Texas ................... Macroscytalus
like setae only, no broad scales present; eyes
situated higher on head, in lateral view with ven- 11(8). Rostrum distinctly tapered apically in dorsal view
tral margin of rostrum directed to lower one-half (Fig. 54); antenna with club truncate at apex ....
of eye or obviously below eye ........................ 6 ............................................................. Tomolips
— Rostrum more or less subparallel or slightly ex-
6(5). Antenna with funicle of 5 or 6 articles ................ 7 panded towards apex in dorsal view (Fig. 55); an-
— Antenna with funicle of 7 articles ..................... 13 tenna with club rounded at apex ................... 12

7(6). Antenna with funicle of 6 articles .... Hexarthrum 12(11). Antenna with club not expanded, article 1 of fu-
— Antenna with funicle of 5 articles ....................... 8 nicle about as wide as club (Fig. 55); widespread
........................................... Pseudopentarthrum
8(7). Dorsal vestiture of obvious, long, very fine hair-like — Antenna with club expanded, article 1 of funicle
setae, each at least as long as a strial puncture or distinctly narrower than club; Quebec ..............
longer ............................................................... 9 ....................................................... Pentarthrum
— Dorsal vestiture of at most indistinct, short, fine
hair-like setae, each shorter than a strial punc- 13(6). Dorsal vestiture of obvious, long, fine, hair-like se-
ture, or obvious vestiture absent .................. 11 tae, each at least as long as a strial puncture or
longer ............................................................. 14
9(8). Body size greater than 2.0 mm, black in color; body — Dorsal vestiture of at most indistinct, short, fine
form subcylindrical, more or less as wide as high; hair-like setae, each shorter than a strial punc-
head not constricted behind eyes ................... ture, or obvious vestiture absent .................. 16
........................................................ Nyssonotus
— Body size less than 2.0 mm, pale or dark brown in 14(13). Front coxae very narrowly separated by much less
color; body form markedly dorsoventrally com- than one-half width of a coxa; elytra with vestiture
pressed, much wider than high; head constricted of long, erect, fine, hair-like setae, each much
behind eyes, area posterior to constriction longer than a strial puncture .............. Pselactus
impunctate and glossy ................................... 10 — Front coxae more widely separated by at least one-
half width of a coxa or more; elytra with vestiture
758 · Family 131. Curculionidae

of shorter hair-like or scale-like setae, each about markedly dorsoventrally compressed; body size
as long as a strial puncture ............................ 15 less than 2.0 mm; Florida ....................... Proeces
— Front coxae situated close to posterior margin of
15(14). Front coxae separated by more or less width of a prosternum, separated from margin by slightly
coxa; elytra with vestiture of erect, stout, scale- less than the length of a coxa; body form
like setae; eyes situated at junction between ros- subcylindrical; body size various; various loca-
trum and head .................................... Apotrepus tions, including Florida .................................. 23
— Front coxae separated by about one-half width of a
coxa; elytra with vestiture of appressed, hair-like 23(22). Middle and hind femora very short, distinctly ex-
setae; eyes situated on head adjacent to base of panded apically and subtriangular in shape,
rostrum .......................................... Carphonotus length about twice width at apex; body size small,
less than 2.0 mm; body color testaceous ..........
16(13). Hind tibia expanded somewhat laterally at apex, ....................................................... Stenomimus
hook-like tooth at outer angle large and stout, — Middle and hind femora moderate in length, slightly
tooth at inner angle, stout, spatulate, much longer expanded apically but more elongate in shape,
than tarsal claw (Fig. 58) ................... Elassoptes length greater than twice width at apex; body
— Hind tibia simple, not expanded laterally at apex, size various, less than 5.0 mm; body color black,
hook-like tooth at outer angle various (Fig. 57), dark brown or testaceous ............... Caulophilus
tooth at inner angle (if present) not spatulate,
smaller than tarsal claw .................................. 17 24(18). Front coxae very narrowly separated by less than
one-half width of a coxa; posterior and anterior
17(16). Rostrum in dorsal view with apical one-half more or prosternal processes acuminate or subacuminate
less abruptly dilated beyond point of antennal apically ........................................................... 25
insertion, wider than basal one-half, ventral mar- — Front coxae moderately to widely separated by more
gin of scrobe visible in dorsal view (Fig. 48); an- than one-half width of a coxa; posterior and ante-
tenna inserted beyond midlength of rostrum .... rior prosternal processes truncate apically .. 27
............................................................ Cossonus
— Rostrum subequal in width throughout length (Fig. 25(24). Elytra at base with intervals 2 to 4 swollen and
56) or tapered towards apex (Fig. 53), scrobe not crenulate or minutely dentate; elytral declivity
visible in dorsal view; antenna with point of in- with numerous small denticles; rostrum very short,
sertion in basal one-half of rostrum ............... 18 wider than long, in dorsal view with lateral mar-
gins subparallel ............................... Stenoscelis
18(17). Rostrum in dorsal view at least twice as long as — Elytra at base with intervals smooth, not swollen,
width of frons between eyes ......................... 19 dentate or crenulate; elytral declivity smooth,
— Rostrum in dorsal view less than twice as long as lacking denticles; rostrum as long as wide to
width of frons between eyes (Figs. 53, 56) ... 24 slightly longer than wide; rostrum with lateral
margins (excluding scrobes if visible in dorsal
19(18). Eyes large and elongate-oval in shape, height more view) convergent apically ............................. 26
or less twice maximum width (Fig. 52) ...............
........................................................ Micromimus 26(25). Eyes at most only slightly visible in dorsal view,
— Eyes moderate in size, round or sub-oval in shape, flat, situated low on head (Fig. 53), in lateral view
height much less than twice width ................ 20 with ventral margin of rostrum directed towards
middle or lower one-half of eye ........................
20(19). Metepisternum wide throughout length, subequal ................................................... Aphanommata
in width to width of antennal club, with 2 or 3 — Eyes clearly visible in dorsal view, slightly convex,
rows of large, distinct, deep punctures; rostrum situated higher on head (Fig. 56), in lateral view
of female with point of antennal insertion basal, with ventral margin of rostrum directed below eye
rostrum long, glabrous, cylindrical and slender, (includes Phloeophagus minor, P. californicus) ..
rostrum of male with point of antennal insertion .................................................. Rhyncolus (part)
at basal one-third, flat and deeply punctate dor-
sally, more quadrate in cross-section ... Mesites 27(24). Rostrum very short, wider than long; pronotum about
— Metepisternum narrow throughout length, width as wide as long ......................... Rhyncolus (part)
much less than width of antennal club, with at — Rostrum longer than wide; pronotum longer than
most one indistinct row of shallow punctures; wide ................................................................ 28
rostrum various in form, similar in both female and
male ................................................................ 21 28(27). Rostrum in dorsal view tapered towards apex, with
apex narrower than width of rostrum at position
21(20). Front coxae very narrowly separated by much less of antennal insertion ..................... Macrancylus
than one-half width of a coxa, anterior and poste- — Rostrum in dorsal view subparallel or slightly wider
rior prosternal processes acuminate apically ... towards apex, with apex subequal in width or
..................................................... Phloeophagus slightly wider than width of rostrum at position of
— Front coxae moderately to widely separated by at antennal insertion .......................................... 29
least one-half width of a coxa, anterior and pos-
terior prosternal processes broadly truncate 29(28). Antenna with apex of scape extended well beyond
apically ........................................................... 22 the hind margin of the eye; eyes flat, or very
slightly convex, slightly visible in dorsal view .
22(21). Front coxae situated distant from posterior margin .................................................. Macrorhyncolus
of prosternum, separated from margin by obvi-
ously more than the length of a coxa; body form
 Family 131. Curculionidae · 759

— Antenna with apex of scape extended at most to 33. Dryotribini LeConte 1876
the posterior margin of the eye; eyes convex,
obviously visible in dorsal view .................... 30
Amaurorhinus Fairmaire 1860, 1 sp., A. beckwickianus (Wollaston
30(29). Antenna with scape extended to the hind margin of 1860), South Carolina; adventive.
the eye; head slightly constricted behind eyes; Mesoxenus Wollaston 1861
elytral declivity lacking fine setae; Florida ........
Mazagranus Pic 1905 (valid subgenus)
........................................................ Stenancylus
— Antenna with scape extended to the midlength of
the eye; head not constricted behind eyes; Caulophilus Wollaston 1854, 4 spp., generally distributed in the
elytral declivity with numerous fine setae; Cali- eastern United States west to Texas, also California; one adven-
fornia .......................................... Trichacorynus
tive species, C. oryzae (Gyllenhal 1838). Native species are associ-
ated with various species of dead trees and grapevines (Blatchley
CLASSIFICATION OF THE NEARCTIC COSSONINAE and Leng 1916). Caulophilus oryzae is the ‘broad-nosed grain wee-
vil’ and is found in stored products as well as in avocado seeds
31. Cossonini Schoenherr 1825 and fruits (Anderson 1993a). See Blatchley and Leng (1916) to
separate some of the species.
Cossonus Clairville 1798, 20 spp., generally distributed. Adults are Allomimus LeConte 1876
found under bark of various tree species, mostly conifers but Tytthomimus Champion 1909
also hardwoods. See Van Dyke (1915, 1916), Blatchley and Leng
(1916) and Hatch (1971) to separate the species. The genus needs Dryotribus Horn 1873, 1 sp., D. mimeticus Horn 1873; Florida and
revision. South Carolina. This species is associated with old boards and
Borophloeus Wollaston 1873 driftwood washed up along the coast (Anderson 1993a).
Isotrogus Wollaston 1873 Thalattodora Perkins 1900
Hyponotus Wollaston 1873 Pentacotaster Chûjô and Voss 1960
Heterophasis Wollaston 1873 (valid subgenus)
Drepocossonus Voss 1939 (valid subgenus) Micromimus Wollaston 1873, 2 spp., Florida. Adults have been
Caenocossonus Voss 1955 (valid subgenus) collected under bark of various trees, especially Bursera simaruba
Odontocossonus Voss 1956 (valid subgenus) (L.) Sarg. (Burseraceae) (Anderson 1993a).
Otiorcossonus Voss 1956 (valid subgenus)
Paralicus O’Brien 1984, 1 sp., P. minyops O’Brien 1984, southern
[Dynatopechus Marshall 1931, 1 sp., D. aureopilosus (Fairmaire 1849), Florida. Adults are found under driftwood and in litter along
intercepted in quarantine; California, Oregon and Washington. beaches (Anderson 1993a).
Not established in North America.]
Stenomimus Wollaston 1873, 1 sp., S. pallidus (Boheman 1845),
Mesites Schoenherr 1838, 2 spp., likely adventive; eastern United generally distributed in the eastern United States. Larvae have
States. Species are associated with driftwood on Atlantic and Gulf been found under bark of Juglans nigra L. (black walnut;
Coast beaches. See Blatchley and Leng (1916) to separate the spe- Juglandaceae) (Anderson 1952).
cies.
Odontomesites Wollaston 1873 (valid subgenus) 34. Onycholipini Wollaston 1873
Stenotrupis Wollaston 1873, 1 sp., S. acicula Wollaston 1873, Florida. Hexarthrum Wollaston 1860, 3 spp., generally distributed in the
This species is associated with dead fronds of Thrinax parviflora eastern United States and southern Canada, disjunct to British
Sw. (Arecaceae) (Anderson 1993a). Columbia and Idaho; one adventive. Adults occur in woodwork
Diodimorpha Broun 1883 of buildings (Blatchley and Leng 1916). See Brown (1966a) to
Pseudaphioda Voss 1956 (valid subgenus) separate the species.
32. Acamptini LeConte 1876 Pselactus Broun 1886, 1 sp., P. spadix (Herbst 1795), eastern United
States and California; adventive. This species is associated with
Acamptus LeConte 1876, 3 spp., generally distributed in the east- driftwood on coastal beaches (Blatchley and Leng 1916).
ern United States and southern Canada, west to Texas and Ari- Codiosoma Bedel 1885; not Kirby 1874
zona. Species are associated with dead limbs and injured spots or Phloeophagia Aurivillius 1924
areas of dead rotten wood such as tree holes or hollowed out
trunks of various trees (Kissinger 1964). See Casey (1895) and Pseudopentarthrum Wollaston 1873, 11 spp., generally distributed
Sleeper (1954b) to separate the species. The genus needs revision. in the eastern United States west to Texas and Arizona. Species
Pseudacamptus Champion 1909 are associated with dead limbs and injured spots or areas of dead
Glyphostethus Marshall 1921 rotten wood such as tree holes or hollowed out trunks of vari-
760 · Family 131. Curculionidae

ous trees (Kissinger 1964; Anderson 1993a). See Blatchley and Macrancyloides Champion 1909
Leng (1916), Blatchley 1922, 1925, 1928) and Sleeper (1954b) to Oocorynus Champion 1909
separate the species. The genus needs revision. Brachytemnoides Folwaczny 1973
Phloeophagomorphus Wollaston 1873
Pentarthrinus Casey 1892 Apotrepus Casey 1892, 1 sp., A. densicollis Casey 1892, Arizona.
Micropentarthrum Champion 1909 This species is associated with scar tissue on wounds of saguaro
Neopentarthrum Mutchler 1925 cactus (Cactaceae) (Kissinger 1964).
Stenotylus Marshall 1933
Carphonotus Casey 1892, 1 sp., C. testaceus Casey 1892, generally
Stenoscelis Wollaston 1861, 2 spp., generally distributed in the distributed in the northern United States and across Canada.
eastern United States and southern Canada. Species are associated Adults have been associated with spruce (Blatchley and Leng 1916).
with the dead wood of various trees (Kissinger 1964). See
Buchanan (1948) to separate the species. Elassoptes Horn 1873, 1 sp., E. marinus Horn 1873, western United
Dendroctonomorphus Wollaston 1873 States and Canada. Adults are associated with driftwood on
Astenoscelis Hustache 1956 (valid subgenus) beaches (Kissinger 1964).
Hexastenoscelis Voss 1964 (valid subgenus)
Himatium Wollaston 1873, 2 spp., generally distributed in the
Trichacorynus Blatchley 1916, 2 spp., Indiana, New Jersey, Penn- eastern United States and southern Canada. Adults have been
sylvania and California. Adults have been reared from Yucca stalks reared from dead branches of Acer saccharum L. (Aceraceae) and
(Liliaceae) in California. See Sleeper (1957b) to separate the spe- have been collected in leaf litter (Anderson 1993a). Adults have
cies. also been associated with the galleries of Ips bark beetles under
pine bark. See Blatchley and Leng (1916) to separate the species.
35. Pentarthrini Lacordaire 1866 Pholidonotus Wollaston 1873
Choerodemas Faust 1898 (valid subgenus)
Macroscytalus Broun 1881, 1 sp., M. chisosensis (O’Brien 1973), Himatinum Cockerell 1906
southwestern Texas. This species is associated with the dead, dry Macrohimatium Konishi 1962
leaves of Agave havardiana Trel. (Amaryllidaceae) (O’Brien 1973).
Rhinanisus Broun 1883 Macrancylus LeConte 1876, 1 spp., M. linearis LeConte 1876, south-
Baeorhopalus Broun 1883 eastern United States west to Texas, Adults are associated with
driftwood on coastal beaches (Blatchley and Leng 1916).
Pentarthrum Wollaston 1854, 1 sp., P. huttoni Wollaston 1854, Haloxenus Perkins 1900
Quebec; adventive. Adults have been found in floor boards of
houses (Warner 1952). Macrorhyncolus Wollaston 1873, 1 sp., M. littoralis (Broun 1880),
Attarus Broun 1909 California; adventive. Adults are associated with driftwood on
Belka Broun 1909 coastal beaches.
Gaurocryphus Broun 1909
Trachyglyphus Broun 1909 Nyssonotus Casey 1892, 1 sp., N. seriatus Casey 1892, Arizona,
California and Texas. This species is associated with Yucca
36. Proecini Voss 1956 (Amaryllidaceae); larvae are in stalks (Anderson 1952).

Proeces Schoenherr 1838, 1 sp., P. depressus (Boheman 1838), Florida. Rhyncolus Germar 1817, 15 spp., generally distributed in the east-
Adults have been collected in curled leaf sheaths of Roystonea ern United States west to Texas and north to Canada, then across
elata Bartr. (F. Harper) (royal palm; Arecaceae) (Anderson 1993a). Canada and south into the western United States. Species are
Stenotis Wollaston 1854 associated with dead wood of various types of trees, mostly
Eucoptus Wollaston 1873 conifers but also willows, aspens and poplars (Salicaceae). Rhyncolus
Borborhynchus Richard 1957 pallens Casey 1892 is associated with Lupinus arboreus Sims.
(Fabaceae) in California. See Casey (1892), Blatchley and Leng
37. Rhyncolini Gistel 1856 (1916) and Buchanan (1946) to separate the species. Some species
in Phloeophagus may be better placed as Rhyncolus (see note under
Rhyncolina Gistel 1856 Phloeophagus). The genus needs revision.
Rhyncholus Gistl 1834
Aphanommata Wollaston 1873, 1 sp., A. tenuis (Casey 1892), south- Eremotes Wollaston 1861
eastern United States west to Texas. Adults have been collected in Syntomocerus Wollaston 1865
tree hollow debris. Hyperemotes Voss 1934 (valid subgenus)
Rhamphocolus Casey 1892 Xylocomesus Thatcher 1940
 Family 131. Curculionidae · 761

Axenomimetes Voss 1955 (valid subgenus) 2(1). Tarsus with claws connate at base ....... Faustinus
— Tarsus with claws separate, not connate at base .
......................................................................... 3
Stenancylus Casey 1892, 2 spp., Florida. Stenancylus colomboi Casey
1892 is associated with Acrostichum (Pteridaceae) in Florida (Ander- 3(2). Antenna with funicle of 5 or 6 articles ................ 4
son 1993a). See Blatchley and Leng (1916) to separate the species. — Antenna with funicle of 7 articles ....................... 6
Liolepta Blatchley 1916
4(3). Body nearly glabrous or with sparse, narrow ap-
Rhinonus Kuschel 1959 pressed scales; elytra with stria 10 ended above
the hind coxa; frequently collected in wetlands
Tomolips Wollaston 1873, 1 sp., T. quercicola (Boheman 1845); .......................................................... Tyloderma
— Body with dense, often erect, scales; elytra with
generally distributed in the eastern United States. Larvae develop
stria 10 ended anterior to, or posterior to hind
in dead wood of various trees (Blatchley and Leng 1916). coxa; not in wetlands ....................................... 5
Wollastonia Horn 1873; not Heer 1852
Wollastoniella Cockerell 1906; not Reuter 1884 5(4). Pronotum with anterior portion with two subparallel
costae, markedly declivitous posteriorly (Fig. 60);
Parahornia Cockerell 1906
southwestern United States, in association with
ants ............................................... Liometophilus
Phloeophagina Voss 1955 — Pronotum with anterior portion simple, lacking cos-
tae, dorsal surface evenly rounded or on same
plane; southern Florida ................. Neoulosomus
Phloeophagus Schoenherr 1838, 5 spp., generally distributed in the
eastern United States north into Canada, west through the north, 6(3). Metasternum short, the distance between middle
then south into the western United States. Phloeophagus minor and hind coxae distinctly shorter than length of
Horn 1873 and P. californicus Van Dyke 1927 appear not to be antennal club; generally, the metepisternal su-
ture is not distinct and wings are absent or re-
congeneric with the remaining species and appear better placed as
duced; scutellum usually not visible, if visible,
Rhyncolus, which is where they are treated in the key. See Blatchley clothed with scales .......................................... 7
and Leng (1916) and Van Dyke (1927) to separate the species. — Metasternum long, the distance between middle
and hind coxae subequal to or distinctly longer
than length of antennal club; generally, the
X. Cryptorhynchinae Schoenherr 1825
metepisternal suture is distinct and wings are
present; scutellum usually visible ................. 15
by Robert S. Anderson
7(6). Abdomen with ventrite 1 about as long as or longer
than length of ventrites 2-5 combined, with large,
Members of this subfamily are easily recognized by the ventral
deep transverse depression or smaller lateral im-
channel on the sternum (Fig. 59), in which the rostrum lies in pressions near posterolateral margins ...............
repose, being extended beyond the prosternum onto the mesos- ........................................................... Eurhoptus
ternum or even the metasternum. Usually, the eyes are covered by — Abdomen with ventrite 1 shorter than length of
ventrites 2-5 combined, simple, lacking any de-
the anterolateral margins of the pronotum when the rostrum is
pressions .......................................................... 8
in respose, and the tibia possess a large, curved apical tooth. The
numbers of genera and species present in North America are very 8(7). Hind tibia abruptly expanded in basal one-third,
small compared to the taxonomic and structural diversity of about twice as wide as at apex ........... Canistes
— Hind tibia subequal in width throughout length or
cryptorhynchines in the Neotropical Region. A detailed study of
slightly expanded towards apex ..................... 9
the genera is much needed.
Cryptorhynchine larvae generally are borers in dead wood 9(8). Ventrite 2 about as long as ventrites 3-5 combined;
although some mine inside living plants and some species are pronotum markedly produced anteriorly over
head, head not visible in dorsal view; elytra with
found in seeds. Most genera are found in terrestrial habitats al-
apices produced, apex broadly truncate ..........
though species of the genus Tyloderma are associated with aquatic ........................................................... Lembodes
habitats. Many species are found in leaf litter and the odd south- — Ventrite 2 distinctly shorter than ventrites 3-5 com-
western species Liometophilus manni Fall 1912 is associated with bined; pronotum at most only slightly produced
anteriorly over head, head visible in dorsal view;
ants. Many species are flightless. No species are serious economic
elytra with apices not produced, apex evenly
pests although the mango weevil, Sternochetus mangiferae (Fabri- rounded .......................................................... 10
cius 1775), is frequently intercepted in quarantine at United States
border inspections. 10(9). Abdomen with suture separating ventrites 1 and 2
deep, nearly straight ...................................... 11
— Abdomen with suture separating ventrites 1 and 2
KEY TO THE NEARCTIC GENERA OF CRYPTORHYNCHINAE shallow, indistinct .......................................... 14

1. Tarsus with claws separate, with basal tooth ........ 11(10). Frons wider than apex of rostrum in dorsal view;
.......................................................... Phyrdenus ventrite 2 slightly longer than ventrites 3 and 4
— Tarsus with claws separate or connate basally, lack- combined ................................................. Calles
ing basal tooth ................................................. 2
762 · Family 131. Curculionidae

13(12). Elytra with stria 10 complete, extended to apex of

60 elytra; scutellum visible, clothed with scales;
59 pronotum in dorsal view widest at base ...........
......................................................... Pseudomus
— Elytra with stria 10 absent or indicated at most only
by a few small punctures in basal one-third of
length; scutellum not visible; pronotum in dorsal
view widest near middle ............. Gerstaeckeria

14(10). Abdomen with ventrite 2 obviously longer than

length of ventrites 3 and 4 combined, not shorter
than ventrite 5; frons wider than apex of rostrum
in dorsal view; sternal channel with posterior mar-
gin anterior to level of anterior margin of middle
coxae .......................................... Pseudoacalles
— Abdomen with ventrite 2 at most slightly longer than
length of ventrites 3 and 4 combined, shorter
than ventrite 5; frons as wide as or narrower than
apex of rostrum in dorsal view; sternal channel
with posterior margin at or posterior to level of
anterior margin of middle coxae ............ Acalles

15(6). Hind tibia lacking apical comb of stout setae;

pronotum markedly produced over head, head
not visible in dorsal view; tibia very short, less
than one-half as long as femur ..... Paracamptus
— Hind tibia with apical comb of stout setae (usually
arranged subparallel to long axis of tibia);
pronotum not produced over head, head visible
in dorsal view; tibia longer than one-half length
of femur .......................................................... 16

16(15). Middle coxae separated by about two-thirds width

of apex of rostrum .......................................... 17
— Middle coxae separated by at least width of apex
of rostrum ....................................................... 19

62 17(16). Abdomen with ventrite 2 longer than length of

ventrites 3 and 4 combined; hind tibia with apical
comb of stout setae longer than width of tibia at
apex; metepisternal suture not visible ..............
63 ......................................................... Maemactes
— Abdomen with ventrite 2 shorter than length of
61 ventrites 3 and 4 combined; hind tibia lacking
apical comb of stout setae; metepisternal suture
FIGURES 59.131-63.131. Cryptorhynchinae. 59. Cryptorhynchus lapathi visible ............................................................. 18
(Linnaeus), thoracic sterna, ventral view; 60. Liometophilus manni Fall,
18(17). Frons much wider than narrowest part of rostrum in
dorsal habitus; 61. Cryptorhynchus lapathi (Linnaeus), head, anterior
dorsal view; elytra with stria 10 complete; hind
view; 62. Cophes oblongus (LeConte); head, anterior view; 63. Zascelis tibia with apical comb composed of a complete
irrorata LeConte, middle tibia. apical row of setae and a confused row above
that; femur simple, not sulcate ventrally ..........
— Frons narrower than apex of rostrum in dorsal view;
............................................................. Euscepes
ventrite 2 subequal to or shorter than length of
— Frons about one-half as wide as narrowest part of
ventrites 3 and 4 combined ........................... 12
rostrum in dorsal view; elytra with stria 10 ended
anterior to hind coxa; hind tibia with short apical
12(11). Sternal channel with posterior margin not extended
comb running perpendicular to long axis and an-
posterior to level of hind margin of front coxae;
other short row at 45 degree angle to apical comb
mesosternum posterior to channel with median
and located above it; femur sulcate ventrally for
longitudinal carina; frons impressed between
reception of tibia ................ Apteromechus (part)
eyes; pronotum with deep, median longitudinal
sulcus ................................................ Peracalles
19(16). Body with suberect or erect, very fine, long, hair-
— Sternal channel with posterior margin extended pos-
like vestiture .................................................. 20
terior to level of hind margin of front coxae; me-
— Body with at most some scattered suberect or erect
sosternum posterior to channel simple, lacking
short, broad scales or scales arranged in tufts,
carina; frons mostly flat between eyes; pronotum
no hair-like vestiture evident ......................... 21
simple, lacking sulcus .................................... 13
20(19). Legs with tibiae with outer margin with large serra-
tions or denticles; antenna with funicle with ar-
 Family 131. Curculionidae · 763

ticle 2 long and slender, more or less twice as ously wider at base than pronotum at base ......
long as article 3 and about one and one-half times .................................................................. Sudus
longer than article 1; body length greater than — Abdomen with ventrite 2 not narrowed laterally, su-
4.0 mm; Arizona ............................ Cnemidoprion ture between ventrites 2 and 3 straight lateral
— Legs with tibiae with outer margin simple; antenna margin; leg with femur with ventral tooth; rostrum
with funicle with article 2 shorter and stouter, with antenna inserted posterior to midlength;
less than twice as long as article 3 and subequal elytra with humerus rounded, only slightly wider
to length of article 1; body length less than 4.0 at base than pronotum at base .... Pseudomopsis
mm; southern Florida ................... Eutinobothrus
28(22). Hind tibia with apical comb composed of a double
21(19). Leg with tibia with outer margin with acute serrate row of setae (may be irregular); elytra either with
or dentate carina, especially in apical one-third intervals 3 and 5 prominent or with no intervals
(Fig. 63) ................................................. Zascelis prominent and no erect setae or nodules present
— Leg with tibia with evenly rounded outer margin, ................................................................ Cophes
lacking carina ................................................. 22 — Hind tibia with apical comb composed of a single
row of setae; elytra either with erect short stout
22(21). Mandible prominent when closed, with obvious setae, nodules, or with sutural interval prominent
acute tooth on inner margin, angle between basal immediately behind scutellum ....................... 29
and apical cusps acute (Fig. 61) ..................... 23
— Mandible not prominent when closed, lacking tooth 29(28). Elytra with large, polished, prominent nodules; an-
on inner margin or if tooth present, angle between tenna with club lacking distinct sutures; pronotum
basal and apical cusps obtuse (Fig. 62) ......... 28 with dense, round overlapping scales and elytra
with small, narrow scales .................... Hohonus
23(22). Frons not or only slightly narrower than narrowest — Elytra with short, stout setae or with sutural interval
part of rostrum in dorsal view (Fig. 61) ........... 24 prominent immediately behind scutellum; an-
— Frons about one-half width of narrowest part of ros- tenna with club with distinct sutures; pronotum
trum in dorsal view ......................................... 26 and elytra with similar scales ......................... 30

24(23). Pronotum distinctly produced over head, head not 30(29). Elytra laterally at base emarginate to receive promi-
visible in dorsal view; eyes almost completely nent hind angle of pronotum; elytra with sutural
covered by postocular lobes when rostrum is in interval prominent immediately behind scutellum,
repose; eyes widely separated by slightly more stria 10 indicated behind level of hind coxa as a
than width of rostrum at base; rostrum more or fine line; pronotum with postocular lobes promi-
less straight ........................................... Troezon nent and rounded; front femur with ventral mar-
— Pronotum at most slightly produced over head, head gin with small tooth; front tarsus of male elongate
partially visible in dorsal view; eyes only partially and with long, fine setae .................... Episcirrus
covered by postocular lobes when rostrum is in — Elytra laterally at base simple, not modified to re-
repose; eyes moderately separated by slightly ceive hind angle of pronotum; elytra with sutural
less than width of rostrum at base; rostrum more interval flat immediately behind scutellum, stria
or less evenly curved .................................... 25 10 not extended behind level of hind coxa;
pronotum with postocular lobes slightly devel-
25(24). Elytra with alternate intervals 3, 5 and 7 variously oped; front femur simple, lacking tooth; front tar-
elevated, sharply carinate or lower and rounded; sus of male normal .............................. Rhynchus
hind femur with ventral margin with single acute
tooth (species with alternate elytral intervals cari-
CLASSIFICATION OF THE NEARCTIC CRYPTORHYNCHINAE
nate) or with two teeth that are broadly joined at
their bases (species with alternate intervals con-
vex) ........................................................ Eubulus 38. Cryptorhynchini Schoenherr 1825
— Elytra with intervals of more or less equal height,
although alternate intervals 3, 5 and 7 may pos-
Cryptorhynchina Schoenherr 1825
sess patches of suberect or erect broad scales
giving an elevated appearance; hind femur with
ventral margin simple or with one or two separate Apteromechus Faust 1896, 4 spp., generally distributed in the east-
acute teeth .............................. Cryptorhynchus ern United States and Canada west to Texas and Arizona. Adults
come to lights and are associated with various trees such as beech,
26(23). Elytra with stria 10 complete, indicated beyond level
of hind coxa; hind tibia with apical comb arranged oak and red bay; larvae have been found mining a dead sassafras
in two discrete parts ......... Apteromechus (part) limb (Kissinger 1964). See Whitehead (1979) to separate the spe-
— Elytra with stria 10 not extended beyond level of cies.
hind coxa; hind tibia with apical comb continu-
Acarlosia Hustache 1940
ous, not separated into two parts .................. 27

27(26). Abdomen with ventrite 2 markedly narrowed later- Cnemidoprion Marshall 1933, 1 sp., C. oblongus (Horn 1895), Ari-
ally, suture between ventrites 2 and 3 markedly zona. Adults come to lights and have been collected on low
angulate posteriorly at lateral margin for distance
roadside vegetation. A Brazilian species was reared from twigs of
about one-half width of ventrite 3; leg with femur
simple; rostrum with antenna inserted anterior to Cienfugosia (Malvaceae) (Anderson 1998).
midlength; elytra with prominent humerus, obvi-
764 · Family 131. Curculionidae

Cryptorhynchus Illiger 1807, 4 spp., generally distributed in the pest of mango but to date has not become established in North
eastern United States north into Canada, west across Canada and America.]
the northern United States to Oregon, Washington and British
Columbia. Adults of C. fuscatus LeConte 1876, C. helvus LeConte Sudus Kissinger 1964, 1 sp., S. floridanus Kissinger 1964, south-
1878 and C. minutissimus LeConte 1876 come to lights and are eastern United States west to Texas.
associated with various species of dead or dying trees.
Cryptorhynchus lapathi (Linnaeus 1758) is associated with living Troezon Champion 1906, 1 sp., T. lutosus (LeConte 1884), Florida
poplar and willow (Salicaceae). See Blatchley and Leng (1916) to and Louisiana. This species is associated with Dahlbergia
separate the species. The genus in North America likely is com- ecastophyllum (L.) Benth. (Fabaceae); larvae feed in the disc-like
posite and needs redefinition. fruits (Blatchley and Leng 1916; Anderson 1993a).
Arachnipes Villa and Villa 1833
Cryptorrhynchus Bedel 1884 Tyloderma Say 1831, 30 spp., generally distributed in the United
Cryptorrhynchus Champion 1906 States (most species in the southeast) and southern Canada. Spe-
Cryptorhynchidius Pierce 1919 cies are associated with various plants, many in wetlands, in the
Atrichopsis Voss 1954 (valid subgenus) families Onagraceae, Polygonaceae, Urticaceae, Haloragaceae, Ro-
Cryptorrhynchobius Voss 1965 saceae, Melostomataceae and Saururaceae (Wibmer 1981). Adults
come to lights. See Wibmer (1981) to separate the species.
Eubulus Kirsch 1870, 3 spp., generally distributed in the eastern Analcis Say 1831; not Wagler 1830
United States into southern Canada, west in the south to Texas. Analcis Schoenherr 1833; not Wagler 1830; not Say 1831
Arizona and California. Adults of E. bisignatus (Say 1831) and E.
parochus (Herbst 1797) have been associated with dead limbs of Zascelis LeConte 1876, 1 sp., Z. irrorata LeConte 1876, southwest-
chestnut, beech, oak and birch (Blatchley and Leng 1916); larvae ern United States. Adults come to lights. A second unnamed
likely mine dead branches of various trees. A single adult of E. (and likely adventive) species is present in southern Florida (Ander-
obliquus (Say 1831) has been associated with Myrica cerifera L. son 1993a).
(Myricaceae) (Anderson 1993a). See Blatchley and Leng (1916) to
separate the species. Tylodina Lacordaire 1866
Eubulosoma Voss 1954 (valid subgenus)
Acalles Schoenherr 1825, 12 spp., generally distributed in the east-
Eutinobothrus Faust 1896, 1 sp., E. pilosellus (Boheman 1844), ern United States into southern Canada, west to Texas, Arizona
southern Florida. This species is associated with Ipomoea and New Mexico in the south. Species are associated with various
(Convolvulaceae) (Anderson 1993a). dead branches or palm fronds on the ground, on dead vines and
Gasterocercodes Pierce 1915 other hanging dead vegetation, and generally in leaf litter (Kissinger
1964; Anderson 1993a). Adults were also found sweeping
Liometophilus Fall 1912, 1 sp., L. manni Fall 1912, Arizona, New Borrichia (Asteraceae), Sesuvium (Aizoaceae) and Salicornia and Suaeda
Mexico and Texas. This species is associated with nests of the ant (Chenopodiaceae) on beaches in southern Florida at night (Ander-
Liometopum apiculatum Mayr (Formicidae). This is perhaps the son 1993a). See Blatchley and Leng (1916) to separate some of
oddest-looking weevil in North America. the species. The genus needs revision and redefinition. Anderson
(1993a) lists six undescribed species as present in southern Florida.
Maemactes Schoenherr 1837, 1 sp., M. cribratus (LeConte 1876), Ulosomus Schoenherr 1825
Texas and Kansas. Microdalotes Gistel 1856
Baropsis LeConte 1876 Trachodius Weise 1891
Baridopsis Rye 1878 Milichacalles Voss 1960 (valid subgenus)
Trichacalles Voss 1960 (valid subgenus)
Neoulosomus O’Brien and Wibmer 1982, 1 sp., N. laticaudis (Suffrian
1872), southern Florida. Adults have been collected on various Calles Kissinger 1964, 1 sp., C. cladotrichis (Pierce 1912), Arizona,
dead limbs and vines (Anderson 1993a). New Mexico and Texas. Adults were reared from roots of
Ulosomus Schoenherr 1826; not Schoenherr 1825 Tidestromia lanuginosa (Nutt.) Standl. (Amaranthaceae) (Pierce
1912). A second undescribed species has been collected in south-
Phyrdenus LeConte 1876, 2 spp., P. divergens (Germar 1824) gener- ern Florida on Salicornia and Suaeda (Chenopodiaceae) at night in
ally distributed in the eastern United States, and P. muriceus (Germar coastal areas (Anderson 1993a) and another is known from Ari-
1824), Florida and Arizona. These species are associated with zona.
Solanum (Solanaceae) (Blatchley and Leng 1916; O’Brien 1961).
Canistes Casey 1892, 1 sp., C. schusteri Casey 1892, eastern United
[Sternochetus Pierce 1917, 1 sp., S. mangiferae (Fabricius 1775), in- States west to Texas. Adults have been collected in leaf litter.
tercepted in quarantine; Florida and California. This species is a
 Family 131. Curculionidae · 765

Eurhoptus LeConte 1876, 2 spp., E. pyriformis LeConte 1876 and Pseudomus Schoenherr 1837, 2 spp., P. sedentarius (Say 1831), Florida
E. sordidus (LeConte 1876), generally distributed in the eastern and P. truncatus LeConte 1876, Georgia and South Carolina. See
and southcentral United States west to Texas and Oklahoma. Blatchley and Leng (1916) to separate the species.
Adults have been commonly collected in leaf litter. At least three
undescribed species are known from Texas; the genus needs revi- 39. Gasterocercini Zherichin 1991
sion.
Eurrhoptus Rye 1878 Cophes Champion 1905, 5 spp., generally distributed in eastern
United States and southern Canada west into Texas. Adults of
Euscepes Schoenherr 1844, 1 sp., E. porcellus Boheman 1844, south- most species are associated with dead wood and come to lights.
ern Florida. Adults have been collected on Ipomoea Cophes texanus Sleeper 1955 has been reared from dead Baccharis
(Convolvulaceae) (Anderson 1993a); larvae are likely in the roots neglecta Britt. (Asteraceae). The genus needs revision and redefini-
or stems. tion. See Sleeper (1955a) and Blatchley and Leng (1916; as
Hyperomorpha Blackburn 1885 Cryptorhynchus) to separate some of the species.
Batatarhynchus Hustache 1933 Coelosternus Schoenherr 1835; not Sahlberg 1823
Sternocoelus Kuschel 1955
Faustinus Berg 1898, 1 sp., F. cubae (Boheman 1844), southern
Florida. Adults and larvae are associated with various Solanaceae Episcirrus Kuschel 1958, 1 sp., E. brachialis (LeConte 1884), Texas
(Anderson 1993a). and Arizona. Adults are associated with Bumelia lanuginosa (Michx.)
Euxenus Faust 1896; not Gistel 1856; not LeConte 1876 Pers. (Sapotaceae); larvae appear to mine dead branches.

Gerstaeckeria Champion 1905, 18 spp., southeastern and western Hohonus Kissinger 1964, 1 sp., H. lacteicollis (Champion 1906),
United States, north into western Canada. Adults and larvae are Texas and Arizona. Adults are associated with Phoradendron
associated with various Cactaceae. Adults are flightless and (mistletoe; Viscaceae); larvae mine the stems (Anderson 1994).
noctural. Larvae mine the pads of Opuntia and hollow out smaller
pincushion cacti such as Mamillaria and Coryphanta. See O’Brien Rhynchus Kissinger 1964, 1 sp., R. apiculatus (Gyllenhal 1837),
(1970b) to separate the species. southeastern United States. Adults are associated with Myrica
Opuntiaphila Pierce 1912 cerifera L. (Myricaceae); larvae mine in dead trunks and larger
Philopuntia Pierce 1912 branches (Ford 1985).

Lembodes Schoenherr 1844, 1 sp., L. solitarius Boheman 1844, south- XI. Cyclominae Schoenherr 1826
ern Florida. Adults are collected on various types of dead vegeta-
tion (Anderson 1993a). by Robert S. Anderson

Paracamptus Casey 1895, 2 spp., P. floridanus Sleeper 1954 and P. This is a small group of three genera of weevils, one of which
subtropicus Casey 1895, southern Florida. Adults of P. subtropicus appears not closely related to the other two. All members have a
have been collected commonly on dead Rhizophora mangle L. (red relatively short snout (but lack any deciduous processes and associ-
mangrove; Rhizophoraceae) branches (Anderson 1993a). See ated scars) and have well-developed postocular lobes (Fig. 66). The
Sleeper (1954b) to separate the species. genus Listroderes, represented by only 3 species, is introduced from
South America, a region of much greater cyclomine diversity.
Peracalles Kissinger 1964, 2 spp., P. pectoralis (Leconte 1876), Illi- Listronotus is a large genus of over 80 species, most of which are
nois, Indiana, Ohio, Kentucky and Missouri, and P. ventrosus associated with semi-aquatic and aquatic habitats. Emphyastes is an
(LeConte 1878), Florida. Adults occur in leaf litter and P. ventrosus odd, unrelated genus found associated with seaweed along Pacific
has been collected from emergent aquatic vegetation at night (C. coastal beaches.
W. O’Brien, pers. comm.). See Blatchley and Leng (1916; as Acalles)
to separate the species. KEY TO THE NEARCTIC GENERA OF CYCLOMINAE

Pseudoacalles Blatchley 1916, 1 sp., P. nuchalis (LeConte 1876), 1. Front tibia prolonged beyond articulation of tarsus
into long, flattened paddle; hind tibia markedly
Florida and South Carolina. Adults are found in leaf litter (Ander-
expanded at apex, wider than maximum width of
son 1993a). hind femur (Fig. 67) ........................ Emphyastes
— Front tibia not prolonged beyond articulation of tar-
Pseudomopsis Champion 1905, 1 sp., P. inflata (LeConte 1876), sus; hind tibia not expanded at apex, not as wide
as maximum width of hind femur ...................... 2
southern Florida. Adults are commonly found on Coccoloba uvifera
L. and C. diversifolia Jacq. (Polygonaceae); larvae feed in fruits 2(1). Pronotum widest subapically, lateral margins straight
(Anderson 1993a). and divergent from base to widest point, trans-
versely quadrate in form (Fig. 66); pronotal disk
766 · Family 131. Curculionidae

Listroderina LeConte 1876

Listroderes Schoenherr 1826, 3 spp., L. costirostris Schoenherr 1826,

L. difficilis Germain 1895, and L. apicalis Waterhouse 1841, south-
eastern United States, Texas, Arizona and California; adventive.
See Morrone (1993) to separate the species.

Listronotus Jekel 1865, 81 spp., generally distributed in the United

States and Canada. Adults are found in wetlands and appear to
66 be associated with a variety of plants (O’Brien 1981, Anderson
65
1993a). Adults of a few species in coastal Florida and Texas have
been collected on Borrichia frutescens (L.) DC. (Asteraceae) and
Salicornia (Chenopodiaceae) (Anderson 1993a). The genus, espe-
cially the smaller species originally placed in Hyperodes, needs revi-
sion. See O’Brien (1981) and Stockton (1963) to separate most of
the species.
Macrops Kirby 1837; not Wagler 1830; not Burmeister 1835
Hyperodes Jekel 1865
Anchodemus LeConte 1876
Lixellus LeConte 1876
Mascarauxia Desbrochers 1898
67 Relistrodes Brèthes 1910
64 Aulametopiellus Brèthes 1926
FIGURES 64.131-67.131. Cyclominae. 64. Listronotus caudatus (Say), Pseudohyperodes Hustache 1939
head, lateral view; 65. Listronotus oregonensis (LeConte), pronotum,
dorsal view; 66. Listroderes costirostris Schoenherr, pronotum, dorsal XII. Entiminae Schoenherr 1823
view; 67. Emphyastes fucicola Mannerheim, hind tibia.
by Robert S. Anderson and Anne T. Howden
broad and flat, lateral profile with anterior margin
at middle elevated above level of pronotal disk;
body size greater than 5 mm; elytra with erect, Entiminae are generally called the ‘broad-nosed’ weevils because this
stout seta-like scales at least 2 to 3 times longer is the group of curculionids in which the snouts are the least devel-
than diameter of adjacent rounded, flat scales . oped. Aside from the possession of a shorter, broader snout, the
......................................................... Listroderes
— Pronotum widest near base or at or before midlength,
best way to recognize them is that nearly all Entiminae have a man-
lateral margins rounded from base to widest point dible that bears a deciduous process that breaks off soon after emer-
(Fig. 65); subcylindrical in form; pronotal disk more gence of the adult leaving a definite scar at the point of attachment
or less evenly convex, lateral profile evenly on the outer face of the mandible. However, not all Entiminae
rounded from basal to anterior margin; body size
various; elytra with or without erect, stout seta-
possess this feature (Thecesternus, Sitona, and members of the tribe
like scales, but if seta-like scales as long as 2 to 3 Alophini) and one must rely on other features in recognizing their
times diameter of adjacent, rounded scales, body inclusion in the subfamily. Entimines also possess only a short
size is less than 5 mm ..................... Listronotus tooth or spine on the inner angle at the apex of the hind tibia and
sexual dimorphism in the form of the rostrum is generally not as
CLASSIFICATION OF THE NEARCTIC CYCLOMINAE evident as in other weevils. The antennal scape of some species also
extends to or beyond the anterior margin of the eye, a feature other-
40. Rhythirrinini Lacordaire 1863 wise only found in Dryophthorinae.
Most entimines have larvae that feed externally in the soil on
Emphyastina Lacordaire 1863 roots whereas the adults tend to feed on fresh foliage or repro-
ductive structures such as flowers or buds. Many species are gen-
Emphyastes Mannerheim 1852, 1 sp., E. fucicola Mannerheim 1852, eralists and feed on a very broad range of plant taxa both as
western coastal United States and Canada north into southern adults and larvae (e.g., Otiorhynchus ovatus (Linnaeus 1758)) whereas
Alaska. Adults and larvae are associated with decaying seaweed others can be very host specific, feeding on a few closely related
washed up and buried on sandy beaches (Anderson 1988b). species or genera. Oviposition usually takes place in the soil or
Korotyaev and Egorov (1975) have suggested that this genus is rarely on the foliage of the host plant, larvae then dropping to
related to Thalasselephus (Molytinae). the ground to feed in the soil. Adults of many species of
Entiminae are flightless and some are parthenogenetic.
 Family 131. Curculionidae · 767

68 70
72 73
69
71

76
74
75
77

FIGURES 68.131-77.131. Entiminae. 68. Mandibles of Entiminae, schematic (after Kissinger 1964). 69. Thecesternus sp., head and thorax,
ventral view (after Kissinger 1964). 70-71. Brachystylus sayi (Alonso-Zarazaga), head, 70. Lateral view; 71. Dorsal view. 72-73. Sciopithes obscurus
Horn, head 72. Lateral view; 73. Dorsal view. 74-75. Dyslobus lecontei Casey, head, 74. Lateral view; 75. Dorsal view. 76-77. Ericydeus lautus
(LeConte), head, 76. Lateral view; 77. Dorsal view.

Entiminae appear to be very adundant and diverse in arid habi- — Rostrum and sides of head not recessed,
tats, particularly in the deserts of the southwestern United States. prosternum not forming such a cavity; mandible
with or without deciduous process; tarsi with
Some have developed adaptations for sand dwelling that include or without pads on ventral surface, tarsal claws
dense long hairs over the body and fossorial legs. Entimines are also free or connate; body length various .......... 2
the weevils most often found at higher elevations.
A number of species are pests of ornamental plants and of 2(1). Mandible large, hemispherical externally, inner
surface slightly cupped; surface of mandible
agricultural produce including citrus and other fruits. These in- densely squamate except narrow median edge
clude Otiorhynchus ovatus (strawberry root weevil), O. sulcatus (Fab- glabrous; mandible without deciduous pro-
ricius 1775) (black vine weevil), Cyrtepistomus castaneus (Roelofs cess; bucal cavity large, maxillary palpus fully
1873) (Asiatic oak weevil), Artipus floridanus Horn 1876, and spe- or mostly exposed; scrobes lateral; rostrum
similar to head in length and width; anterior
cies of Naupactus and Sitona. edge of prothorax straight laterally, not lobed
Entiminae are the most diverse subfamily in North America beneath eye ........................................ Sitona
with 124 genera in 23 tribes recognized. In some tribes such as — Mandible various; if large and densely squamate,
Peritelini, identification of the genera is very difficult and generic then with postocular lobe and/or with scrobe
dorsal and/or with deciduous process; other
definitions need to be reassessed. characters various ........................................ 3
The portions of the key from couplet 68 to 84 and 105 to
122 are slightly modified from Kissinger (1964). The portion of 3(2). Mandible prognathous, like a thin roof extending
the key from couplet 85 to 104 was adapted from Sleeper (1955b) over mouth parts; mandible without a decidu-
ous process, or process very small and incon-
and O’Brien (1984). An application is before the International spicuous; all or most of mouthparts exposed in
Commission on Zoological Nomenclature to maintain ventral view; elytra with humeri rounded; tar-
Trachyphloeini (Alonso-Zarazaga and Lyal 1999, p. 8). sal claws free ................................................ 4
— Mandible not prognathous; mandible with decidu-
ous process or its scar (Fig. 68); elytra with
KEY TO THE NEARCTIC GENERA OF ENTIMINAE humeri various; tarsal claws free or not ....... 6

1. Rostrum and sides of head recessed in deep cav- 4(3). Mentum as wide as bucal cavity, maxillae only
ity formed by the sides of the prothorax and by briefly exposed basally; mandible with decidu-
prosternum in the form of a small triangular plate ous process or its scar present on extreme
originating between fore coxae (Fig. 69); when outer angle; elytra flat basally in vicinity of
positioned within the cavity, only dorsal sur- scutellum; pronotum with glabrous median ca-
faces of rostrum and frons visible; mandible rina; body length up to 9.0 mm .... Byrsopages
without deciduous process; tarsi without pads — Mentum as wide as one-third width of bucal cav-
on ventral surface; tarsal claws free; body ity; mouthparts completely exposed in ventral
length 6.5-9.0 mm ..................... Thecesternus view; deciduous process of mandible lacking
768 · Family 131. Curculionidae

or very small; elytra not flat basally; body length — Head without a sulcus across ventral surface of
up to 6.0 mm ................................................. 5 head-rostrum junction; humeri rounded ..... 13

5(4). Dorsal surface clothed with appressed oval or fan- 13(12). Base of elytra carinate; rostrum dorsally with fine,
shaped scales with fine ribs radiating from point deep median sulcus and shorter lateral sulcus
of attachment; body surface not encrusted with at base that is abruptly turned laterally toward
dirt; scrobe evanescent dorsally, not reaching antennal scrobe; body length 4.3-5.6 mm .....
anterior edge of eye; scape reaching posterior ........................................................... Sapotes
edge of eye; body length 3.5-5.6 mm ............... — Base of elytra not carinate, sloping to mesotho-
...................... Dirotognathus and Lepidophorus rax; rostrum with or without sulci, not with right-
— Dorsal surface dirt-encrusted, with papillae; angled dorsolateral sulcus; postocular lobe
scales not readily visible; scrobe well-defined with row of setae of graduated lengths; body
dorsally, continuing above dorsal edge of eye; length 5.0-25.0 mm ..................................... 14
scape abruptly thickened distally, reaching
anterior edge of eye; body length 3.0-3.3 mm 14(13). Rostrum with dorsal margin of scrobe well-de-
.......................................................... Vitavitus fined, with a sharp upper angle; antenna with
scape and funicle, and dorsal surfaces of tar-
6(3). Dorsal surface dirt-encrusted, with papillae, with- sal articles all with round overlapping scales;
out scales; mentum almost completely cover- body lacking long, fine, erect hairs ...............
ing bucal cavity; scrobe ending well before ..................................................... Ophryastes
eye; tarsal claws free; body length 3.5-6.3 mm — Rostrum with dorsal margin of scrobe less well-
..................................................... Leptopinara defined, with a rounded upper angle; with one
— Dorsal surface squamate, without papillae, with or more of antennal scape and funicle, and dor-
or without crust of dirt; other characters vari- sal surfaces of tarsal articles, lacking round
ous ................................................................ 7 overlapping scales; body with or without long,
fine, erect hairs .......................................... 15
7(6). Bucal cavity deeply recessed, occupying approxi-
mately one-third width of rostrum, lateral walls of 15(14). Mandible with process or its scar linear, very small,
cavity perpendicular, posterior wall formed by rarely observed; sides of mandible large, flat-
distal edge of gula, anterior wall formed by ven- tened or slightly concave, irregularly sculp-
tral surface of mandibles when closed; mentum tured; scape and funicle without broad scales;
distally occupying full width of bucal cavity; max- rostrum longer than head; eye teardrop shape
illae slightly exposed proximally; densely cov- or oval; size large, length 10.0-17.0 mm .... 16
ered with shiny, appressed, whitish, slightly opal- — Mandibular process or its scar usually much larger
escent scales; tarsal claws connate; body length and distinct (Fig. 68); sides of mandible not as
2.6-3.8 mm .................................. Connatichela above; antennal scape and funicle various; size
— Bucal cavity not so deeply recessed, if at all; various ........................................................ 19
mentum of various widths; vestiture dense or
not; tarsal claws connate or not; other charac- 16(15). Scutellum triangular, not conspicuous in dorsal
ters various ................................................... 8 view; integument shiny through sparse, elon-
gate scales and stiff dark setae . Acmaegenius
8(7). Side of prothorax with anterior margin produced — Scutellum rectangular, readily visible in dorsal
into slight to very large rounded postocular view, conspicuously densely clothed with ap-
lobe (Fig. 74); eye flat, tear-drop shaped (Fig. pressed setae/scales ................................. 17
74) ................................................................. 9
— Side of prothorax with anterior margin straight, 17(16). Rostrum with three very deep sulci; one median
not produced (Figs. 70, 72, 76); eye various . dorsal sulcus from interantennal line to frons;
.................................................................... 44 one lateral sulcus on each side from near an-
tennal insertion to upper edge of eye, thence
9(8). Mandible with four or more large setae; femur vari- following anterior edge of eye to scrobe .....
ous .............................................................. 10 ..................................................... Triglyphulus
— Mandible with three large setae; femur with a — Rostrum with sulci, if present, not present along
tooth on inner edge distally ....................... 39 edge of eye ................................................ 18

10(9). Ventral edge of postocular lobe very abrupt, 18(17). Elytra with slight humeral angle; striae with deep
angulate ...................................................... 11 elongate punctures ........................ Plinthodes
— Ventral edge of postocular lobe gradual, more — Elytra with humeral angle rounded; striae with shal-
rounded ...................................................... 12 low punctures ............................ Trichalophus

11(10). Elytra oval, scutellum seldom visible dorsally; 19(15). All surfaces densely squamate with smooth, shiny
body length 2.7-3.5 mm ................ Aracanthus scales; with long fine erect hairs as much as
— Elytra with humeral angle slanted from stria 7 out- 1.0 mm long on dorsal surface including legs
wards; scutellum very wide; prothorax and and antennal funicle (Fig. 79); front tibia with
elytra patterned with lines and other markings; outer angle expanded, middle tibia less so, hind
body length 4.1-8.0 mm ............... Eudiagogus tibia with outer and sometimes inner angle
greatly expanded; tibiae edged distally with
12(10). Head with a deep sulcus across ventral surface row of stout spines; many species with tarsal
of head-rostrum junction, sulcus continuous articles spinose ventrally, particularly article
with scrobe; humeri quadrate ......... Colecerus
 Family 131. Curculionidae · 769

25(24). Dorsum very irregularly, coarsely sculptured; ros-

trum with three broad sulci; head with swelling
above eye; elytra with costae on intervals 3, 5,
79 7; with large knob on apical umbone and large
78
knob on declivity ............................ Rhigopsis
— Surface not irregularly sculptured as above; other
characters various ...................................... 26

26(25). Metepisternal suture present, well defined at

least in basal half, usually complete, rarely oblit-
erated; if suture is obliterated, then
metepisternum with finer, smaller, and sparser
scales than metasternum ............................ 27
— Metepisternal suture obliterated entirely or in
large part at least in basal half, metepisternum
clothed with scales similar in coarseness, size,
and density ................................................. 28

27(26). Dorsal margin of scrobe indistinct posteriorly;

scape rests on eye when retracted next to
head; rostrum separated from frons by distinct
80c transverse impression; abdominal ventrite 2
80a 80b
longer than 3 and 4 united, suture between
ventrites l and 2 deep, straight; article 1 of front
tarsus dorsally clothed with sparse round
scales .............................................. Adaleres
— Abdominal ventrite 2 not longer than 3 and 4
united, suture between ventrites 1 and 2 deep,
straight; article 1 of front tarsus lacking round
scales on dorsal surface ................. Dyslobus

28(26). Each puncture of elytral stria covered by round

FIGURES 78.131-80.131. Entiminae. 78-79. Dorsal habitus, 78.
scale; elytra usually with sparse, coarse, erect
Pandeleteius rotundicollis (Fall); 79. Miloderes nelsoni Kissinger. 80. Hind setosity ......................................... Panscopus
tibiae of various Entiminae, schematic. a) open corbel, b) semi-closed — Each puncture of elytral stria either with a fine
corbel, c) closed corbel. seta, apparently glabrous, or puncture not dis-
tinct; elytral vestiture various .................... 29
1; lacking complete pads on all tarsal articles;
associated with sandy environments ........ 20
29(28). Scape of antenna clothed with round, flat scales
— Vestiture dense or not; without long fine erect
.................................................................... 30
hairs and expanded tibial apices (if long erect
— Scape of antenna not clothed with round scales,
hairs and distinct postocular lobes present, see
only suberect, fine setae ........................... 35
Paracimbocera); other characters various . 24
30(29). Tibia with corbel open (Fig. 80a); body usually
20(19). Epistoma abruptly perpendicular, posterior mar-
dirt-encrusted; with long, erect, sparse, spatu-
gin carinate .......................... Trigonoscutoides
late setae; dorsal margin of scrobe poorly de-
— Epistoma on same plane as remainder of rostrum
fined posteriorly; scape extended over middle
.................................................................... 21
of eye ................................................ Phyxelis
— Tibia with corbel closed (Fig. 80c); other charac-
21(20). Eye convex ................................... Trigonoscuta
ters various ................................................. 31
— Eye flat ........................................................... 22
31(30). Elytra clothed with long, fine, erect setae, each
22(21). Tarsal claws connate basally or with a single claw
seta about five times as long as diameter of an
................................................. Eucilinus (part)
adjacent scale; hind tibia with corbel closed,
— Tarsal claws free ............................................ 23
corbel plate clothed with flat, round scales ..
........................................................ Diamimus
23(22). Antennal scrobe shallow and greatly widened
— Elytra either lacking erect setosity (Orimodema)
posteriorly .................................... Cimbocera
or with stout bristles not more than three times
— Antennal scrobe deep and only slightly widened
as long as diameter of scales (Dichoxenus); hind
posteriorly ...................................... Miloderes
tibia with corbel closed, corbel plate lacking
scales ......................................................... 32
24(19). Body with long fine erect hairs and distinct pos-
tocular lobes (as in Cimbocera, but tibial api-
32(31). Body elongate in form; elytra lacking erect
ces not expanded); tarsal article 3 with apical
setosity, scales of elytra not at all overlapping;
tufts (except females of P. robusta) ...............
dorsal margin of scrobe poorly defined poste-
................................................ Paracimbocera
riorly; in dorsal view lateral margins of rostrum
— Body without long fine erect hairs; tarsal article 3
distinctly converging from base at anterior
various ........................................................ 25
margin of eye to about the middle, thence
770 · Family 131. Curculionidae

nearly straight to point of insertion of antenna 40(39). Femora each with very large tooth bearing one or
..................................................... Orimodema two smaller teeth on distal edge; scape reach-
— Body shorter and stouter in form; elytra with ing anterior third of prothorax; base of elytron
sparse, erect bristles; in dorsal view lateral forming a large lobe between scutellum and
margins of rostrum slightly, evenly convergent interval 5; dorsal elytral setae minute; color dark
from base at anterior margin of eye to point of brown-black with whitish scales, with irregular
insertion of antenna ................................... 33 pattern created by areas of very small scales
allowing integument to show through; body
33(32). Sternite 8 of female compressed distally, the verti- length 6.0 -7.0 mm ........................ Myllocerus
cal dimension much greater than horizontal di- — Femora each with single small tooth; dorsal
mension; elytra with scales not overlapping, on elytral setae long, slender, arcuate; base of
dorsal surface with short, erect, clavate setae; elytra straight; color and body length various
scape densely squamate ............. Paranametis .................................................................... 41
— Sternite 8 of female shaped like a horizontal
shovel, with broad, laterally expanded process 41(40). Eye small, separated from anterior margin of pro-
distally; other characters various .............. 34 thorax by three or more scales; elytral inter-
vals flat; pronotum and elytra evenly, densely
34(33). Scape densely scaled; elytral setae blunt squamose; body length 3.3-4.0 mm ................
apically; ventral margin of scrobe clearly de- ...................................................... Oedophrys
fined .............................................. Dichoxenus — Eye large, separated from anterior margin of pro-
— Scape mostly setose with a few broad scales; thorax by single row of scales; elytral intervals
elytral setae fine, acute; ventral margin of convex; pronotum and disc of elytra with
scrobe poorly defined ..................... Anametis scales very sparse or absent, replaced with
minute setae; scales becoming more numer-
35(29). Basal margin of elytra produced strongly and ous laterally; body length 4.5-5.8 mm ...........
abruptly perpendicularly before merging with ................................................. Cyrtepistomus
sclerites covered by prothorax ................. 36
— Elytra lacking distinct basal margin, evenly 42(39). Pterygium open apically (Fig. 73); eye with ap-
rounded to sclerites covered by prothorax (or proximately 14 facets along longest axis; pro-
elytra lacking erect setosity, Melanolemma); thorax with distinct lobe on anterior edge be-
dorsal margin of scrobe poorly defined poste- low eye, prothorax here 1.2x longer than
riorly ........................................................... 37 length of pronotum; body length 3.5-3.8 mm
.................................................. Calomycterus
36(35). Dorsal margin of scrobe not defined posteriorly, — Pterygium various; prothorax straight on anterior
scape passing over middle of eye; corbel of edge, longer dorsally than ventrally; other char-
hind tibia open; body lacking erect setae or acters various ............................................. 43
scales; tarsal claws connate ....... Tropiphorus
— Dorsal margin of scrobe well defined posteriorly, 43(42). Pterygium closed apically (Fig. 71); eye small, flat-
scape passing over bottom of eye; corbel of tened, with approximately 20 facets along long-
hind tibia narrowly closed; body with erect fine est axis; prothorax 1.4x longer dorsally than
setae usually numerous; tarsal claws free ..... ventrally; elytral scales not sculptured; body
.......................................................... Peritaxia length 3.0-4.0 mm ........................ Neoptochus
— Pterygium open apically; eye with approximately
37(35). Elytra clothed with long, fine, erect setae; elytral 10 facets along longest axis; elytral scales
stria 10 extending posteriorly to above margin sculptured in a fan shape; body length 2.4-4.0
of hind coxa, there joining stria 9 .... Crocidema mm .................................................... Myosides
— Elytra lacking erect setae; elytral stria 10 obscure,
not joining stria 9 ........................................ 38 44(8). Anterior edge of prothorax laterally with postocu-
lar vibrissae in a cluster or tuft, or lacking; with-
38(37). Rostrum flat on dorsal surface, there with fine, out lobe (except Pachnaeus); with or without
longitudinal, glabrous median line; frons two- tooth or knob .............................................. 45
fifths wider than distance between lateral mar- — Anterior edge of prothorax without postocular
gins of rostrum at point of insertion of antenna vibrissae ..................................................... 55
in dorsal view; rostrum lacking deep, fine sul-
cus distad of eye ........................ Pseudorimus 45(44). Elytra with humeral angle well-developed, distinct
— Rostrum on dorsal surface convex, there with (Fig. 78; less so in Pandeleteius simplarius) 46
very vague, low, median longitudinal carina; — Elytra with humerus rounded ......................... 52
frons much less than one-fifth wider than dis-
tance between lateral margins of rostrum at 46(45). Eye large, flattened; rostrum thick; front coxae
point of insertion of antenna in dorsal view; contiguous or apparently so; body length 5.0-
rostrum at base slightly distad of anterior mar- 12.0 mm ...................................................... 47
gin of eye with short, deep, fine sulcus per- — Eye smaller; front coxae distinctly separated by
pendicular to longitudinal axis of rostrum .... continuous prosternal integument; body length
.................................................. Melanolemma less than 5.0 mm ......................................... 49

39(9). Elytra with humeral angle ............................... 40 47(46). Hind tibia with straight comb of setae on outer
— Elytra with humerus rounded ......................... 42 edge, comb at least as long as width of tibia at
apex; postocular vibrissae set on edge of pro-
thorax; color dorsally dark, vaguely patterned
 Family 131. Curculionidae · 771

at most; corbel open; front coxae contiguous; 54(53). Side of rostrum with tooth-like extension over
body length 5.5-l0.0 mm .............. Tanymecus scrobe just apicad of eye ............. Piscatopus
— Hind tibia without straight comb of setae; — Side of rostrum without extension over scrobe
postocular vibrissae set on knob or rounded ..................................................... Minyomerus
tooth on edge of prothorax; color pale or bril-
liant metallic; corbel various; front coxae con- 55(44). Scrobe dorsal or dorsolateral, indefinite caudad
tiguous or apparently so; body length 6.4-12.0 of antennal insertion; scape in repose not situ-
mm ............................................................... 48 ated in scrobe, usually passing over eye .. 56
— Scrobe lateral; scape in repose situated in scrobe
48(47). Front leg not larger, front femur not more swollen .................................................................... 96
than middle and hind femora; eye large, flat-
tened, oval, slightly diagonal; color pastel gray, 56(55). Corbel closed (Fig. 80c); tarsal claws free; hu-
green, or bluish; many with pollinosity; meral angle well-developed ....................... 57
postocular vibrissae set on a prominent knob — Corbel open or not distinctly closed (Fig. 80a, b);
....................................................... Pachnaeus tarsal claws free or connate; humeral angle
— Front leg larger than middle and hind legs, front rounded ...................................................... 61
femur enlarged; eye moderately convex,
slightly transverse; postocular vibrissae lack- 57(56). Rostrum twice as long as head, abruptly narrowed
ing (or rudimentary in H. opalina); color glossy immediately caudad of antennal insertion where
white or metallic blue, blue-green, green or it is one-half as wide as dorsally .......... Evotus
copper; without pollinosity ... Hadromeropsis — Rostrum shorter, not shaped as above .......... 58

49(46). Tarsal claws fused; rostrum extremely short; 58(57). Scrobe completely dorsal; apex of rostrum with
postocular vibrissae vestigial ......... Isodrusus approximately l6 long setae of graduated
— Tarsal claws free; rostrum short; postocular vibris- lengths; scape moderately thick, reaching to a
sae various ................................................. 50 point midway between eye and prothorax; dor-
sal outline of rostrum and head continuously
50(49). Anterior margin of abdominal ventrites 3, 4, and 5 flat; humeri obsolete; corbel not abruptly dif-
without modification; contour and vestiture ferentiated proximally, enclosed space filled
more or less uniform; mandible without scales; with long scaly setae; body length 4.4-5.6 mm
postocular vibrissae various, well-developed .................................................... Achrastenus
in most species; front legs distinctly to greatly — Scrobe lateral or dorsolateral; corbel closed (Fig.
larger than middle and hind legs (Fig. 78) ...... 80c), corbel plate various .......................... 59
.................................................... Pandeleteius
— Anterior margin of abdominal ventrites 3, 4 and 5 59(58). Scape very thick, short, no longer than thickness
conspicuously modified; postocular vibrissae of rostrum (Fig. 70, 71); apex of rostrum with
present or absent; front legs slightly larger than 20-30 long setae (Fig. 71); mandible with many
middle and hind legs .................................. 51 very long setae directed to mandibular scar
(Fig. 71); body length 5.5-7.5 mm ...................
51(50). Anterior margin of abdominal ventrites 3, 4, and 5 ................................................... Brachystylus
deeply, narrowly sulcate across width of abdo- — Scape longer, extended to or beyond eye ... 60
men; posterior margin of sulcus carinate, right-
angled in female, more rounded in male ........ 60(59). Eye lateral; anterior margin of prothorax without
..................................................... Scalaventer modified setae; humeri oblique, not prominent;
— Anterior margin of abdominal ventrites 3, 4, and 5 base of each elytron arcuately produced be-
shallowly sulcate across width of abdomen or tween scutellum and stria 6; corresponding area
medially only; posterior edge rounded .......... on prothorax depressed, with different vestiture;
.................................................. Pandeleteinus body length 7.5-12.5 mm ................... Compsus
— Eye slightly encroaching on dorsum; anterior mar-
52(45). Tarsal articles 1, 2, and 3 on ventral surface with gin of prothorax with a row of 20 or more very
complete pad of dense fine setae; scrobe right- fine long setae of graduated lengths directed
angled, reaching ventral surface, glabrous ... toward lower edge of eye; humeri quadrate, promi-
........................................................ Isodacrys nent; base of elytra very slightly produced; body
— Tarsal articles 1, 2, and 3 on ventral surface lack- length 13.5-18.0 mm ......................... Diaprepes
ing complete pads; at most with small pad at
apex of tarsal article 3; pads replaced with 61(56). Funicle with six articles; tarsal claws free ..... 62
spines or long recumbent setae or neither; — Funicle with seven articles; tarsal claws various
scrobe not right-angled, squamate ............ 53 .................................................................... 63

53(52). Dorsal surface with very fine long setae, recum- 62(61). Prothorax lacking median sulcus; surface punc-
bent or semi-erect, setae longer on sides and tate; elytral intervals flat; corbel plate large,
ventral surface; prothorax distinctly wider than glabrous, oval; body length 5.8-6.8 mm ........
long ............................................ Miloderoides ......................................................... Agraphus
— Dorsal surface with inconspicuous short, recum- — Prothorax with median longitudinal sulcus; elytral
bent setae, similar on sides and ventral sur- intervals 3, 5, and 7 more prominent; corbel
face; prothorax distinctly longer than wide ... plate indeterminate; body length 6.2 mm .......
.................................................................... 54 .................................................... Paragraphus

63(61). Tarsal claws connate ..................................... 64

772 · Family 131. Curculionidae

— Tarsal claws free ............................................ 84 twice as long; scrobe not reaching eye; scape
slightly arcuate; abdominal ventrite 2 as long
64(63). Eye large, almost touching prothorax; anterior mar- as 3 plus 4 combined .................... Dysticheus
gin of prothorax with postocular lobe; corbel — Apical comb on hind tibia made up of spines al-
narrowly closed; elytra very convex, sides most uniform in length and coarseness; other
greatly rounded .................. Pseudocneorhinus characters various ...................................... 73
— Eye smaller, prothorax without postocular lobe;
corbel various; elytra various ..................... 65 73(72). Dorsal and ventral margins of scrobe distinctly
defined up to eye, scrobe a completely flat
65(64). Femora with large tooth on inner surface; dorsum bottomed channel; elytra with erect, stout,
with slender, decumbent setae only, or with moderately long, acute setae distinctly longer
very small metallic scales; scrobe more lateral than scales composing decumbent vestiture;
than dorsal ...................................... Phyllobius dorsal surface of head and rostrum in lateral
— Femora without a tooth; other characters various view interrupted by erect, stout, acute setae;
.................................................................... 66 abdominal ventrite 2 shorter than ventrites 3
and 4 combined; scape strongly arcuate .......
66(65). Rostrum in dorsal view more or less rectangular ........................................................... Eucyllus
in outline; entire body and appendages — Dorsal margin of scrobe indicated up to slightly
densely scaled; scrobe completely dorsal .... distad of eye, ventral margin either merging
........................................................ Aphrastus with dorsal margin distad of eye or else sepa-
— Rostrum in dorsal view not rectangular ......... 67 rated from dorsal margin by raised, convex area;
elytra with suberect, short, wide, rounded
67(66). Mandible with more than five setae; elytra glo- scales slightly longer than those composing
bose; body shiny black with white setae ....... decumbent vestiture; dorsal surface of head
.............................................................. Omias and rostrum in lateral view slightly interrupted
— Mandible with three setae; elytra more elongate; by suberect, short, rounded scales; abdominal
body color and vestiture various ............... 68 ventrite 2 as long as ventrites 3 and 4 com-
bined; scape slightly arcuate ...... Thinoxenus
68(67). Tarsal claws free at base; femora with minute tooth
on inner edge distally; vestiture of sparse, 74(69). Female with hind tibia unarmed at apex; scrobe
erect, fine setae; without scales .... Stomodes situated more dorsally on rostrum, short, some-
— Tarsal claws connate at base ......................... 69 what convergent posteriorly ..................... 75
— Female having hind tibia armed with one or more
69(68). Inner apical surface of hind tibia adjacent to tar- small spines or teeth at apex ..................... 76
sal insertion clothed with dense, round scales;
funicle and dorsal surface of tarsi clothed with 75(74). Rostrum longer than head, antenna inserted at
round scales; tarsi on ventral surface with distance from anterior margin of eye about
coarse setae ............................................... 70 twice diameter of eye ................... Thricolepis
— Inner apical surface of hind tibia adjacent to tar- — Rostrum about as long as head, antenna inserted
sal insertion glabrous; articles 4 to 7 of funicle at distance from anterior margin of eye less
and dorsal surface of tarsi clothed with elon- than the diameter of eye ............... Peritelinus
gate, rather fine, hairlike scales; pubescence
on ventral surface of tarsi fine ................... 74 76(74). Female with hind tibia armed with one spine or
small tooth at apex ..................................... 77
70(69). Elytra and prothorax clothed with long, very fine, — Female with hind tibia armed with two or more
erect pile, pile longer on lateral portion of body, spines or small teeth at apex ..................... 80
at least twice as long as width of an interval;
dorsal surface of prothorax in lateral view 77(76). Scrobes lateral, not convergent posteriorly on
slightly gibbous ....................... Eucilinus (part) dorsal surface of rostrum ........................... 78
— Elytra and prothorax with short, stout setae, not — Scrobes dorsal, more or less convergent posteri-
longer than width of an interval; dorsal surface orly on dorsal surface of rostrum ............... 79
of prothorax nearly flat in lateral view ....... 71
78(77). Antenna with scape much shorter than funicle
71(70). Dorsal comb of setae on apex of hind tibia about ...................................................... Aragnomus
twice as long as apical comb and situated al- — Antenna with scape as long as funicle .............
most at 45 degree angle to longitudinal axis of .................................................... Geodercodes
tibia; abdominal ventrite 2 as long as ventrites
3 and 4 combined; ventral and dorsal margin of 79(77). Hind tibia at narrowest point in apical third at least
scrobe not distinct at distance one-half length two-thirds as wide as widest point of tibia at
of eye distad of front margin of eye .............. apex; epistoma separated from rostrum by fine,
...................................................... Rhypodilius moderately raised, acute carina; in dorsal ante-
— Dorsal comb of setae on apex of hind tibia about rior view dorsal margins of rostrum distinctly,
as long as apical comb and almost parallel to broadly emarginate above point of antennal
longitudinal axis of tibia; other characters vari- insertion, greatly expanded toward apical re-
ous .............................................................. 72 gion, in apical region subparallel sided,
subparallel region about one-third wider than
72(71). Apical comb on hind tibia with anterior spines narrowest point between points of antennal
very short and coarse and posterior spines on insertions ........................................ Anchitelus
ascending portion distinctly finer and almost
 Family 131. Curculionidae · 773

— Hind tibia at narrowest point in apical third less — Scrobe superior, very short and deep, not reach-
than one-half as wide as tibia at widest point at ing eye, not directed below; only front and
apex; epistoma indistinctly separated from ros- middle tibiae with apical spine or tooth ..... 90
trum; in dorsal anterior view dorsal margins of
rostrum straight, slightly diverging from behind 88(87). Appressed vestiture of dense radiate-pectinate
point of antennal insertions to apex of rostrum, scales; surface encrusted ........ Chaetechidius
subparallel region distad of points of antennal — Appressed vestiture of imbricate, oval scales, or
insertions only slightly wider than narrowest scales concealed by encrustation ............. 89
point between points of antennal insertions .
.................................................. Orthoptochus 89(88). Vestiture short, suberect, broad, spatulate scale-
like setae; eye with more than 5 facets across
80(76). Female with hind tibia armed with three spines or greatest width; elytra more elongate, parallel-
small teeth at apex; funicle with article 7 dis- sided; Oregon ........................ Cathormiocerus
tinctly longer than wide ............................. 81 — Vestiture of longer, suberect, fine hair-like setae;
— Female with hind tibia armed with two small spines eye with less than 5 facets across greatest
or teeth at apex .......................................... 82 width; elytra more rounded, subglobose; south-
ern United States .............. Trachyphloeosoma
81(80). Suture between abdominal ventrites 1 and 2 al-
most straight; scape with coarse, elongate 90(87). Rostrum short and broad; prementum not emar-
scales; elytra with obvious (at 56X) erect, spatu- ginate on apical margin; only labial palpi visible
late scales .................................. Stenoptochus ............................................. Pseudocercopeus
— Suture between abdominal ventrites 1 and 2 ar- — Rostrum long, narrow, almost cylindrical;
cuate; scape with fine setae; elytra (at 56X) prementum broadly emarginate; both labial and
lacking obvious erect scales, at most with very maxillary palpi visible .................. Cercopedius
fine, short, subdecumbent setae ..................
...................................................... Peritelopsis 91(84). Antennal scape stout, short, not exceeding eye
.................................................................... 92
82(80). Antennal funicle with article 7 distinctly longer — Antennal scape reaching beyond anterior margin
than wide .................................... Nemocestes of prothorax ................................................ 94
— Antennal funicle with article 7 about as wide as
long ............................................................. 83 92(91). Eye not prominent; pronotum smooth, polished;
with moderate punctures becoming larger and
83(82). Suture between abdominal ventrites 1 and 2 al- closer laterad; body length 4.0-4.5mm ..........
most straight; prothorax with coarse, erect ...................................................... Lupinocolus
setiform scales more or less uniformly distrib- — Eye very prominent; pronotum with pronounced
uted over dorsal surface, lacking a distinct sculpture .................................................... 93
clump at anterior margin ............ Paraptochus
— Suture between abdominal ventrites 1 and 2 ar- 93(92). Pronotum with large, irregularly spaced, flat-
cuate; prothorax with coarse, setiform scales topped cylindrical protuberances with central
subdecumbent on dorsal surface except at an- depression set with a seta; integument and
terior margin with elongate clump of about 10 vestiture black; body length 8.3-9.6 mm ........
erect scales on either side of middle line ..... ................................................. Agasphaerops
....................................................... Peritelodes — Pronotum with contiguous nodules each slightly
depressed caudally; vestiture of slender, elon-
84(63). Antenna with scape with vestiture of fine setae gate coppery, green, or reddish metallic
and round flat scales; body size small, 2.3-4.5 scales/setae on dorsum; oval, opalescent or
mm ............................................................... 85 metallic scales in groups forming pattern on
— Antenna with scape with vestiture of fine setae sides of prothorax, elytra; elytra with intervals
only or with at most a few scattered elongate, 3 and 5 slightly elevated; body length 7.5-9.0
recumbent scales intermixed; body size larger, mm .................................................. Hormorus
3.2-13.0 mm, most larger than 5.0 mm ........ 91
94(91). Eye small, round, prominent, almost touching pro-
85(84). Epistoma large, distinct, occupying approximately thorax; rostrum greatly narrowed between an-
half the anterior margin of rostrum, triangular, tennal insertions, less than half the width of
limited by distinct carina ............................ 86 rostrum between outer edge of pterygia; scape
— Epistoma very small, indistinct ....................... 87 arcuate; body length 3.2-4.0 mm ...... Agronus
— Eye moderate; rostrum wider between pterygia;
86(85). Scrobe dorsolateral, usually reaching and often scape arcuate or straight; body length various
enclosing eye; all tibiae with single strong, al- .................................................................... 95
most horizontal apical tooth ... Trachyphloeus
— Scrobe dorsal, very short and deep, not reaching 95(94). Elytra of most species with few sparse scales or
eye; front and middle tibiae with single, strong, no scales; if densely squamose, pronotum simi-
almost horizontal apical tooth; hind tibia with larly squamose or not and nodulate; femora with
pair of short vertical apical spines ................. or without a tooth on inner surface; body length
........................................................ Cercopeus 4.0-13.0 mm .............................. Otiorhynchus
— Elytra and prothorax densely squamose; femora
87(85). Scrobe lateral, long, passing backward and be- not dentate; body length 4.9-6.6 mm (Figs. 72-
low the lower angle of eye; all tibiae with apical 73) ................................................... Sciopithes
spine or tooth ............................................. 88
774 · Family 131. Curculionidae

96(55). Tarsal claws connate ..................................... 97 — Eyes lateral; humeri rounded (except Ericydeus,
— Tarsal claws free .......................................... 105 southwestern United States only); rostrum with
longitudinal sulcus or impressed line reaching
97(96). Corbel closed (Fig. 80c), corbel plate covered from interantennal line to head, continuing or
with shiny round scales; epistoma concave, not with fine impressed line reaching beyond
steeply angled, posterior margin conspicu- eyes, in some genera to occiput .............. 116
ously carinate; body length 4.5-8.0 mm ........
....................................................... Philopedon 107(106). Rostrum separated from frons by distinct trans-
— Corbel open (Fig. 80a) .................................... 98 verse sulcus or depression; dorsal aspect of
tarsal articles with scales; tarsi clothed ven-
98(97). Elytra with humerus prominent ...................... 99 trally with “coarse setae” (stiff decumbent se-
— Elytra with humerus rounded ....................... 100 tae) ............................................................ 108
— Rostrum not separated from frons when viewed
99(98). Rostrum with a conspicuous curved, glabrous cal- laterally, frons lacking tubercle above eye; max-
losity extending between the antennal inser- illae concealed by mentum ...................... 110
tions and paralleling the glabrous epistoma,
surface squamose between the glabrous areas; 108(107). Hind tarsus with article 3 bilobed and wider than
elytral scales elongate ............... Pachyrhinus article 2 ...................................... Stereogaster
— Rostrum without a glabrous callosity; elytral scales — Hind tarsus with article 3 not wider than article 2
round; some species with a minute tooth on .................................................................. 109
fore femur ..................................... Polydrusus
109(108). Mentum not concealing maxillae; frons above
100(98). Head constricted dorsally behind eyes; eyes each eye with distinct tubercle which conceals
small, prominent; epistoma marked by fine ca- eye in dorsal view; elytra with inconspicuous,
rina; scape extended to posterior margin of eye sparse, short, suberect spatulate scales or pa-
................................................... Strophosoma pillae; body length 2.5-3.0 mm ...... Calyptillus
— Head not constricted behind eyes; other charac- — Mentum concealing maxillae; frons lacking dis-
ters various ............................................... 101 tinct tubercle above eye; eye visible in dorsal
view; elytra with suberect, fine, acute setae;
101(100). Epistoma not defined; body form narrow elongate body length greater than 3.0 mm ...................
.................................................... Brachyderes .................................................. Cryptolepidus
— Epistoma carinate or not (Barypeithes); eye flat-
tened; humeri rounded ............................. 102 110(107). Head conspicuously constricted behind eye;
eyes very prominent ........ Bradyrhynchoides
102(101). Vestiture of several sizes of very fine setae, lack- — Head not constricted behind eye; eyes moder-
ing scales; color castaneous; epistoma minute, ately convex but not protuberant ............ 111
almost undetectable .................... Barypeithes
— Vestiture of scales with or without setae; epistoma 111(110). Mesepimeron triangular, anterior margin running
distinct ...................................................... 103 straight to angle between elytron and peduncle
of mesothorax, mesepisternum not touching
103(102). Femora with short, broad tooth ......... Sciaphilus side margin of elytron; scutellum well devel-
— Femora without tooth ................................... 104 oped; metepisternal suture complete ...... 112
— Mesepimeron short trapezoidal, anterior margin
104(103). Striae on disc of elytra almost as wide as inter- running to side margin of elytron,
vals; striae composed of large foveae; each mesepisternum touching elytron on broad con-
interval with row of long stiff, erect setae; also tact ............................................................ 113
with small appressed elongate scales, 2 to 4
abreast; elytra rotund; length less than 3.0 mm 112(111). Humeri well-developed, quadrate; scales on body
.................................................. Brachysomus sparse, not imbricate, prothorax and elytra lack-
— Elytral striae fine; stria 10 distinct; elytra narrow, ing erect scales or setae; article 7 of funicle
elongate; length more than 3.0 mm ................ distinctly longer than wide ........... Lachnopus
........................................................ Mitostylus — Humeri rounded; scales on body dense, imbri-
cate; prothorax and elytra with short, suberect
105(96). Apex of rostrum with keel across entire width of scales and setae; article 7 of funicle more or
rostrum, keel forming posterior edge of less wider than long .......................... Omileus
epistoma; humeri distinct, quadrate; corbel
semi-closed (Fig. 80b); mandibular cusp situ- 113(111). Metepisternal suture complete; rostrum lacking
ated at the apex of an anterior projection of impressions on dorsal surface; eye distant from
the mandible; front coxae very narrowly sepa- anterior margin of prothorax by more than half
rated ............................................. Polydacrys its greatest diameter; base of elytra as wide as
— Epistoma indistinct, without carinate or keeled base of prothorax ... Stamoderes and Amotus
margin (except Glaphyrometopus, recognized — Metepisternal suture obliterated in basal half; ros-
by frons with deep, broad transverse concav- trum on dorsal surface with distinct, lateral, lon-
ity between eyes); humeri mostly rounded; gitudinal impression; eye separated from ante-
other characters various .......................... 106 rior margin of prothorax by half its diameter or
less; base of elytra distinctly wider than base
106(105). Eyes partly encroaching on head; epistoma of prothorax .............................................. 114
poorly defined; humeri rounded (except
Lachnopus, southern Florida only) ........... 107
 Family 131. Curculionidae · 775

114(113). Dorsal surface of tarsi clothed with broad, setae (except some P. viridis); corbel plate
rounded scales; tarsi on ventral surface with small, squamate ........................... Phacepholis
coarse setae on anterior part of articles 1 to 3 — Funicle with articles 2 and 1 with various ratios;
.................................................. Graphorhinus other characters various .......................... 123
— Dorsal surface of tarsi clothed with fine setae or
elongated scales; tarsi on ventral surface 123(122). Scutellum glabrous, glossy; antennal funicle with
clothed with fine, dense setae ................. 115 article 2 approximately as long as article 1; cor-
bel plate narrow ......................... Atrichonotus
115(114). Rostrum (excluding mandibles) in dorsal view from — Scutellum squamate; antennal funicle with article
anterior margin of eye to apex slightly longer 2 approximately 1.5 to 2.0x longer than article
than greatest width in apical region; antenna 1; corbel plate absent or present ..................
with short, narrow scales and sparse, long, fine ........................... Naupactus and Pantomorus
setae; prosternum lacking two close adjacent
tubercles behind front coxae ......... Epicaerus
— Rostrum (excluding mandibles) in dorsal view from
anterior margin of eye to apex distinctly longer CLASSIFICATION OF THE NEARCTIC ENTIMINAE
than greatest width in apical region; antenna
with both short and long, fine setae; prosternum 41. Agraphini Horn 1876
with two closely adjacent tubercles behind
front coxae ..................................... Barynotus
Agraphus Say 1831, 1 sp., A. bellicus (Say 1831), eastern coastal
116(106). Rostrum trisulcate (Fig. 77) ........................... 117 United States from New York south to Florida. Adults are asso-
— Rostrum with median sulcus only ................ 118 ciated with sandy habitats along the Atlantic coast and in central
Florida.
117(116). Frons with deep, broad transverse concavity be-
tween eyes; rostrum with all three longitudinal Agraphus Schoenherr 1834; not Say 1831
sulci reaching transverse concavity; humerus
absent; length 4.5-5.0 mm ............................. Paragraphus Blatchley 1916, 1 sp., P. setosus Blatchley 1916, Florida.
........................................... Glaphyrometopus
— Frons without transverse impression; rostrum with
longitudinal sulci extending as far as between 42. Alophini LeConte 1876
eyes (Fig. 77); humerus prominent; length 10.0-
18.0 mm ........................................... Ericydeus Acmaegenius LeConte 1876, 2 spp., A. granicollis Van Dyke 1927,
Wyoming, and A. hylobinus LeConte 1876, Idaho and Oregon.
118(116). Rostrum with very deep, wide median longitudi-
nal “crater” from apex to at least middle; scape
very thick, slightly curved; head with fine lon- Lepidophorus Kirby 1837, 10 spp., generally distributed in the west-
gitudinal impressed line; funicle with articles 1 ern United States and Canada, north into Alaska, Yukon Terri-
and 2 equal in size; elytral stria 10 evanescent
tory and the Northwest Territories, including L. setiger Hamilton
or absent beyond level of hind coxa ............
....................................................... Platyomus 1895, generally distributed in the eastern United States from New
— Rostrum not sulcate from apex; scape not thick; York and Ohio south into Virginia and Tennessee. Adults of all
straight; other characters various ............ 119 species are flightless and collected in leaf litter or under rocks. The
western species are often collected at high elevations. See Buchanan
119(118). Epistoma conspicuous, very wide, occupying
most of anterior edge of rostrum; prothorax and (1936a) to separate the species. This genus is questionably dis-
elytra with very irregularly shaped, randomly tinct from Dirotognathus (Tropiphorini), we could find no charac-
situated large foveae .......................... Artipus ter to reliably distinguish the two.
— Epistoma inconspicuous, occupying half or less
Lophalophus LeConte 1876
of anterior edge of rostrum; prothorax and
elytra with only regular sculpture ............ 120
Plinthodes LeConte 1876, 2 spp., P. foveirostris (Chittenden 1925),
120(119). Head and rostrum in dorsal view forming a flat Ohio, North Carolina, Tennessee, and Virginia, and P. taeniatus
triangle two times wider than apex of rostrum;
(LeConte 1857), British Columbia, Oregon and Washington.
epistoma small, inconspicuous; rostral longitu-
dinal sulcus fine, reaching pronotum; corbel
plate narrow ..................................... Aramigus Trichalophus LeConte 1876, 8 spp., generally distributed in the
— Head and rostrum less triangular, not flat .... 121 western United States north into Canada and Alaska, then east
across the north to Manitoba and Ontario. Adults are collected
121(120). Elytral stria 10 complete, but sometimes con-
cealed by vestiture, separate from stria 9; cor- on several different kinds of plants. The genus needs revision.
bel open; some species resembling a Sitona or See Hatch (1971) to separate some of the species.
Tanymecus, others with more slender rostrum
and tuberculate prothorax ......... Mesagroicus
Triglyphulus Cockerell 1906, 2 spp., T. ater (LeConte 1876) and T.
— Elytral stria 10 evanescent or absent beyond hind
coxae ........................................................ 122 nevadensis Van Dyke 1938, California, Oregon, Washington and
Nevada.
122(121). Funicle with article 2, l.0-1.6x longer than article Triglyphus LeConte 1876; not Loew 1840; not Fraas 1866
1; scales of pronotum in clusters surrounding
776 · Family 131. Curculionidae

43. Anypotactini Champion 1911 Calomycterus Roelofs 1873, 1 sp., C. setarius Roelofs 1873, gener-
ally distributed in the eastern United States west to Iowa and
Polydacrys Schoenherr 1834, 1 sp., P. depressifrons Boheman 1840, Nebraska; adventive. Adults generally feed on foliage.
southern Texas. Synolobus Faust 1886

44. Brachyderini Schoenherr 1826 Myllocerina Pierce 1913

Brachyderes Schoenherr 1823, 1 sp., B. incanus (Linnaeus 1758), Myllocerus Schoenherr 1823, 1 sp., M. undatus Marshall 1916,
northeastern United States; adventive. Florida; adventive.
Thylacites Germar 1817 Macrocorynus Schoenherr 1823
Brachylophus Fischer von Waldheim 1829 (valid subgenus) Hyperstylus Roelofs 1873
Poloposes Gistel 1848 Exmyllocerus Voss 1937 (valid subgenus)
Eumonima Gistel 1856 Pachymyllocerus Voss 1937 (valid subgenus)
Gastraspis Flach 1907 (valid subgenus) Isomyllocerus Marshall 1954 (valid subgenus)
Sulciurus Flach 1907 Pseudocanoixus Voss 1958 (valid subgenus)
Echopus Desbrochers 1909 Allomycterops Voss 1959 (valid subgenus)
Calomyllocerus Voss 1959 (valid subgenus)
Strophosoma Billberg 1820, 1 sp., S. melanogrammum (Forster 1771), Mylloceroversus Hoffmann 1961 (valid subgenus)
northeastern United States and adjacent southern Canada, also Corigetellus Hoffmann 1964 (valid subgenus)
Washington and British Columbia; adventive. Adults feed gen-
erally on foliage. Neoptochus Horn 1876, 1 sp., N. adspersus (Boheman 1834), Florida,
Strophosomus Schoenherr 1823 Georgia and South Carolina.
Strophosomum Gistel 1856
Leucostrophus Flach 1907; not Rothschild and Jordan 1903 47. Eudiagogini LeConte 1874
Morphostrophus Flach 1907 (valid subgenus)
Aracanthus Say 1831, 1 sp., A. pallidus Say 1831, generally distrib-
45. Cneorhinini Lacordaire 1863 uted in the southeastern United States west to Iowa and Texas.

Philopedon Schoenherr 1826, 1 sp., P. plagiatum (Schaller 1783), Colecerus Schoenherr 1840, 2 spp., C. dispar (LeConte 1874), Ari-
Newfoundland, New Brunswick, Nova Scotia and Prince Ed- zona and Texas, and C. marmoratus (Horn 1876), Texas. Adults
ward Island; adventive. This species appears to be found in sandy feed on foliage of various Fabaceae, especially Prosopis (mesquite),
habitats. Mimosa and Acacia. See LeConte and Horn (1876) to separate the
Philopedum Agassiz 1846 species.
Dactylorhinus Tournier 1876 Coleocerus Agassiz 1846
Dactylorrhinus Rye 1878 Coleocerus Gemminger and Harold 1871; not Agassiz 1846
Bathyris LeConte 1874
46. Cyphicerini Lacordaire 1863
Eudiagogus Schoenherr 1840, 3 spp., generally distributed in the
Cyphicerina Lacordaire 1863 southeastern United States west to Texas, also Arizona and Cali-
fornia. Adults feed on foliage of species of Sesbania (Fabaceae);
Cyrtepistomus Marshall 1913, 1 sp., C. castaneus (Roelofs 1873), larvae feed on nitrogen-fixing root nodules in the soil (Kovarik
generally distributed in the eastern and southeastern United States and Burke 1989). See Warner (1979) to separate the species.
west to Texas; adventive. Adults generally feed on foliage. This
species is known as the “Asiatic oak weevil”. 48. Eustylini Lacrodaire 1863

Myosides Roelofs 1873, 1 sp., M. seriehispidus Roelofs 1873, Mary- Achrastenus Horn 1876, 1 sp., A. griseus Horn 1876, Texas.
land, Massachusetts and Connecticut; adventive.
Brachystylus Schoenherr 1824, 2 spp., B. sayi Alonso-Zarazaga 1994,
Oedophrys Marshall 1941, 1 sp., O. hilleri (Faust 1889), eastern southeastern United States, west to Texas, and B. microphthalmus
United States from Connecticut and Pennsylvania south into Champion 1911, southern Texas. Adults of B. sayi feed on foli-
Virginia; adventive. Adults feed generally on foliage. age of Diospyros (persimmon; Ebenaceae).
Syntomostylus Scudder 1893
Acanthotrachelina Marshall 1944
 Family 131. Curculionidae · 777

Compsus Schoenherr 1823, 1 sp., C. auricephalus (Say 1824), south- Asteraceae). See Van Dyke (1935b) to separate the species. This
eastern United States west to Texas and Colorado. Adults gener- genus is questionably distinct from Amotus (Tanymecini). We can
ally feed on foliage. find no characters to reliably distinguish these two genera.
Callopistus Say 1831
Stereogaster Van Dyke 1936, 1 sp., S. globosa Van Dyke 1936, Cali-
Diaprepes Schoenherr 1823, 1 sp., D. abbreviatus (Linnaeus 1758), fornia.
adventive, Florida. Adults feed on foliage and are pests of citrus
(Woodruff 1968, 1979). Trigonoscuta Motschulsky 1853, 64 spp., western United States
and adjacent southern Canada. Adults are associated with sandy
49. Geonemini Gistel 1856 habitats and feed on various plants. In a posthumous publica-
tion, Pierce (1975) described all but a few of the species as well as
Barynotus Germar 1817, 3 spp., northeastern United States and numerous subspecies. Most of these taxa are of questionable
eastern Canada, also British Columbia; adventive. Adults gener- validity and need reassessment. See Pierce (1975) to separate the
ally feed on foliage. See Brown (1950) to separate the species. species and subspecies.
Merionus Dejean 1821 Panormus Casey 1888 (valid subgenus)
Kissodontus Desbrochers 1909 Eremocatoecus Pierce 1975 (valid subgenus)
Nesocatoecus Pierce 1975 (valid subgenus)
Bradyrhynchoides Pierce 1913, 1 sp., B. constrictus Pierce 1913, Texas.
50. Hormorini Horn 1876
Calyptillus Horn 1876, 1 sp., C. cryptops Horn 1876, New Mexico,
Colorado, Kansas and Nebraska. Agasphaerops Horn 1876, 1 sp., A. niger Horn 1876, California,
Oregon, Washington and British Columbia. Adults are associ-
Cryptolepidus Van Dyke 1936, 7 spp., Arizona, California and Ne- ated with lilies (Liliaceae).
vada. Adults are associated with Artemisia (sagebrush; Asteraceae)
and other shrubs. See Ting (1940) to separate the species. Hormorus Horn 1876, 1 sp., H. undulatus (Uhler 1856), generally
Lepidopus Van Dyke 1936; not Gouan 1770; not Dana 1852 distributed in the northeastern United States and southern
Pseudoeucyllus Tanner 1950 Canada.

Epicaerus Schoenherr 1834, 11 spp., generally distributed in the Lupinocolus Van Dyke 1936, 1 sp., L. blaisdelli Van Dyke 1936,
United States. Adults generally feed on foliage. The genus needs California and Nevada.
revision. See Pierce (1913) to separate most of the species.
Epagrius Schoenherr 1840
Diorynotus Sharp 1891 (valid subgenus) 51. Naupactini Gistel 1856
Cacochromus Sharp 1891
Bradyrhynchus Sharp 1891 Aramigus Horn 1876, 1 sp., A. tesselatus (Say 1824), central United
Melbonus Casey 1895 States; adventive. Adults are pests on alfalfa and some other
Epagriopsis Champion 1911 crops in Argentina (Lanteri and Díaz 1994). Lanteri and Díaz
(1994) describe a number of morphotypes of A. tesselatus.
Graphorhinus Say 1831, 1 sp., G. vadosus Say 1831, Texas, Colorado, Aomopactus Jekel 1876
Kansas, Wyoming and Missouri.
Graphorhinus Schoenherr 1833; not Say 1831 Artipus Sahlberg 1823, 1 sp., A. floridanus Horn 1876, Florida.
This species is a pest of the Florida citrus industry (McCoy et al.
Lachnopus Schoenherr 1840, 3 spp., southern Florida. Lachnopus 1985). (Volume 1, Color Fig. 2)
floridanus Horn 1876 is native and feeds on foliage of various Artipus Schoenherr 1823; not Sahlberg 1823
plants. Lachnopus argus (Reiche 1840) and L. hispidus (Gyllenhal
1834) are adventive species which doubtfully are established in Atrichonotus Buchanan 1939, 1 sp., A. taeniatulus (Berg 1881),
Florida (Anderson 1993a). southeastern United States west to Texas; adventive. Adults feed
Menoetius Dejean 1821 on foliage of various plants, but most frequently Fabaceae (Lanteri
Ptilopus Schoenherr 1823 and O’Brien 1990).
Floresianus Hustache 1939
Omileus Horn 1876, 1 sp., O. epicaeroides Horn 1876, Texas. Floresianellus Lanteri 1981

Stamoderes Casey 1888, 2 spp., S. lanei (Van Dyke 1935), British Ericydeus Pascoe 1880, 2 spp., E. lautus (LeConte 1856) Arizona,
Columbia, Oregon and Washington, and S. uniformis Casey 1888, California, Colorado, Utah and New Mexico, and E. placidus (Horn
California. Adults of S. lanei are found on Artemisia (sagebrush;
778 · Family 131. Curculionidae

1876), Arizona and California. Adults appear to be associated 53. Ophryastini Lacordaire 1863
with Fabaceae. See Lanteri (1995) to separate the species.
Ophryastes Germar 1829, 35 spp., generally distributed in western
Glaphyrometopus Pierce 1913, 1 sp., G. ornithodorus Pierce 1913, Texas. United States and adjacent southern Canada. Adults are flightless
and are associated with various arid habitat shrubs, mostly in the
Mesagroicus Schoenherr 1840, 9 spp., generally distributed. Adults family Asteraceae, but also Larrea tridentata (DC.) Cov. (creosote
generally feed on foliage. See Buchanan (1929a) and Burke (1960) bush; Zygophyllaceae) and Atriplex (saltbush; Chenopodiaceae).
to separate the species. See Kissinger (1970) to separate the species.
Mesagroecus Agassiz 1846 Ophryastes Say 1831; not Germar 1929
Lepidocricus Pierce 1910 Dystirus Pascoe 1872
Eupagoderes Horn 1876
Naupactus Dejean 1821, 4 spp., N. godmanni (Crotch 1867), N. leucoloma Caccophryastes Sharp 1891
Boheman 1840, N. minor (Buchanan 1942) and N. peregrinus (Buchanan Tosastes Sharp 1891
1939), generally distributed in the southeastern United States; ad- Amydrogmus Pierce 1913
ventive. Adults are considered pests and feed on foliage of various
plants. See Lanteri and Marvaldi (1995) and Lanteri (1986) to sepa- Sapotes Casey 1888, 2 spp., S. longipilis Van Dyke 1934 and S.
rate the species. We have not seen specimens of Pantomorus pallidulus puncticollis Casey 1888, Arizona, New Mexico and Texas. Adults
Emden 1936 and thus cannot separate it from Naupactus. are associated with arid habitat shrubs, especially Larrea tridentata
Asynonychus Crotch 1867 (DC.) Cov. (creosote bush; Zygophyllaceae). See Van Dyke (1934)
Mimopactus Jekel 1875 to separate the species.
Archopactus Heller 1921
Graphognathus Buchanan 1939 54. Otiorhynchini Schoenherr 1826

Pactorrhinus Ancey 1881, 1 sp., P. grisescens Ancey 1881, Arizona. Agronus Horn 1876, 3 spp., California, Oregon, Montana, Alberta
This genus and species are unknown to us but likely refer to and British Columbia. See Buchanan (1929b) to separate the spe-
Ericydeus lautus (LeConte 1856). It does not appear in the key. cies.

Pantomorus Schoenherr 1840, 1 sp., P. pallidulus Emden 1936, Texas. Otiorhynchus Germar 1822, 14 spp., generally distributed; all ad-
We have not seen specimens of Pantomorus pallidulus Emden 1936 ventive. Adults and larvae generally feed on a variety of plants.
and thus cannot separate it from Naupactus. This genus includes a number of common pest species; O. ovatus
Pantoplanes Schoenherr 1840 (Linnaeus 1758), the strawberry root weevil, and O. sulcatus (Fab-
Symmathetes Schoenherr 1847 ricius 1775), the black vine weevil. Adults are flightless and a
Pantopactus Jekel 1876 number of species have very restricted distributions in eastern
Athetetes Pascoe 1886 coastal North America. See Warner and Negley (1976) to separate
Antelmia Hustache 1919 the species. Alonso-Zarazaga and Lyal (1999:168-170) list 105
Pseudeudius Voss 1934 valid subgeneric names, not including synonyms. For brevity,
these are not repeated here.
Phacepholis Horn 1876, 5 spp., central United States. See Lanteri
(1990) to separate the species. Sciopithes Horn 1876, 6 spp., California, Oregon, Washington
and British Columbia. Adults feed generally on foliage. See Van
Platyomus Sahlberg 1823, 1 sp., P. flexicaulis (Schaeffer 1905), south- Dyke (1935b) to separate the species.
ern Texas.
Platyomus Schoenherr 1823; not Sahlberg 1823 55. Peritelini Lacordaire 1863
Pseudocyphus Schaeffer 1905
Eustylomorphus Pierce 1915 Anchitelus Van Dyke 1936, 1 sp., A. alboviridis Van Dyke 1936,
Pachyus Kuschel 1955 California.

Aragnomus Horn 1876, 3 spp., western United States.

52. Omiini Shuckard 1840
Dysticheus Horn 1876, 2 spp., D. insignis Horn 1876 and D.
Omias Germar 1817, 6 spp., California, Oregon, Washington, rotundicollis Van Dyke 1953, California. See Van Dyke (1953) to
Idaho and British Columbia. Adults generally feed on foliage. separate the species.
See Hatch (1971) to separate the species.
Mylacus Boheman 1843 Eucilinus Buchanan 1926, 3 spp., California, Utah, Idaho and
Omorus Gistel 1856 Arizona. See Kissinger (1960) to separate the species.
 Family 131. Curculionidae · 779

Thysanocorynus Van Dyke 1938 on the foliage of various plants. See Blatchley and Leng (1916) to
separate the species.
Eucyllus Horn 1876, 8 spp., generally distributed in the far west- Aphrastus Schoenherr 1833; not Say 1831
ern United States. Adults are flightless and mostly nocturnal and Micronychus Provancher 1877; not Motschulsky 1861
are associated with various shrubs in arid habitats. See Pelsue and
Sleeper (1972) to separate the species. Evotus LeConte 1874, 1 sp., E. naso (LeConte 1857), northwestern
United States and adjacent Canada. Adults feed on foliage of
Geodercodes Casey 1888, 1 sp., G. latipennis Casey 1888, California, various plants.
Oregon, Washington, Idaho, Montana and British Columbia.
Phyllobius Germar 1824, 3 spp., northeastern United States and
Nemocestes Van Dyke 1936, 9 spp., generally distributed in the far adjacent Canada; adventive. Adults feed on foliage of various
western United States and adjacent British Columbia, including N. trees. No key to the three species in North American species is
horni (Van Dyke 1936) also in Michigan, New York, Wisconsin, available but the status of P. glaucus as established is undeter-
Ontario, Quebec, New Brunswick and Nova Scotia. Adults are flight- mined. See Côté and Bright (1995) to separate two of the species.
less and mostly nocturnal and feed on foliage of various plants. See Alonso-Zarazaga and Lyal (1999:173-174) list 19 valid subgeneric
Van Dyke (1936a) and Hatch (1971) to separate the species. names, not including synonyms. For brevity, these are not re-
peated here.
Orthoptochus Casey 1888, 1 sp., O. squamiger Casey 1888, California.
57. Polydrusini Schoenherr 1823
Paraptochus Seidlitz 1868, 3 spp., California, Oregon and British
Columbia. See Van Dyke (1935b) to separate the species. [Liophloeus Germar 1817, 1 sp., L. tessulatus (Mueller 1776), inter-
cepted in quarantine; New York. Not established in North
Peritelinus Casey 1888, 3 spp., far western United States and adja- America.]
cent British Columbia. See Van Dyke (1936a) to separate the spe-
cies. Pachyrhinus Schoenherr 1823, 8 spp., generally distributed in the
western United States and Canada east to Nova Scotia and south
Peritelodes Casey 1888, 1 sp., P. obtectus Casey 1888, California. to Pennsylvania and Connecticut. Adults are associated with Pinus
(pine; Pinaceae). The genus needs revision. See Fall (1901) to
Peritelopsis Horn 1876, 1 sp., P. globiventris (LeConte 1857), Cali- separate the species.
fornia. Scythropus Schoenherr 1826
Carpomanes Gistel 1856
Rhypodillus Cockerell 1906, 2 spp., R. brevicollis (Horn 1876), Ari-
zona, New Mexico, Texas and Colorado, and R. dilatatus (Horn Polydrusus Germar 1817, 7 spp., generally distributed in the north-
1876), California. See LeConte and Horn (1876) to separate the eastern United States and adjacent southern Canada, also in Ari-
species. zona and New Mexico; includes 3 adventive species, P. cervinus
Rhypodes Horn 1876 (Linnaeus 1758), P. impressifrons (Gyllenhal 1834), and P. sericeus
(Schaller 1783). Adults generally feed on foliage of various trees.
Stenoptochus Casey 1888, 2 spp., S. inconstans Casey 1888 and S. The genus needs revision and the relationships between the na-
vanduzeei Van Dyke 1935, California. See Van Dyke (1935b) to tive and adventive species reassessed. See Sleeper (1957c) to sepa-
separate the species. rate most of the species. Alonso-Zarazaga and Lyal (1999:175)
list 24 valid subgeneric names, not including synonyms. For brevity,
Stomodes Schoenherr 1826, 1 sp., S. gyrosicollis Boheman 1843, these are not repeated here.
Maine; adventive.
58. Sciaphilini Sharp 1891
Thinoxenus Horn 1876, 1 sp., T. squalens Horn 1876, California.
Barypeithes Jacquelin du Val 1854, 1 sp., B. pellucidus (Boheman
Thricolepis Horn 1876, 2 spp., T. inornata Horn 1876, generally 1834), generally distributed in the northeastern United States and
distributed in the western United States, and T. simulator Horn adjacent southern Canada, also in California, Oregon, Washing-
1876, California. See LeConte and Horn (1876) to separate the ton and British Columbia; adventive. Adults feed on foliage of a
species. wide variety of plants.
Barypithes Gemminger and Harold 1871
56. Phyllobiini Schoenherr 1826 Exomias Bedel 1883 (valid subgenus)
Moroderia Reitter 1915
Aphrastus Say 1831, 3 spp., generally distributed in the eastern
United States west to Texas, and southern Canada. Adults feed
780 · Family 131. Curculionidae

Brachysomus Schoenherr 1823, 1 sp., B. echinatus (Bonsdorff 1785), Isodacrys Sharp 1911, 2 spp., I. ovipennis (Schaeffer 1908) and I.
Massachusetts, Minnesota, Quebec and Newfoundland; adven- burkei Howden 1961, Texas. Adults have been collected sweeping
tive. herbaceous Asteraceae. See Howden (1961) to separate the spe-
Pavrosomus Fischer de Waldheim 1829 cies.
Platytarsus Schoenherr 1840
Thricolepoides O’Brien 1979 Isodrusus Sharp 1911, 1 sp., I. debilis Sharp 1911, Texas.

Mitostylus Horn 1876, 3 spp., southwestern United States includ- Miloderoides Van Dyke 1936, 3 spp., Nevada, Colorado, Utah,
ing Texas and Oklahoma. Adults are found on various kinds of Wyoming and Idaho. See Tanner (1942) to separate the species.
low vegetation. The genus needs revision. See Van Dyke (1936b)
and Burke (1963) to separate the species. Minyomerus Horn 1876, 6 spp., southwestern United States, north
Derosomus Sharp 1891 into Kansas, Wyoming, Montana and Alberta. Adults have been
found on various kinds of low vegetation. The genus needs
Sciaphilus Schoenherr 1823, 1 sp., S. asperatus (Bonsdorff 1785), revision and the relationships of this genus to Piscatopus need
generally distributed in the northeastern United States and adja- reassessment. No key to species exists.
cent southern Canada, also Idaho, South Dakota and British Elissa Casey 1888
Columbia; adventive. Adults generally feed on foliage. Pseudelissa Casey 1888
Lygophilus Fischer von Waldheim 1829; not Rafinesque 1815
Sphaerilethmus Gistel 1856 Pachnaeus Schoenherr 1826, 2 spp., P. litus (Germar 1824), south-
ern Florida, and P. opalus (Olivier 1807), coastal southeastern United
59. Sitonini Gistel 1856 States from northern Florida north to New Jersey. Adults are
general foliage feeders and are citrus pests. The larvae feed on
Sitona Germar 1817, 11 spp., generally distributed; includes 5 roots. See Woodruff (1981) to separate the species.
adventive species. Adults are associated with various herbaceous Docorhinus Schoenherr 1823
species of Fabaceae. Larvae feed in the soil on roots. Some species Pachneus Gemminger and Harold 1871
such as the pea leaf weevil, S. lineatus (Linnaeus 1758), the
sweetclover weevil, S. cylindricollis (Fahraeus 1840), and the clover Pandeleteinus Champion 1911, 3 spp., generally distributed in the
root curculio, S. hispidulus (Fabricius 1776), are agricultural pests. southwestern United States. Adults have been collected on vari-
See Bright (1994) to separate the species. ous species of trees. See Howden (1959) to separate the species.
Charagmus Schoenherr 1826 (valid subgenus)
Clyptus Villa and Villa 1833 Pandeleteius Schoenherr 1834, 13 spp., generally distributed in the
Sitones Schoenherr 1840 southwestern and eastern United States and adjacent southern
Parasitones Sharp 1896 Canada. Adults are frequently found on Quercus (oak; Fagaceae)
Sitonidius Mueller 1913 or on various trees in the family Fabaceae. See Howden (1959) to
Coelositona González 1971 (valid subgenus) separate the species.
Pandeletius Agassiz 1846
60. Tanymecini Lacordaire 1863 Menetypus Kirsch 1868
Pandeletius Gemminger and Harold 1871; not Agassiz 1846
Tanymecina Lacordaire 1863 Pandeletejus Horn 1876
Exmenetypus Voss 1954 (valid subgenus)
Amotus Casey 1888, 3 spp., California. Adults are associated with
Artemisia (Asteraceae) and perhaps other shrubs and trees. See Piscatopus Sleeper 1960, 1 sp., P. griseus Sleeper 1960. Adults have
Van Dyke (1935b) to separate the species. This genus is question- been found on Larrea tridentata (Zygophyllaceae). The relation-
ably distinct from Stamoderes (Geonemini). We can find no charac- ships of this genus to Minyomerus need reassessment.
ters to reliably distiunguish these two genera.
Mimetes Schoenherr 1847; not Eschscholtz 1818; not Leach Scalaventer Howden 1970, 1 sp., S. subtropicus (Fall 1907), south-
1820; not Huebner 1821; not Vigors 1827; not Gloger ern Florida. Adults have been collected on Bumelia celastrina H.B.K.
1841 (Sapotaceae) and various other trees (Anderson 1993a).

Hadromeropsis Pierce 1913, 1 sp., H. opalinus (Horn 1876), Arizona. Tanymecus Germar 1817, 3 spp., generally distributed in the east-
Adults have been collected on Calliandra eriophylla Benth. and ern, central and southwestern United States north into the prairie
Acacia (Fabaceae) (Howden 1982). provinces of Canada. Adults are usually collected on low herba-
Hadromerus Schoenherr 1834; not Schoenherr 1823 ceous plants. Adults of T. lacaena (Herbst 1797) have been col-
Hadrorestes Howden 1982 (valid subgenus) lected commonly on Sesuvium portulacastrum (L.) L. (Aizoaceae) in
 Family 131. Curculionidae · 781

southern Florida (Anderson 1993a). See Van Dyke (1935b) and Trachyphloeosoma Wollaston 1869, 1 sp., T. advena Zimmerman
Blatchley and Leng (1916) to separate the species. 1956, Alabama, Florida, Georgia, Mississippi, North Carolina
Hynnulus Villa and Villa 1833 and South Carolina; adventive.
Episomechus Reitter 1903 (valid subgenus) Trachyphloeops Roelofs 1873
Geomecus Reitter 1903 (valid subgenus)
Indomecus Pajni and Gandhi 1987 Trachyphloeus Germar 1817, 4 spp., T. aristatus (Gyllenhal 1827),
T. asperatus Boheman 1843, T. angustisetulus Hansen 1915, and T.
Trigonoscutoides O’Brien 1977, 1 sp., T. texanus O’Brien 1977, Texas. bifoveolatus (Beck 1817), northeastern United States and adjacent
Adults are collected on and under Quercus havardii Rydb. (O’Brien Canada, also northwestern United States and adjacent Canada;
1977a). adventive. See Brown (1965) and Borovec (1989) to separate the
species. This genus is questionably distinct from Cathormiocerus.
61. Thecesternini Lacordaire 1863 We can find no characters to reliably distinguish these two genera.
Phyllastolus Gistel 1856
Thecesternus Say 1831, 7 spp., eastern, central and southwestern Lacordairius Brisout 1866 (valid subgenus)
United States north into Alberta. Adults are flightless and found Chaetechus Horn 1876
under rocks and cow dung (Kissinger 1964). Larvae of T. hirsutus Paratrachyphloeus Desbrochers 1895
Pierce 1909 feed on the roots of Parthenium hysterophorus L. Pseudolacordairius Escalera 1923 (valid subgenus)
(Asteaceae) (McClay and Anderson 1985). The genus needs revi-
sion. See Pierce (1909) to separate the species. 63. Tropiphorini Marseul 1863
Lithodus Germar 1834
Thicosternus Gemminger and Harold 1871 Adaleres Casey 1895, 3 spp., California. See Casey (1895) to sepa-
rate the species.
62. Trachyphloeini Lacordaire 1863
Anametis Horn 1876, 2 spp., A. granulata (Say 1831), generally
Trachyphloeina Lacordaire 1863 distributed in the eastern United States and adjacent southern
Canada, and A. subfusca Fall 1907, Arizona, Texas, New Mexico
Cathormiocerus Schoenherr 1842, 1 sp., C. curvipes Wollaston 1854, and Colorado. The relationships of this genus to Dichoxenus
Oregon; adventive. This genus is questionably distinct from Horn 1876, Peritaxia Horn 1876 and some species of the Mexi-
Trachyphloeus. We can find no characters to reliably distinguish can genus Amphidees Schoenherr 1842 need to be reassessed. See
these two genera. Fall and Cockerell (1907) to separate the species.
Scoliocerus Wollaston 1854
Mitomermus Jacquelin du Val 1854 (valid subgenus) Byrsopages Schoenherr 1842, 1 sp., B. villosus Boheman 1842, Alaska.
Schaumius Brisout 1866 (valid subgenus) Strongylophthalmus Motschulsky 1860
Cathormiocerinus Escalera 1918 (valid subgenus) Strongylophthalmus Faust 1894; not Motschulsky 1860
Kurilonus Sharp 1896
Cercopedius Sleeper 1955, 1 sp., C. artemisiae (Pierce 1910), western
United States from Nevada, Utah and Colorado north, into Brit- Cimbocera Horn 1876, 4 spp., generally distributed in the western
ish Columbia. Adults are found on Artemisia (sagebrush; inland United States and Canada. Adults are found on various
Asteraceae). woody shrubs at night. See Van Dyke (1935a) and Tanner (1941)
to separate the species.
Cercopeus Schoenherr 1842, 11 spp., generally distributed in the
eastern United States west into Texas. Adults are flightless and Connatichela Anderson 1984, 1 sp., C. artemisiae Anderson 1984,
found in leaf litter. The genus needs revision. See Sleeper (1955b), Yukon Territory and Alaska. Adults are associated with Artemisia
Burke (1963) and O’Brien (1977b) to separate the species. frigida Willd. (Asteraceae) (Anderson 1984).
Cercopius Gemminger and Harold 1871
Crocidema Van Dyke 1934, 5 spp., Arizona, Texas, Utah and Cali-
Chaetechidius Sleeper 1955, 1 sp., C. speciosus Sleeper 1955, Colo- fornia. Adults are found on various woody shrubs at night. The
rado. Adults were found under a stone. genus needs revision and its relationships to Pseudorimus Van
Dyke 1934 and Melanolemma Van Dyke 1935 reassessed. See Van
Pseudocercopeus Sleeper 1955, 1 sp., P. setosus Sleeper 1955, Arizona. Dyke (1934, 1951) to separate the species.

Pseudocneorhinus Roelofs 1873, 1 sp., P. bifasciatus (Roelofs 1880), Diamimus Horn 1876, 1 sp., D. subsericeus Horn 1876, western
eastern United States; adventive. Adults and larvae feed on a wide inland United States north to Montana.
variety of plants (Maier 1983).
782 · Family 131. Curculionidae

Dichoxenus Horn 1876, 4 spp., eastern and central United States Paracimbocera Van Dyke 1938, 3 spp., Wyoming, Idaho, Colorado,
from Illinois and Arkansas west to Texas, Colorado and Wyo- Nevada and California. Paracimbocera robusta (Van Dyke 1935) has
ming. The relationships of this genus to Anametis Horn 1876, been recorded from Ephedra nevadensis S. Wats. (Ephedraceae)
Peritaxia Horn 1876 and some species of the Mexican genus (Sleeper and Jenkins 1967) and P. artemisiae Ting 1940 from Arte-
Amphidees Schoenherr 1842 need to be reassessed. See Sleeper misia (Asteraceae) (Ting 1940). See Ting (1940) to separate the
(1956b) to separate the species. species.

Dirotognathus Horn 1876, 2 spp., D. punctatus Hatch 1971, Or- Paranametis Burke 1960, 1 sp., P. distincta Burke 1960, Texas.
egon, and D. sordidus Horn Arizona, California, Nevada and Colo-
rado. See Hatch (1971) to separate the species. This genus is ques- Peritaxia Horn 1876, 7 spp., southwestern United States north to
tionably distinct from Lepidophorus (Alophini). Wyoming and Montana. Adults are noctural and collected on
various kinds of plants. The genus needs revision and the rela-
Dyslobus LeConte 1869, 34 spp., generally distributed in the west- tionships of this genus to Dichoxenus Horn 1876, Anametis Horn
ern United States and adjacent Canada. Adults are flightless and 1876 and some species of the Mexican genus Amphidees Schoenherr
noctural and are found on foliage of various plants. The genus 1842 need to be reassessed. There is no key to species.
needs revision. See Van Dyke (1933) and Hatch (1971) to separate
some of the species. Phyxelis Schoenherr 1842, 2 spp., generally distributed in the east-
Lepesoma Motschulsky 1845; not Spix 1825 ern United States and adjacent southern Canada. The genus needs
Lepidosoma Agassiz 1846; not Wagler 1830; not Swainson revision since at least two undescribed species are known. See
1839 Blatchley and Leng (1916) to separate the species.
Ledidosoma Gemminger and Harold 1871; not Wagler 1830; Geoderces Horn 1876
not Swainson 1839; not Agassiz 1846
Melamomphus Horn 1876 Pseudorimus Van Dyke 1934, 2 spp., Arizona and New Mexico.
Amnesia Horn 1876 The relationships of this genus to Melanolemma Van Dyke 1935
Thricomigus Horn 1876 and Crocidema Van Dyke 1934 need to be reassessed. See Van Dyke
(1934) to separate the species.
Leptopinara O’Brien 1981, 2 spp., L. papillata O’Brien 1981, New
Mexico and Texas, and L. flemingi Anderson 1993, Texas. See Rhigopsis LeConte 1874, 1 sp., R. effracta LeConte 1874, Califor-
Anderson (1993c) to separate the species. nia.

Melanolemma Van Dyke 1935, 1 sp., M. montana Van Dyke 1935, Tropiphorus Schoenherr 1842, 3 spp, Newfoundland, Nova Scotia
Colorado. The relationships of this genus to Pseudorimus Van and Quebec; adventive. See Brown (1967) to separate the species.
Dyke 1934 and Crocidema Van Dyke 1934 need to be reassessed. Brius Dejean 1821
Tropidophorus Gistel 1856; not Duméril and Bibron 1839
Miloderes Casey 1888, 6 spp., Utah, Nevada, Arizona and Califor- Tropidophorus Gemminger and Harold 1871; not Duméril
nia. See Kissinger (1960) to separate some of the species. and Bibron 1839; not Gistel 1856; not Jan 1865
Synirmus Bedel 1883
Orimodema Horn 1876, O. protracta Horn 1876, generally distrib- Dochorhynchus Desbrochers 1897
uted in the southwestern United States.
Vitavitus Kissinger 1974, 1 sp., V. thulius Kissinger 1974, Alaska,
Panscopus Schoenherr, 1842, 28 spp., generally distributed through- Yukon Territory, Northwest Territories and Nunavut. Adults are
out the United States and southern Canada. Adults are mostly flightless and are collected in tundra and dry south-facing slopes
noctural and are associated with various types of plants. Most (Anderson 1997).
species are in forests but a few are found in more arid habitats.
The genus needs revision. See Buchanan (1936b) and Hatch (1971) XIII. Hyperinae Marseul 1863
to separate most of the species.
Nocheles LeConte 1874 (valid subgenus) by Robert S. Anderson
Phymatinus LeConte 1876 (valid subgenus)
Nomidus Casey 1895 (valid subgenus) Only the genus Hypera occurs in North America. It is recognized
Neopanscopus Pierce 1913 (valid subgenus) by a short snout (Fig. 81) (lacking deciduous processes and asso-
Panscopidius Pierce 1913 ciated scars), the pronotum lacks a postocular lobe, and at least
Pseudopanscopus Buchanan 1927 (valid subgenus) some of the scales of the body are bifid. In Mexico, Central and
Dolichonotus Buchanan 1936 (valid subgenus) South America several related genera are found. Larvae of all
Parapanscopus Buchanan 1936 (valid subgenus) hyperines feed externally on plant foliage and make loosely wo-
ven cocoons, which they attach to the host plants, in which they
 Family 131. Curculionidae · 783

of lixines can be short and wide or long and slender and most
species are grey or otherwise dull in color and scale pattern.
Larvae of most species mine in the roots and stems of
various plants but a few such as Rhinocyllus and Larinus have
larvae that feed in reproductive structures on seeds. Many species
are associated with Asteraceae and Fabaceae but members of the
genus Lixus are also found in semi-aquatic and aquatic habitats.
Several genera have been introduced into North America for bio-
logical control of noxious or pest weeds especially in rangelands
of western North America.

KEY TO THE NEARCTIC GENERA OF LIXINAE

FIGURE 81.131. Hyperinae. 81. Hypera punctata (Fabricius), head, 1. Rostrum short and broad in dorsal view, from apex
of epistoma to anterior margin of eye, more or
lateral view.
less as long as greatest width (Fig. 82); body size
pupate. Plants in the Polygonaceae and Fabaceae appear to be the small, at most slightly greater than 5 mm ......... 2
— Rostrum more elongate, in dorsal view, from apex
primary hosts. of epistoma to anterior margin of eye, much longer
than greatest width; body size moderate to large,
CLASSIFICATION OF THE NEARCTIC HYPERINAE subequal to or greater than 5 mm .................... 4

2(1). Elytron with intervals each with a row of conspicu-

64. Hyperini Marseul 1863 ous, erect setae in addition to appressed hair-
like scales; anterolateral margin of pronotum
Hypera Germar 1817, 17 species, generally distributed through- straight, with long postocular vibrissae immedi-
out the United States and Canada, north into Alaska; 6 species are ately behind eye; eye more or less round .........
........................................................ Microlarinus
adventive. Adults and larvae feed externally on foliage of various — Elytron with intervals lacking erect setae, with only
Fabaceae and Polygonaceae (Titus 1911, Puttler et al. 1973). A appressed to suberect hair-like scales; anterolat-
number of pest species are included in the genus, namely, H. eral margin of pronotum with rounded postocular
postica (Gyllenhal 1813), the alfalfa weevil; H. brunneipennis lobe and short postocular vibrissae; eye distinctly
elongate-oval ................................................... 3
(Boheman 1834), the Egyptian alfalfa weevil; H. nigrirostris (Fab-
ricius 1775), the lesser clover weevil; H. punctata (Fabricius 1775), 3(2). Prosternum with a pair of prominent lateral ridges
the clover leaf weevil; H. meles (Fabricius 1792), the clover head which form a deep ventral channel; dorsal
weevil; and H. rumicis (Linnaeus 1758). The genus needs revision. vestiture of pronotum and elytra with scales in
part bifid ....................................... Bangasternus
See Titus (1911) to separate most of the species. — Prosternum lacking ridges, no ventral channel evi-
Phytonomus Schoenherr 1823 dent; dorsal vestiture of pronotum and elytra with
Dapalinus Capiomont 1868 (valid subgenus) scales simple .................................... Rhinocyllus
Eririnomorphus Capiomont 1868 (valid subgenus)
4(1). Ventral and dorsal surfaces of body with numerous
Tigrinellus Capiomont 1868 (valid subgenus) long, fine, erect hairs, some hairs about as long
Phytonomidius Capiomont 1868 as antennal scape; white vestiture of both dorsal
Antidonus Bedel 1886 (valid subgenus) and ventral surfaces, especially prosternum and
Spongifer Petri 1901 abdomen, composed of numerous pectinate
suberect or appressed scales ........ Eustenopus
Heteromorphus Petri 1901 — Ventral and dorsal surfaces of body with at most
Boreohypera Korotyaev 1999 (valid subgenus) suberect short hair-like scales, or erect hairs if
present, sparse and many times shorter than an-
XIV. Lixinae Schoenherr 1823 tennal scape; white vestiture of both dorsal and
ventral surfaces composed of simple appressed
scales, some scales of mesosternum and coxae
by Robert S. Anderson bifid or pectinate ............................................. 5

This is a relatively small subfamily of generally large-sized weevils 5(4). Pronotum dorsally and laterally with numerous
shiny, glabrous tubercles, lacking distinct punc-
associated mostly with arid habitats. Lixines are readily recog- tures (Figs. 83-84) ............................................. 6
nized by the large tooth at the apex of the hind tibia, their larger — Pronotum dorsally and laterally with distinct punc-
size, and short, globular and telescoping labial palpi of 3 articles tures, with at most the outer margins of punc-
(but appearing composed of 1 article) (Fig. 90), ventrally situated tures at lateral margins swollen, glabrous and
shiny ................................................................. 7
at the apex of the large prementum. Females possess large paired
symbiont sacs attached to the vagina near the base of the
gonocoxites but this can only be seen in dissections. The rostrum
784 · Family 131. Curculionidae

8(7). Tibia with apical flange rounded (Fig. 89); elytron

83 with all intervals equally flat or with at most only
humerus and very base of interval 3 swollen and
convex; pronotum with disk with scale pattern
various; prosternum with or without swellings,
swellings, if present, situated immediately ante-
rior to each front coxal cavity and lateral to each
prosternal impression ...................................... 9
— Tibia with apical flange sharp, carinate; elytron with
82 all intervals equally flat or with humerus and vari-
ously sutural interval, intervals 3, 5, 7 and 9 el-
evated and convex throughout the greater part
of their length; pronotum with disk with large
white scales in a lateral stripe of various width,
small and fine in moderately broad to very broad
apically narrowed median stripe, median area
largely black in color, underlying dark cuticle not
obscured by overlying scales; prosternum with
85 or without swellings, swellings, if present, situ-
84 ated immediately anterior to each prosternal im-
pression ......................................................... 11

9(8). Antenna with article 2 of funicle distinctly longer

than wide; distinctly longer than each of articles
3 to 6, slightly shorter than to distinctly longer
than article 1; pronotum with anterolateral margin
straight, slightly sinuate, or with at most variously
developed (usually small) acute, postocular pro-
87 jection, postocular vibrissae unequal in length,
greatest length (more or less equal to one-half
width of eye) behind base of each eye (Fig. 87);
86 femur with ventral surface dentate or not ........
................................................................... Lixus
— Antenna with article 2 of funicle more or less as
long as wide; more or less subequal in length to
each of articles 3 to 6, shorter than article 1;
pronotum with anterolateral margin straight or
with slightly to well-developed rounded postocu-
lar lobe, postocular vibrissae uniformly short (less
89 than one-half width of eye in length) to unequal
88 in length, greatest length (more or less equal to
90 or greater than one-half width of eye) behind base
of each eye; femur with ventral surface not den-
FIGURES 82.131-90.131. Lixinae. 82. Bangasternus orientalis
tate ................................................................. 10
(Capiomont), head, dorsal view. 83-87. Head and pronotum, lateral
view, 83. Cleonis pigra (Scopoli); 84. Cyphocleonus achates (Fahraeus); 10(9). Elytra elongate-narrow (width at midlength less than
85. Apleurus albovestitus (Casey); 86. Stephanocleonus plumbeus LeConte; 0.65 times length); pronotal disk with distinct
87. Lixus scrobicollis Boheman. 88-89. Lixus scrobicollis Boheman, 88. white scales of various sizes; elytra with white
Fore tarsus; 89. Fore tibia. 90. Labial palpi, Lixinae, schematic (after scales, various in size, but more or less obscur-
Anderson 1988a) ing view of underlying cuticle over large part of
elytral surface .......................... Scaphomorphus
6(5). Rostrum with single median sulcus; pronotum with — Elytra more robust (width at midlength greater than
anterolateral margin with rounded postocular lobe 0.65 times length); pronotal disk with at most only
(Fig. 83) ................................................... Cleonis very short fine setae, distinct scales absent; elytra
— Rostrum carinate medially throughout length, lat- with scattered patches of elongate fine hair-like
eral margins raised towards base, appearing as scales in addition to very short setae, underlying
lateral carinae; pronotum with anterolateral mar- cuticle not obscured by scales ............ Larinus
gin straight behind eye, lacking postocular lobe
(Fig. 84) ........................................ Cyphocleonus 11(8). Epistoma with anterior margin emarginate; pronotum
with well-developed, rounded postocular lobes,
7(5). Mesosternum with mesosternal process markedly postocular vibrissae indistinct, uniformly short
tumescent; male with aedeagus expanded later- (Fig. 86); eyes widest near upper margin, flat or
ally from midlength to apical one-third ............. only slightly convex in dorsal view ..................
.................................................... Apleurus (part) ................................................ Stephanocleonus
— Mesosternum with mesosternal process flat or at — Epistoma with anterior margin rounded; pronotum
most slightly convex, not tumescent; male with with anterior margin straight behind eyes or with
aedeagus more or less uniform in width through- small acute projection immediately behind base
out median portion of length ........................... 8 of eye, postocular vibrissae distinct and long,
longest immediately behind base of eye (Fig. 85);
 Family 131. Curculionidae · 785

eyes widest near midheight, protruding and mod- Cleonis Dejean 1821, 1 sp., C. pigra (Scopoli 1763), far northeastern
erately to markedly convex in dorsal view ......
United States and adjacent southern Canada, adventive. Adults
.................................................... Apleurus (part)
are associated with Cirsium arvense (L.) Scop. (Canada thistle) and
CLASSIFICATION OF THE NEARCTIC LIXINAE C. vulgare (Savi) Tenore (bull thistle) (Anderson 1988a).
Geomorphus Schoenherr 1823
65. Lixini Schoenherr 1823 Cleonus Schoenherr
Xerobia Gistel 1856
Eustenopus Petri 1907, 1 sp., E. villosus (Boheman 1843), locally
distributed in the western United States. Introduced for the bio- Cyphocleonus Motschulsky 1860, 1 sp., C. achates (Fahraeus 1842),
logical control of Centaurea solstitialis L. (yellow star-thistle) Colorado, Wyoming, Montana, Oregon and British Columbia.
(Asteraceae). This species was introduced as a biological control agent for Cen-
taurea maculosa Lam. (spotted knapweed) and C. diffusa Lam. (dif-
Larinus Dejean 1821, 3 spp., locally distributed in the northeast- fuse knapweed) (Asteraceae) (Lang 1997c). It has been introduced
ern and northwestern United States, British Columbia, Manitoba in other states but does not appear to be established.
and Nova Scotia, adventive. In North America, the species L.
planus (Fabricius 1792), L. obtusus Gyllenhal 1836, and L. minutus Scaphomorphus Motschulsky 1860, 19 spp., generally distributed
Gyllenhal 1836 have been introduced for the biological control of in the western and central United States and adjacent southern
Cirsium arvense (L.) Scop. (Canada thistle), and Centaurea solstitialis Canada, also along the eastern coastal United States from Florida
L. (yellow star-thistle), C. maculosa Lam. (spotted knapweed) and north into New York. Adults mostly are associated with various
C. diffusa Lam. (diffuse knapweed) (all Asteraceae) (Lang 1997a, arid habitat Fabaceae and Asteraceae (Anderson 1988a). See Ander-
b). There is no key to the North American species. son (1988a; as Cleonidius) to separate the species.
Rhinobatus Germar 1817; not Walbaum 1792; not Schneider Scaphidomorphus Lacordaire 1863; not Hope 1841
1801 Cleonidius Casey 1891
Larinus Germar 1824; not Dejean 1821 Lixestus Reitter 1916
Phyllonomeus Gistel 1856 (valid subgenus)
Larinodontes Faust 1898 Stephanocleonus Motschulsky 1860, 6 spp., western montane United
Cryphopus Petri 1907 (valid subgenus) States north into the Yukon Territory and east across Canada to
Lariniorhynchus Reitter 1924 Newfoundland. See Anderson (1988a, 1989b) to separate the
Larinomesius Reitter 1924 (valid subgenus) species.
Eucleonus Faust 1904; not Gistel 1856
Lixus Fabricius 1801, 69 species, generally distributed throughout Deracanthopsis Voss 1967 (valid subgenus)
the United States and Canada. Adults are associated with various Eremocleonus Ter-Minasian 1974 (valid subgenus)
plants in the Asteraceae and Polygonaceae. The genus needs revision. Taeniocleonus Ter-Minasian 1974 (valid subgenus)
See Blatchley and Leng (1916) to separate some of the species. Alonso- Sanzia Alonso-Zarazaga and Lyal 1999 (valid subgenus)
Zarazaga and Lyal (1999:190) list 18 valid subgeneric names, not
including synonyms. For brevity, these are not repeated here. 67. Rhinocyllini Lacordaire 1863

Microlarinus Hockhuth 1847, 2 spp., M. lareynii (Jacquelin du Val Bangasternus Gozis 1882, 2 spp., B. orientalis (Capiomont 1873)
1852) and M. lypriformis (Wollaston 1861), southwestern United and B. fausti Reitter 1890, California, Montana, Nebraska and
States and Washington. Adults have been introduced for the Oregon. These species have been introduced for the control of
biological control of Tribulus terrestris L. (puncturevine; Centaurea solstitialis L. (yellow star-thistle), C. diffusa Lam. (dif-
Zygophyllaceae) (Kirkland and Goeden 1977, 1978a, b). See Hatch fuse knapweed) and C. maculosa Lam. (spotted knapweed)
(1971) to separate the species. (Asteraceae) (Lang 1997d). There is no key to separate the two
species in North America.
66. Cleonini Schoenherr 1826 Coelostethus Capiomont 1873; not LeConte 1861

Apleurus Chevrolat 1873, 6 spp., southwestern United States east Rhinocyllus Germar 1817, 1 sp., R. conicus (Froelich 1792), locally
to Texas, north to Idaho. Adults are associated with various arid distributed throughout most of the United States and adjacent
habitat plants (Anderson 1988a). See Anderson (1988a) to sepa- southern Canada. This species was introduced for the biological
rate the species. control of Carduus nutans L. (nodding or musk thistle; Asteraceae);
Cleonopsis LeConte 1876 larvae feed in flowerheads (Kok 1998). Louda et al. (1997) report
Cleonaspis LeConte 1876 the species has apparently expanded its host range and is now a
Centrocleonus LeConte 1876; not Chevrolat 1873 threat to native species of Cirsium at various locations in the
Dinocleus Casey 1891 United States.
Gibbostethus Anderson 1988 (valid subgenus)
786 · Family 131. Curculionidae

CLASSIFICATION OF THE NEARCTIC MESOPTILIINAE

91 68. Laemosaccini Lacordaire 1866

Laemosaccus Schoenherr 1823, 2 spp., L. nephele (Herbst 1797),

generally distributed in the eastern United States and adjacent
southern Canada west into Texas, New Mexico and Arizona;
and, L. texanus Champion 1903, southern Texas and Arizona.
92 Adults of L. nephele are associated with species of Quercus
(Fagaceae) and Prosopis (Fabaceae), and L. texanus with various
Malvaceae (Blatchley and Leng 1916); larvae mine twigs or stems.
The genus needs revision; L. nephele is a composite of a number
of distinct host-specific species. See Champion (1903) to separate
the species.
93
69. Magdalidini Pascoe 1870
FIGURES 91.131-93.131. Mesoptiliinae. 91. Magdalis lecontei Horn,
dorsal habitus; 92. Laemosaccus nephele complex, lateral habitus; 93.
Magdalis lecontei Horn, hind tibia. Magdalis Germar 1817, 25 spp., generally distributed, M. barbicornis
(Latreille 1804) is adventive. Adults are associated with various
XV. Mesoptiliinae Lacordaire 1863 trees; larvae mine in bark of dead or dying trees (Blatchley and
Leng 1916). The genus needs revision. See Fall (1913), Blatchley
by Robert S. Anderson and Leng (1916) and Hatch (1971) to separate some of the spe-
cies.
This is a very small subfamily of only three genera in North Rhina Latreille 1802; not Schneider 1801
America. They are recognized by the presence of a large hook-like Edo Germar 1819 (valid subgenus)
apical tooth on the hind tibia (Fig. 93), the pronotum is only Porrothus Dejean 1821 (valid subgenus)
slightly narrower than the base of the elytra (Fig. 91), and the Rhinodes Dejean 1821
elytra have the basal margin from intervals 2-4 extended anteri- Panus Schoenherr 1823 (valid subgenus)
orly and overlapping the base of the pronotum (Fig. 91). The Thamnophilus Schoenherr 1823; not Vieillot 1816
genus Laemosaccus are compact cylindrical beetles with a short, Magdalinus Germar 1843
straight rostrum (Fig. 92), whereas Magdalis and Trichomagdalis Porrhothus Agassiz 1846
have a very different, anteriorly tapered form, with the width Scardamyctes Gistel 1848
across the apices of the elytra generally the widest part of the Panopsis Daniel 1903 (valid subgenus)
beetle. Neopanus Reitter 1916
Larvae of all species mine in wood (both hardwoods and Laemosaccidius Smreczynski 1972 (valid subgenus)
conifers) or in stems of herbaceous plants. The genus Magdalis Odontomagdalis Barrios 1984 (valid subgenus)
does not extend far into Mexico, but the genus Laemosaccus has
many Neotropical species, most undescribed. The center of di- Trichomagdalis Fall 1913, 3 spp., California. See Fall (1913) to sepa-
versity for Mesoptilinae appears to be in Chile. rate the species.

KEY TO THE NEARCTIC GENERA OF MESOPTILIINAE XVI. Molytinae Schoenherr 1823

1. Front coxae separated by prosternum; rostrum stout, by Robert S. Anderson

short and straight (Fig. 92); elytra black with red
markings or entirely black ............ Laemosaccus
— Front coxae contiguous; rostrum elongate, cylin- Like Curculioninae, this subfamily is also somewhat of a conglom-
drical, curved ventrally or straight; elytra erate of likely unrelated forms. They are grouped together here pri-
unicolorous, black or reddish .......................... 2 marily because they all share a large, hook-like apical tooth on the
2(1). Elytra more or less parallel sided in dorsal view,
hind tibia, or have various modifications to the apex of the hind
with numerous fine, elongate scales; tarsus with tibia related to the development of the tooth (Figs. 99-101).
claws simple; pronotum with anterolateral angles Most taxa of molytines are associated with woody plants
smooth, not serrate .................. Trichomagdalis and have larvae that feed in dead wood or other dead and decay-
— Elytra usually widened posteriorly in dorsal view
(Fig. 91), nearly glabrous; tarsus with claws simple
ing plant material. Some taxa such as Lepilius, Epacalles and
or toothed basally; pronotum with anterolateral Lymantes are associated with leaf litter and likely have larvae that
angles serrate or smooth .................... Magdalis develop in fallen plant debris. Larvae of others such as Cholus
mine stems, or some such as Conotrachelus, feed in the seeds,
 Family 131. Curculionidae · 787

94 96

95

97

99

98 101
100
FIGURES 94.131-101.131. Molytinae. 94-95. Lateral habitus, 94. Lepyrus sp.; 95. Microhyus setiger LeConte. 96. Dioptrophorus repens (Casey),
head and pronotum, lateral view. 97. Hilipinus nearcticus O’Brien, head, lateral view. 98. Heilus bioculatus (Boheman), hind coxae and first
abdominal ventrite. 99-101. Hind tibia, 99. Conotrachelus posticatus Boheman; 100. Pachylobius picivorus (Germar); 101. Pissodes strobi (Peck).

fruits or reproductive structures of living plants. Several taxa rep- — Rostrum in repose not received into ventral chan-
nel, but may rest between front, middle and/or
resented in the United States by only one species, such as Hilipinus,
hind coxae ..................................................... 13
Heilus and Heilipus, are significantly more diverse in the Neotro-
pical Region with hundreds of species found there. Odontopus 5(4). Tarsus with claw simple, free or connate, lacking
and Piazorhinus have larvae that mine leaves. basal tooth ........................................................ 6
— Tarsus with claw with basal tooth ..................... 10
With over 500 species, the genus Conotrachelus may prove to
be the most diverse genus of weevil in the Americas. The odd 6(5). Tarsus with claws connate at base; pronotum
genera Thalasselephas and Hormops are associated with seaweed and coarsely strigose or punctate; some with metal-
tree squirrel nests, respectively. lic sheen ..................................... Chalcodermus
— Tarsus with claws free; pronotum finely and shal-
lowly punctate; not with metallic sheen .......... 7
KEY TO THE NEARCTIC GENERA OF MOLYTINAE
7(6). Pronotum and elytra with sparse, long, erect stout
1. Eyes absent or reduced in size to less than 12 fac- seta-like scales (Fig. 95); body length less than
ets .................................................................... 2 2.5 mm ............................................... Microhyus
— Eyes present, well-developed, composed of more — Pronotum and elytra lacking long, erect vestiture,
than 12 facets .................................................. 3 with only appressed scales; length various .... 8

2(1). Antenna with funicle of 8 articles; eyes absent; 8(7). Body form elongate; California ......... Micromastus
southern Florida ............................ Caecossonus — Body form oval; eastern United States west into
— Antenna with funicle of 7 articles; eyes absent (but Texas ................................................................ 9
indicated by setose swelling on basal portion of
rostrum) or present, composed of up to 11 fac- 9(8). Pronotum and elytra with scattered short, clavate,
ets; eastern United States (not southern Florida) recurved setae and appressed scales; eyes lat-
west into southwestern Texas .......... Lymantes eral in placement, separated dorsally by a dis-
tance slightly greater than width of rostrum at
3(2). Head with eyes obviously situated on basal portion base; metasternum steeply sloped between hind
of rostrum, head distinctly constricted and globu- coxae; extreme southwestern Texas ... Lepilius
lar behind eyes (Fig. 96) ............. Dioptrophorus — Pronotum and elytra with only appressed scales;
— Head with eyes obviously situated on head, not on eyes lateral in placement, but with upper portion
rostrum (Fig. 97); head may be constricted and encroached on dorsal surface of head, separated
globular behind base of rostrum ...................... 4 dorsally by a distance slightly less than width of
rostrum at base; metasternum gradually sloped
4(3). Rostrum in repose received into ventral channel between hind coxae; eastern United States into
on prosternum .................................................. 5 central Texas ...................................... Epacalles
788 · Family 131. Curculionidae

10(5). Pronotum with only slightly developed postocular — Eyes separated by more than width of antennal club;
lobe; dorsum with at most a few scattered, unpig- front femur lacking obvious tooth; tarsal claw
mented scales; pronotum coarsely strigose or simple ............................................................. 20
rugulose or regularly punctate ... Rhyssomatus
— Pronotum with distinct postocular lobe; dorsum with 19(18). Front femur with large, serrate tooth; rostrum about
more or less dense, appressed, pigmented scales; as long as pronotum, subcylindrical; elytra lack-
pronotum regularly finely or coarsely punctate, ing scales ......................................... Odontopus
median impunctate line or carina present or not — Front femur with small, simple tooth; rostrum shorter
....................................................................... 11 than length of pronotum, dorsoventrally com-
pressed, spatulate, especially towards apex;
11(10). Elytra with all intervals carinate or alternate inter- elytra with narrow scales ................ Piazorhinus
vals carinate or at least swollen throughout most
of their length, in some specimens carina or swell- 20(18). Front coxae widely separated by the width of a
ing only or most evident on declivity; femora, coxa; body form markedly dorsoventrally com-
especially middle and hind with distinct tooth on pressed, upper contour flat ..................... Nanus
ventral margin ............................ Conotrachelus — Front coxae very narrowly separated by much less
— Elytra with all intervals flat or slightly but evenly than one-third the width of a coxa; body form
swollen throughout most of their length; femora more subcylindrical, upper contour rounded ...
with or without tooth of ventral margin ......... 12 ....................................................................... 21

12(11). Dorsum of pronotum and elytra with long, erect se- 21(20). Hind tibia with apex with apical comb of stout setae
tae and appressed scales; rostrum long and slen- oriented longitudinally along axis of tibia on outer
der ..................................................... Pheleconus margin (Fig. 101) ................................... Pissodes
— Dorsum of pronotum and elytra with only appressed — Hind tibia with apex with apical comb transverse or
scales; rostrum short and stout ....... Micralcinus obliquely oriented to long axis of tibia across
outer apical margin ......................................... 22
13(4). Front coxae very narrowly to widely separated by
prosternal processes ..................................... 14 22(21). Body size moderate, greater than 3.0 mm; scutel-
— Front coxae contiguous, not separated by lum large and distinct; California, adventive .....
prosternal processes ..................................... 24 ................................................................ Tranes
— Body size small, less than 3.0 mm; scutellum minute
14(13). Metepisternal suture absent; pronotum deeply, or not visible; coastal beaches of Pacific states,
coarsely punctate, distance between punctures British Columbia, and Florida .......................... 23
less than the diameter of a puncture; elytra with
numerous, low, setiferous tubercles ................. 23(22). Pronotum sculptured with paramedian broad impres-
............................................................ Anchonus sion, lateral impressions and low lateral tubercles
— Metepisternal suture present, although may be de- at anterior one-third; elytra with alternate inter-
fined only in anterior one-half; pronotum various, vals elevated especially so on declivity; body
but not deeply, coarsely punctate; elytra smooth color dark brown or black; Florida .... Gononotus
or with two large tubercles at about midlength on — Pronotum smooth and virtually impunctate, lateral
interval 2 ........................................................ 15 margins evenly rounded; elytra with intervals flat;
body color pale brown; Pacific coastal states and
15(14). Metepisternal suture visible and subcarinate in only British Columbia ....................... Thalasselephas
anterior one-half; elytra with scattered tufts of
suberect broad scales ...................... Trachodes 24(13). Tarsus with claws connate at base ................... 25
— Metepisternal suture visible throughout length; — Tarsus with claws free at base .......................... 26
elytra with appressed scales or fine vestiture ..
....................................................................... 16 25(24). Elytra with acute lateral subhumeral tubercle; eyes
moderate in size, widely separated ventrally; front
16(15). Hind tibia with equally large tooth at both outer and tibia with tooth on inner margin ...... Sternechus
inner apical angles; mandibles with apices diver- — Elytra with lateral margins simple, lacking tubercle;
gent, not overlapping .................................... 17 eyes very large and elongate, subcontiguous
— Hind tibia with large tooth at outer apical angle only ventrally; front tibia with inner margin simple ...
(Fig. 101), or also with much smaller tooth at inner ............................................................ Hormops
apical angle; mandibles convergent, overlapping
....................................................................... 18 26(24). Metepisternal suture absent; body size small to mod-
erate, length less than 6.0 mm ....................... 27
17(16). Pronotum with anterolateral margin with distinct pos- — Metepisternal suture present; body size moderate
tocular lobe; adventive, in greenhouses .......... to large, length greater than 6.0 mm .............. 28
................................................................. Cholus
— Pronotum with anterolateral margin almost straight, 27(26). Metasternum and abdominal ventrites 1 and 2 with
lacking distinct postocular lobes; native, extreme large, deep excavations ............. Gastrotaphrus
southern Arizona ......................... Neoerethistes — Metasternum and abdominal ventrites 1 and 2
smooth, lacking large, deep excavations .........
18(16). Eyes narrowly separated by less than width of an- ............................................................. Sthereus
tennal club; front femur with obvious tooth; tar-
sal claw with basal process ........................... 19 28(26). Pronotum markedly constricted and tubulate at
base; rostrum very long and fine, about twice as
 Family 131. Curculionidae · 789

long as length of pronotum; body black and glossy, area immediately behind hind coxa (Fig. 98); elytra
lacking distinct broad scales ............. Sicoderus with single, rounded patch of black scales form-
— Pronotum not constricted or tubulate at base, broad; ing an ‘eyespot’ at posterior two-fifths .... Heilus
rostrum moderate in length and somewhat stout, — Abdomen with ventrite 1 with anterior margin evenly
shorter than length of pronotum; body various, developed immediately behind hind coxa, no ex-
not black and glossy, with at least some broad panded area evident; elytra with scale pattern
appressed scales ........................................... 29 various, lacking ‘eyespots’ ............................ 35

29(28). Hind tibia markedly expanded at apex, wider at apex 35(34). Rostrum above scrobe coarsely punctate, not at all
than apex of femur; area adjacent to tarsal articu- sulcate; pronotum and elytra with contrasting
lation on hind femur with large flat flange (Fig. pattern of black cuticle and bright white, glossy
100); front tibia with outer angle produced and scales arranged around periphery; eyes sepa-
spatulate ......................................... Pachylobius rated dorsally by less than one-half width of the
— Hind tibia not significantly expanded at apex, width rostrum at base ..................................... Heilipus
at apex narrower than width of femur at apex; — Rostrum above scrobe with shallow, longitudinal
area adjacent to tarsal articulation on hind femur sulcus oriented parallel to dorsal margin of scrobe
with at most a low cariniform extension; front tibia (Fig. 97); pronotum and elytra scale pattern vari-
with outer angle rounded, not produced or spatu- ous, not contrasting; eyes separated dorsally by
late .................................................................. 30 a distance greater than or subequal to width of
the rostrum at base ........................................ 36
30(29). Femur with inner margin lacking tooth, evenly
rounded at apical one-third; tibia with inner mar- 36(35). Elytra with punctures each with a fine hair-like scale;
gin simple, not cariniform or expanded, outer rostrum with a slight longitudinal impression im-
margin more or less straight ........................... 31 mediately above scrobe, impression not defined
— Femur with inner margin with variously developed dorsally but more or less continuous with dor-
tooth at apical one-third; tibia with inner margin sum of rostrum; hind tibia with apical comb com-
cariniform and slightly to markedly expanded at posed of a single row of setae ... Hylobius (part)
point of occlusion with femoral tooth, outer mar- — Elytra with punctures each with a broad scale; ros-
gin arcuate ..................................................... 33 trum with a distinct, moderately deep longitudi-
nal impression immediately above scrobe, impres-
31(30). Pronotum with anterolateral margins lacking distinct sion defined dorsally by a low but distinct carina,
postocular lobes; eyes rounded, distinctly con- not continuous with dorsum of rostrum; hind tibia
vex; metepimeron visible, with vestiture and with apical comb composed of a long apical row
sculpture similar to metepisternum (Fig. 94) ...... of setae and a second short preapical row to-
.............................................................. Lepyrus wards the dorsal end of the apical row .............
— Pronotum with anterolateral margins with distinct ............................................................. Hilipinus
postocular lobes; eyes elongate-oval, flat or
slightly convex; metepimeron not visible, con-
CLASSIFICATION OF THE NEARCTIC MOLYTINAE
cealed by elytra, if metepimeron visible because
of displaced elytra, vestiture and sculpture finer
than on metepisternum .................................. 32 70. Molytini Schoenherr 1823
32(31). Antenna with funicle with article 2 longer than 1;
Plinthina Lacordaire 1863
elytra with pattern of brown scales with patches
of paler scales surrounded by black scales along
length of interval 4 and at apical callus, scales Gastrotaphrus Buchanan 1936, 1 sp., G. barberi Buchanan 1936, far
about twice as long as wide, striae with small, western United States and British Columbia. Adults have been
rounded punctures, each with a broad flat scale;
collected in moss and leaf litter (Anderson 1988b).
associated with Taxodiaceae ....... Eudociminus
— Antenna with funicle with article 2 shorter than 1;
elytra with pattern of scattered, white or cream Steremnius Schoenherr 1835, 3 spp., S. carinatus (Boheman 1842)
colored scales, scales many times longer than and S. tuberosus Gyllenhal 1836, far western United States, British
wide, striae with large, deep, elongate punctures,
Columbia and Alaska, and S. shermani (Fiske 1906), North Caro-
each with a fine hair-like seta; associated with
Pinaceae ..................................... Hylobius (part) lina, Tennessee and Virginia (at high elevations). Adults have
been collected in leaf litter; larvae feed in phloem of slash or
33(30). Metasternum between middle and hind coxae roots of dead conifers (Anderson 1988b). See Brown (1966b) to
shorter than length of a middle coxa; pronotum
separate the species.
and elytra with numerous, small, glossy, round
nodules; surface sculpture coarse and irregular Paraplinthus Faust 1892
......................................................... Steremnius
— Metasternum between middle and hind coxae Sthereus Motschulsky 1845, 4 spp., far western United States, Brit-
longer than length of a middle coxa; pronotum
ish Columbia, Alaska, Nova Scotia and Newfoundland. Adults
and elytra punctate or rugose (pronotum); sur-
face sculpture regular and more or less smooth of S. multituberculatus Buchanan 1936, S. quadrituberculatus
....................................................................... 34 Motschulsky 1845, and S. horridus (Mannerheim 1952) have been
associated with various conifers and collected in leaf litter; adults
34(33). Abdomen with ventrite 1 with raised anterior mar-
of S. ptinoides have been collected under driftwood on beaches
gin with posteriorly expanded, slightly crenulate
790 · Family 131. Curculionidae

(Anderson 1988b). See Hatch (1971) or Zimmerman (1964) to Ardoleucus Checrolat 1881
separate the species. Atroniscus Desbrochers 1906
Stereus Mannerheim 1846
Lobosoma Buchanan 1936 Rhinastina Vaurie 1973
Philostratus Zimmerman 1964
Lobosoma Zimmerman 1964 Neoerethistes O’Brien and Wibmer 1982, 1 sp., N. arizonicus (Sleeper
1954), Arizona.
71. Trachodini Gistel 1848
75. Cleogonini Gistel 1856
Trachodes Germar 1824, 1 sp., T. hispidus (Linnaeus 1758), New-
foundland; adventive. Rhyssomatus Schoenherr 1837, 17 spp., generally distributed with
Blastophila Gistel 1856 the exception of the northwestern United States and adjacent
Metrachodes Marshall 1948 Canada. Adults are associated with various plants in the families
Atrachodes Morimoto 1962 (valid subgenus) Asclepiadaceae, Asteraceae, Convolvulaceae and Fabaceae (Blatchley
and Leng 1916; Kissinger 1964; Anderson 1993a). The genus
72. Anchonini Imhoff 1856 needs revision. See Casey (1895) and Blatchley and Leng (1916) to
separate some of the species.
Anchonus Schoenherr 1825, 4 spp., Florida. Adults frequently are Polydus Pascoe 1872
collected in association with driftwood and in the litter of coastal Sermysatus Casey 1895 (valid subgenus)
hardwood hammocks (Thomas and O’Brien 1999). See Thomas
and O’Brien (1999) to separate the species. 76. Conotrachelini Jekel 1865
Choristorhinus Fairmaire 1899
[Chaleponotus Casey 1892, 1 sp., C. elusus Casey 1892, Indiana.
73. Camarotini Schoenherr 1833 This genus and species are known only from the type specimen,
labelled from “Indiana”. There is considerable doubt that this is
Alonso-Zarazaga and Lyal (1999) place these weevils as a North American taxon. At the time of its description Casey was
Curculioninae but the structure of the uncus at the tibial apex studying Brazilian Baridinae and there is the possibility that the
suggests they are Molytinae or related. Here they are placed as a locality reference is to Indiana, Brazil and not the state of Indiana
tribe within Molytinae. in the United States.]

Prionomerina Lacordaire 1863 Conotrachelus Dejean 1835, 63 spp., generally distributed through-
out the United States and Canada. Adults are associated with
Odontopus Say 1831, 1 sp., O. calceatus (Say 1831), generally distrib- various plants; many come to lights. Larvae feed in developing
uted in eastern United States. This species is associated with Sas- fruits and in injured or dying wood (Kissinger 1964). A number
safras (Lauraceae) and Liriodendron (Magnoliaceae); larvae mine of species are associated with Quercus (Fagaceae) and other hard-
leaves. woods. Conotrachelus nenuphar (Herbst 1797) is the plum curculio
Prionomerus Schoenherr 1835 and C. crataegi Walsh 1863 is the quince curculio. The genus needs
revision; a number of undescribed species are known from Florida
74. Cholini Schoenherr 1825 (Anderson 1993a). See Schoof (1942) and Blatchley and Leng
(1916) to separate some of the species. This treatment of
Cholina Schoenherr 1825 Conotrachelus includes C. parvulus Champion 1904, and Pheleconus
cribricollis (Say 1831) and P. infector (Boheman 1845); see also
Cholus Germar 1824, 1 sp., C. cattleyae (Champion 1916), Wash- notes about Micralcinus and Pheleconus. Relationships of these
ington, DC, New Jersey and Wisconsin; adventive. This species genera need to be reassessed.
has been found in greenhouses; it is not established in the wild in Cyphorhynchus Schoenherr 1837
North America. Glycaria Pascoe 1880
Archarias Dejean 1821 Edesius Pascoe 1881
Dionychus Germar 1824 Enops Pascoe 1889
Litomerus Schoenherr 1833 Loceptes Casey 1910
Polyderces Schoenherr 1833 Pseudocomus Varga 1951 (valid subgenus)
Aphyoramphus Guérin-Méneville 1844 (valid subgenus)
Lonchocerus Chevrolat 1879 Epacalles Kissinger 1964, 1 sp., E. inflatus Blatchley 1916, eastern
Sternoxus Chevrolat 1879 United States west into central Texas. Adults are collected in leaf
Platypachys Chevrolar 1879 litter.
Gymnonotus Chevrolat 1879
 Family 131. Curculionidae · 791

Lepilius Champion 1905, 1 sp., undescribed, extreme southwest- Eudociminus Leng 1918, 1 sp., E. mannerheimi (Boheman 1836),
ern Texas. Adults of an undescribed species have been collected southeastern United States west to Louisiana. Adults are associ-
in leaf litter in Big Bend National Park, Texas. ated with Taxodium distichum (L.) Rich. (bald cypress; Taxodiaceae).
Eudocimus Boheman 1836; not Wagler 1832
Micralcinus LeConte 1876, 3 spp., southeastern United States west
into Texas. Adults of M. cribratus LeConte 1876 have been asso- Heilipus Germar 1824, 1 sp., H. apiatus (Olivier 1807), Florida,
ciated with Amaranthus (Amaranthaceae) and adults of M. Georgia, South Carolina, North Carolina and Tennessee. Adults
maculatus (Blatchley 1916) with Sesuvium portulacastrum (L.) L. have been associated with various plants. In Florida the larvae
Aizoaceae (Anderson 1993a). I have not seen specimens of M. bore into the cambium at the base of Persea americana Mill. (Ameri-
kalmbachi Buchanan 1927. See Sleeper (1955c) to separate the spe- can avocado; Lauraceae) (Woodruff 1963). Two additional spe-
cies (note that M. stehri Sleeper 1955 is a junior synonym of cies, H. lauri Boheman 1845 and H. pittieri Barber 1919 have been
Conotrachelus parvulus Champion 1904; Wibmer and O’Brien found in greenhouses.
1989). Hilipus Agassiz 1846
Hilipus Gemminger and Harold 1871; not Agassiz 1846
Microhyus LeConte 1876, 1 sp., M. setiger LeConte 1876, eastern
United States and adjacent southern Canada. Adults have been Heilus Kuschel 1955, 1 sp., H. bioculatus (Boheman 1843), south-
associated with dead Fagus (beech; Fagaceae). ern Florida, adventive. Adults and larvae have been associated
Echinaspis Blatchley 1922; not Haeckel 1881 with Bursera simaruba (L.) Sarg. (Burseraceae) (Anderson 1993a).

Micromastus LeConte 1876, 1 sp., M. gracilis (Boheman 1859), Hilipinus Champion 1902, 1 sp., H. nearcticus O’Brien 1982, Florida,
California. Louisiana and Mississippi. Adults come to lights.

Pheloconus Roelofs 1875, 1 sp., P. hispidus (LeConte 1876) generally Hylobius Germar 1817, 8 spp., generally distributed throughout
distributed in the eastern United States west to Louisiana. Adults the eastern and central United States and all of Canada; one ad-
of P. hispidus (LeConte 1876) have been associated with Malvastrum ventive species. Adults and larvae of the native species are associ-
corchorifolium (Desc.) Britt. (Malvaceae). Two additional species, P. ated with conifers (Warner 1966). Hylobius transversovittatus (Goeze
infector (Boheman 1845) and P. cribricollis (Say 1831) have been 1777) has been introduced from Europe for the biological con-
considered as Pheleconus but do not fit the generic definition and trol of Lythrum salicaria (purple loosestrife; Lythraceae) and is
are likely Conotrachelus. See Blatchley and Leng (1916; as now established in New York, Pennsylvania, Maryland, Virginia,
Conotrachelus groups III and VI) to separate this complex of Ohio, Indiana, Illinois, Iowa, Michigan, Wisconsin, Minnesota,
species. South Dakota, Colorado, Montana, Oregon and Washington in
the United States (Weeden 2000), and in British Columbia, Alberta,
77. Cycloterini Lacordaire 1863 Manitoba and Nova Scotia in Canada (Harris 2001). See Warner
(1966) to separate the seven native species.
Cycloterina Lacordaire 1863 Callirus Dejean 1821 (valid subgenus)
Hypomolyx LeConte 1876
Gononotus LeConte 1876, 1 sp., G. angulicollis (Suffrian 1871), Hylobitelus Reitter 1923
Florida. Adults are common under debris on beaches (Anderson Poiyaunbus Kôno 1934
1993a).
Nemosinus Faust 1892 Pachylobius LeConte 1876, 1 sp., P. picivorus (Germar 1824), gener-
ally distributed in the eastern United States and adjacent southern
78. Erodiscini Lacordaire 1863 Canada. Adults are associated with Pinus (Pinaceae); larvae mine
the inner bark of roots and stumps of dying or injured trees
Alonso-Zarazaga and Lyal (1999) place these weevils as (Franklin and Taylor 1970).
Curculioninae but the structure of the uncus at the tibial apex
suggests they are Molytinae or related. Here they are placed as a 80. Lepyrini Kirby 1837
tribe within Molytinae.
Lepyrus Germar 1817, 6 spp., generally distributed in the north-
Sicoderus Vanin 1986, 1 sp., S. tinamus (LeConte 1884), Florida. ern and western montane United States and across Canada in-
This species appears to be associated with Bumelia celastrina (Nutt.) cluding the far north and Alaska. Adults are often associated with
R. W. Long (Sapotaceae) (Anderson 1993a). Salix (willow; Salicaceae) but larvae likely feed on the roots of
other plants (Anderson 1997). The genus needs revision. Several
79. Hylobiini Kirby 1837 subspecies of questionable status are recognized in North America
and no attempt has been made to compare the North American
Hylobiina Kirby 1837
792 · Family 131. Curculionidae

fauna to those of Asia. See Van Dyke (1928) to separate the 85. Sternechini Lacordaire 1863
forms.
Dirus Dejean 1821 Chalcodermus Dejean 1835, 7 spp., generally distributed in the east-
ern and central United States west into Texas and Arizona. Adults
81. Lymantini Lacordaire 1866 of C. aeneus Boheman 1837 have been associated with Vigna
luteola (Jacq.) Benth. (Fabaceae); larvae develop in seed pods
Caecossonus Gilbert 1955, 1 sp., C. dentipes Gilbert 1955, southern (Ainslie 1910). Adults of C. collaris Horn 1873 have been reared
Florida. Adults are frequently collected in soil and leaf litter from seed pods of Cassia chamaechrista L. (Fabaceae) (Alsterlund
(Howden 1992). 1937a, b). Adults of C. martini Van Dyke 1930 have been collected
from two species of Brickellia (Asteraceae) in Arizona and Texas.
Dioptrophorus Faust 1892, 1 sp., D. repens (Casey 1892), California, The genus needs revision. See Blatchley and Leng (1916) to sepa-
Oregon and Washington. Adults have been collected in leaf litter. rate some of the species.
Metopotoma Casey 1892 Anthobates Gistel 1848
Anculopus Van Dyke 1927
Sternechus Schoenherr 1826, 2 spp., S. armatus (Casey 1895), south-
Lymantes Schoenherr 1838, 4 spp., southeastern United States eastern United States north to Illinois and New Jersey, and S.
north to Ohio and west to western Texas and Oklahoma. Adults paludatus (Casey 1895), Arizona. Some tropical species are associ-
have been collected in leaf litter. The genus needs revision. See ated with Fabaceae (Anderson 1993b).
Sleeper (1965) to separate the species. Sternuchus Gemminger and Harold 1871; not LeConte 1850
Typhloglymma Dury 1901 Sternuchus Suffrian 1871; not LeConte 1850; not Gemminger
Stewpeckia Osella 1980 and Harold 1871
Plectromodes Casey 1895
82. Petalochilini Lacordaire 1863
86. Thalasselephantini Alonso-Zarazaga and Lyal 1999
Hormops LeConte 1876, 1 sp., H. abducens LeConte 1876, south-
eastern United States north to Ohio and west to Texas. Adults Thalasselephas Egorov and Korotyaev 1976, 1 sp., T. testaceus
are found in the nests of fox and grey squirrels (Blatchley 1918). (LeConte 1876), California, Oregon and British Columbia. Adults
are found under seaweed on sandy coastal beaches (Anderson
83. Piazorhinini Lacordaire 1863 1988b). Korotyaev and Egorov (1975) have suggested that this
genus is related to Emphyastes (Cyclominae).
Alonso-Zarazaga and Lyal (1999) place these weevils as Phycocoetes LeConte 1876; not Agassiz 1846
Curculioninae but the structure of the uncus at the tibial apex Neophycocoetes O’Brien and Wibmer 1982
suggests they are Molytinae or related. Here they are placed as a
tribe within Molytinae. 87. Trypetidini Lacordaire 1866

Piazorhinus Schoenherr 1835, 4 spp., generally distributed in east- Nanus Schoenherr 1844, 1 sp., N. uniformis Boheman 1844, south-
ern United States and southeastern Canada. Species are associated ern Florida. Adults are associated with palms and banana trees.
with Quercus (Fagaceae) and Coccoloba diversifolia Jacq. (Polygonaceae) Homaloxenus Wollaston 1873
(Anderson 1993a, b). See Blatchley and Leng (1916) to separate
the species. Incertae sedis
Acamatus Schoenherr 1833
Polyponus Kirsch 1875 Tranes Schoenherr 1843, 1 sp., T. internatus Pascoe 1870, Califor-
Piazorrhinus Champion 1903 nia, adventive. Adults have been collected in association with
introduced Encephalartos (Cycadaceae) from Australia. It is not
84. Pissodini Gistel 1856 known whether the genus is established in North America.
Platyphaeus Pascoe 1877
Pissodina Gistel 1856
XVII. Scolytinae Latreille 1807
Pissodes Germar 1817, 22 spp., generally distributed throughout
the United States and Canada. Adults and larvae are associated by Robert J. Rabaglia
with various conifers. Some species are of economic importance.
The genus needs revision. See Hopkins (1911) to separate the Subfamily common name: The bark and ambrosia beetles
species.
Piniphilus Dejean 1821 Subfamily synonyms: Hylurgidae Zimmerman 1868; Ipidae
Epipissodes Voss 1956 (valid subgenus) Latreille 1804
 Family 131. Curculionidae · 793

The general body shape of these small beetles ranges from very Habits and habitats. Most bark and ambrosia beetles live in
stout to moderately elongate and cylindrical. Typically the body is injured, weakened or dying woody plants. Hosts must contain
brownish with moderate pubescence. The geniculate antennae sufficient moisture for development and most species complete
have a distinct club. only one generation in a host. A few species breed in roots and
Description: (modified from Wood 1982) Very small to stems of non-woody plants, others breed in seed or cones, but
small in size, 1-9 mm, mostly 1-3 mm; shape stout to cylindrical; the majority of species are considered bark beetles or ambrosia
color brownish or piceous; pubescence sparse to abundant, mostly beetles. Bark beetles feed on the phloem of the inner bark of
consisting of very fine, short setae or stout, flat setae. their woody host plant. Fewer than half the species in the family
Head prominent, or withdrawn into pronotum; surface are bark beetles, but they are the majority of species in the tem-
punctate to granulate. Antennal scape well developed, funicle one perate regions. Ambrosia beetles cultivate and feed on symbiotic
to seven segmented, club large, either solid, annulated or rarely ambrosia fungi in the xylem of the host plant. Most tropical
pseudolamellate; inserted on the sides of head between eyes and species exhibit this habit.
mandibles. Labrum absent; mandibles short, curved, the apices Typically, adult bark and ambrosia beetles bore through the
blunt, dentate; maxillary palpi three segmented, segments short outer bark and construct an egg gallery either in the phloem-
and stout. Gular region reduced to a small pregula, gular sutures cambial region (bark beetles) or in the xylem (ambrosia beetles).
confluent; mentum moderate, variable; labial palpi three seg- Females lay eggs at regular intervals on either side of the gallery.
mented, small, stout, apically acute. Eyes lateral, moderate, flat, Among bark beetles, larval feeding mines radiate out from the
transverse. egg gallery, and engrave the inner bark or wood or both. These
Pronotum slightly broader than head; shape truncate characteristic engravings can often be found under the bark of
anteroventrally, quadrate to subcircular, borders margined or not; dead or dying trees. Ambrosia beetle larvae feed on the ambrosia
surface punctate, asperate, rugose or striate; pleural region broad; fungus in small cradles off of the egg gallery. After pupation, the
prosternum short in front of coxae, some with a small median next generation of bark beetles emerges through individual exit
process projecting posteriorly; procoxal cavities closed behind. holes in the bark, giving it a characteristic “shot hole” appearance.
Legs moderate in length; trochantins not exposed; anterior coxae Ambrosia beetle adults usually emerge through the parental en-
globular, contiguous to widely separated; middle coxae round, trance hole.
flat, separate; hind coxae subtriangular, separate; trochanters small, Most of the life stages of these beetles occur within the host
triangular; femora swollen, short; tibiae compressed, mostly plant, however, upon emergence adults must find suitable host
toothed with apical hooks or, with marginal teeth or denticles; material in which to feed and breed. They are often among the
tarsal formula 5-5-5, apparently 4-4-4, slender, third segment nar- first insects to colonize a dying tree; therefore, rapid location of
row or dilated, fourth segment minute; claws large, simple diver- hosts is an important part of their biology. In many species, host
gent. Scutellum small, quadrate, triangular or absent. Elytra en- location is mediated by olfactory responses to host odors (e.g.,
tire, apically rounded, mostly declivous and often with tubercles, terpene hydrocarbons), tree degradation products (e.g., alcohols)
denticles or spines apically; striae mostly distinct, punctate; or conspecific semiochemicals (pheromones). Several species uti-
epipleural fold obscure. Wing venation and folding pattern not lize pheromones not only for attraction of potential mates, but
described. also for mass aggregation to overcome resistance of the host tree.
Abdomen with five visible sterna, sutures entire; surface The pheromone biology of species of Dendroctonus, Ips and
microrugose to punctate. Male genitalia with penis stout, apically Scolytus, among others, has been well studied, and the complex
blunt, basally with a pair of slender, articulating struts; parameres inter- and intraspecific interactions elucidated (Wood, D.L. 1982,
absent; pars basalis reduced to a slender complete or incomplete Borden 1982, Raffa et al. 1993).
ring and a curved, slender basal strut of variable length. Female Many bark and ambrosia beetle species have distinctive, sub-
genitalia undescribed. social behaviors. Social organization associated with reproductive
Larvae C-shaped, subcylindrical, fleshy; size 2 mm - 10 mm behavior ranges from simple monogamy to heterosanguineous
in length; vestiture ranges from absent to a few, simple setae; polygyny to consanguineous polygyny. Division of labor in gal-
color near white. Head partly retracted or distinctly exserted, lery construction and maintenance is marked by sexual dimor-
mouthparts hypognathous or nearly prognathous with a faint phism, especially in structures on the head and elytral declivity.
epicranial suture surrounding the frons. Antennae very small to Ecologically and economically this is a very important group
absent. Mandible mostly short, stout, gouge-shaped, subtrian- of beetles. Members of Dendroctonus and Ips kill or degrade vast
gular without mola or retinaculum; maxilla with cardo, fused expanses of forest each year. Species of Scolytus are well known as
stipes and mola; maxillary palpi one or two segmented. Stem- vectors of the Dutch elm disease fungus. In the tropics, ambro-
mata absent in most. Thorax frequently broader than abdomen; sia beetles stain and degrade valuable wood products. In North
legs absent, but with fleshy lobes ventrally. Abdomen with three America, several species of exotic xyleborines cause damage to
or more plicae on each segment; nine or ten segmented, seg- young, stressed trees in the landscape and nurseries, and species
ments 8 -10 in some with pigmented tubercles dorsally. Spiracles of Gnathotrichus, Monarthrum and Trypodendron degrade wood
on mesothorax and abdominal segments one to eight, annular, products in the Pacific Northwest (Furniss and Carolin 1977).
annular-biforous or biforous, or inconspicuous.
794 · Family 131. Curculionidae

There have been numerous studies on the biology, chemical ecol- beyond spine of inner apical angle; antennal fu-
nicle 7-segmented; lateral prosternal area bear-
ogy and control of many of the economically important genera.
ing a sharply elevated ridge from coxa to anterior
Status of the classification. This book treats bark and margin; crenulations on elytral bases rather small
ambrosia beetles as a subfamily of Curculionidae following (Bothrosternina) ................................................ 3
Crowson (1967); however, the following Key and Classification — Prothorax punctate or asperate, never longitudinally
strigose; all tibiae bearing several teeth, none
of Tribes and Genera follow Wood (1973) and a family catalog by
extending beyond tarsal insertion; antennal fu-
Wood and Bright (1992), but with the status of the subfamilies nicle and prosternal area variable ................... 4
and tribes reduced to tribes and subtribes. See Wood (1973 and
1986) for a discussion of this issue.
Bark and ambrosia beetles occur on all continents except
Antarctica. In North America, the fauna has been well studied
within the past century, and is now well known. Wood (1982)
published a monograph on the bark and ambrosia beetles of
North and Central America, including a key to all genera and
species in the region (at the time about 1430 species were recog-
nized). Wood and Bright (1992) published a catalog of the world-
wide Scolytidae, followed by a recent update (Bright and Skidmore
1997). The taxonomic status of tribes and genera in the Key and
Classification sections below follows these catalogs. The two ex-
ceptions are the new genera Dryoxylon Bright and Rabaglia (1999)
and Pseudips Cognato (2000) which have been added to the key.
Distribution. There are approximately 5,800 species world-
wide, with about 525 species and subspecies described from the
United States and Canada. Bark beetles can be found from the
subalpine forests of the north to the subtropical forests of Florida. 102 103
Distinctive faunas exist in the desert plateau of the southwest,
the deciduous forests of the southeast, the northern coniferous
forests, the Pacific Coast and southern Florida. Bark beetles tend
to be more restricted by host than ambrosia beetles. Within a
bark beetle genus, most species are restricted to a limited number
of host species; for example, Phloeosinus are found mostly in
Cupressaceae and Pseudopityophthorus are found almost exclusively
in Quercus. 105
104
Wood (1977) estimated that there were 37 Old World spe-
cies established in North and Central America. Since then ap-
proximately 10 additional species new to North America have
been reported (Atkinson et al. 1990, Hoebeke 1991, Haack and
Kucera 1993, Rabaglia and Cavey 1994, Bright and Rabaglia 1999,
Vandenberg et al. 2000, Hoebeke 2001 and Mudge et al. 2001).
106 107

KEY TO THE NEARCTIC GENERA OF SCOLYTINAE

(Modified from D. E. Bright, unpublished 2000)

1. Anterior margins of elytra procurved and bearing a

series of crenulations; pronotum unarmed in
most; head visible from above (Fig. 102)
(Hylesinini) ........................................................ 2 108
— Anterior margins of elytra forming a straight line 109
across body, unarmed, smooth and either
rounded or with a fine raised line; pronotum, in
most, armed by granules or asperities on at least FIGURES 102.131-109.131. Scolytinae. 102-103. Dorsal habitus,
anterior third; head concealed from above (Fig. 102. Dendroctonus pseudotsugae Hopkins; 103. Dr yocoetes affaber
103) (Scolytini) ............................................... 24 Mannerheim. 104-105. Protibia, 104. Scolytus sp.; 105. Procryphalus sp.
106-107. Lateral habitus, 106. Hypothenemus sp.; 107. Pityophthorus sp.
2(1). Prothorax longitudinally strigose; prothoracic tibia 108-109. Antennal club, 108. Cryptocarenus sp.; 109. Hypothenemus sp.
with a curved bifid process, meso- and metatho- (Figures 102-103 after Swaine 1918; Figure 104 after Chamberlin
racic tibiae with a single curved spine extending 1958; Figures 105-109 after Wood 1982.)
 Family 131. Curculionidae · 795

3(2). Sutures of antennal club straight; rostrum distinctly accommodate elytral margins; xylophagous spe-
wider than distance between eyes; body and cies (Phloeosinina, part) ................. Dendrosinus
frons not as below ............................... Cnesinus — Scutellar notch between elytra emarginate, but not
— Sutures of antennal club procurved; rostrum width deeply grooved; elytra not extended anteriorly,
at tip equal to distance between eyes; body oval; pronotum not grooved; phloeophagous species
frons excavated, with median tubercle just above (Tomicina) ....................................................... 13
epistoma ........................................... Pagiocerus
13(12). Fore coxae widely separated ............................ 14
4(2). Prothoracic precoxal area rather large, lateral mar- — Fore coxae contiguous, or at most very narrowly
gin strongly elevated from anterior margin to coxa separated ....................................................... 16
......................................................................... 5
— Prothoracic precoxal area short, lateral prosternal 14(13). Elytral vestiture hair-like; antennal club slightly flat-
ridge poorly developed or absent ................... 8 tened, segment 1 occupying one-fourth of club
length; in Ulmus ........................... Hylurgopinus
5(4). Crenulations on elytral bases forming a single row — Elytral vestiture scale-like; antennal club conical,
of teeth; first and second segments of antennal segment 1 occupying less than one-fourth of
club subequal in length; body rather stout, length club length; in conifers .................................. 15
less than 2.5 mm; in roots of herbaceous legumes
(Hylesinina, part) ............................... Hylastinus 15(14). Each elytral interspace bearing a row of erect, flat-
— Crenulations on elytral bases obsolete, if visible, tened scales in addition to recumbent ground
then irregularly placed, not forming a definite cover; antennal funicle 5-segmented ...............
single row; first segment of antennal club dis- .......................................................... Xylechinus
tinctly longer than second; body mostly larger — Elytral interspaces bearing a row of erect, hairlike
than 3 mm, very slender if smaller; not in herba- setae, ground cover scale-like or stout setae; an-
ceous legumes (Hylastina) ............................... 6 tennal funicle 7-segmented .... Pseudohylesinus

6(5). Anterior coxae widely separated; surface of elytra 16(13). Antennal funicle 5-segmented; antennal club with
and between punctures on pronotum dull; sutures slightly procurved; anterior margin of
vestiture sparse, recumbent, yellow; body color pronotum distinctly emarginate; 2.5-9.0 mm in
dull reddish brown ................................. Scierus length .......................................... Dendroctonus
— Anterior coxae narrowly separated, almost contigu- — Antennal funicle 6-segmented .......................... 17
ous; surface between punctures on pronotum
and elytra smooth and glossy; the longer hairlike 17(16). Elytra with erect interstrial setae abundant, ran-
vestiture erect; mature color glossy, dark brown domly placed; a short median carina on frons ex-
or black ............................................................. 7 tending from epistomal margin to level of anten-
nal insertion, ending dorsally in an acute eleva-
7(6). Pronotum, in most, constricted anteriorly, discal sur- tion; elytra densely rugose ................. Hylurgus
face with about equal numbers of small and large — Elytra with erect interstrial setae in uniseriate rows;
punctures intermixed; third tarsomere broad, bi- a fine median carina on frons extending from
lobed ................................................. Hylurgops epistoma to middle of frons, of equal height
— Pronotum not noticeably constricted anteriorly, throughout; elytra smooth ................... Tomicus
discal surface with punctures uniformly large,
with very few small punctures; third tarsomere 18(9). Antennal club pseudolamellate, constricted at su-
narrower, emarginate .......................... Hylastes tures and movable at intersegmental lines
(Phloeotribina) ................................ Phloeotribus
8(4). Scutellum visible, elytral bases notched for its re- — Antennal club fused at sutures, sutures oblique or
ception ............................................................. 9 partly to entirely obsolete (Phloeosinina, part) ..
— Scutellum not visible, elytral bases straight ..... 20 ....................................................................... 19

9(8). Antennal club symmetrical, sutures transverse .... 19(18). Antennal club with three oblique sutures; funicle
....................................................................... 10 attached to base of club; pronotum unarmed; eye
— Antennal club with sutures oblique, deeply emarginate; hosts Cupressinine trees,
pseudolamellate or absent ............................. 18 rarely other conifers ....................... Phloeosinus
— Antennal club solid and unmarked by sutures; fu-
10(9). Pronotum asperate on anterolateral areas nicle attached to side of club; pronotum, in most,
(Hylesinina, part) ............................................ 11 armed by a few asperities in anterolateral areas;
— Anterolateral areas of pronotum unarmed ......... 12 eye entire; hosts mostly hardwoods ................
......................................................... Chramesus
11(10). Eye entire; vestiture scalelike; costal margins of
elytra ascending slightly at apex, abdomen as- 20(8). Eye emarginate or completely divided; pronotum
cending to meet them; hosts Fraxinus species never armed by asperities; crenulations at bases
............................................................ Hylesinus of elytra widely distributed, extending laterally
— Eye shallowly emarginate; vestiture hairlike; costal beyond interstriae 5; antennal funicle 5- or 6-seg-
margins of elytra descending to apex, abdomen mented (Polygraphina) .................................... 21
horizontal; hosts Alnus species ...... Alniphagus — Eye sinuate or entire; pronotum armed by a few
scattered or clustered asperities; crenulations at
12(10). Scutellar notch between elytra very deep, acute; bases of elytra restricted to area between suture
elytra extended anteriorly over pronotum, pos- and interstriae 5; antennal funicle 4- or 5-seg-
terolateral area of pronotum abruptly grooved to mented (Hypoborina) ...................................... 23
796 · Family 131. Curculionidae

21(20). Eye completely divided into two parts; antennal on outer apical angle exceeding tarsal insertion;
club solid, unmarked by sutures ... Polygraphus procoxae separated; (Micracina) . .................. 30
— Eye less than half divided by an emargination; an- — Fore tibia much wider apically, armed on lateral mar-
tennal club marked by sutures ...................... 22 gin by several denticles; procoxae contiguous
....................................................................... 36
22(21). Antennal funicle 5-segmented ......... Carphoborus
— Antennal funicle 6-segmented ........... Carphobius 30(29). Antennal club small, greatest width through basal
half, apex narrowly rounded, sutures straight,
23(20). Antennal funicle 4-segmented, sutures of club in- transverse ...................................................... 31
dicated only by marginal notches; elytra with — Antennal club larger, greatest width through apical
uniseriate rows of erect, broad interstrial scales half, apex broadly rounded, sutures procurved
and recumbent strial hair of equal length; ....................................................................... 32
pronotum armed by 3 or 4 pairs of median tu-
bercles, the anterior pair marginal .. Liparthrum 31(30). Elytral declivity subvertical, bisulcate, obtusely
— Antennal funicle 5-segmented, sutures of club angulate behind; sutures of antennal club dis-
transverse, distinct; elytral vestiture without con- tinctly marked by rows of setae; antennal pedicle
spicuous recumbent hair; pronotum armed by 2 and scape about equal in length ... Stenocleptus
or 3 widely separated paired clusters of lateral — Elytral declivity more gradual, evenly convex, rather
teeth ........................................... Chaetophloeus narrowly rounded behind; sutures of antennal
club indicated only by marginal notches; scape
24(1). Lateral margin of anterior and posterior tibia unarmed distinctly longer than pedicle ...........................
except for a single curved process at outer api- ...................................... Pseudothysanoes (part)
cal angle that curves toward and extends be-
yond process of inner apical angle (Fig. 104); lat- 32(30). Elytra broadly rounded behind; margins of antennal
eral line of pronotum sharply elevated; antennal club, in most, constricted at first suture ....... 33
club flattened, the sutures strongly procurved; — Elytra acuminate behind; antennal club without su-
antennal funicle 7-segmented (Scolytina) ..... 25 tural constrictions at sides ............................ 34
— Lateral margin of anterior tibia armed by several
toothlike processes, none of which curve toward 33(32). Pronotum wider than long, widest near base, sum-
the inner process (Fig. 105); lateral line of mit more prominent; fore tibia more slender,
pronotum raised or not; antennal club and funicle apically obliquely truncate, mucro often bifur-
variable ........................................................... 26 cate ............................... Psuedothysanoes (part)
— Pronotum longer than wide, widest near middle, sum-
25(24). Elytra slightly if at all declivous behind, the abdo- mit less prominent; fore tibia rather broad, more
men ascending abruptly behind to meet them; nearly truncate apically, mucro undivided ........
scutellum depressed; antennal scape distinctly .......................................................... Thysanoes
shorter than funicle .............................. Scolytus
— Elytral declivity rather steep, descending to meet 34(32). Sutures of antennal club broadly procurved, the
the horizontal abdomen; scutellum small, flush first appearing bisinuate and extending less than
with surface of elytra; antennal scape at least as one-third length of club; scape club-shaped, with
long as funicle ................................. Cnemonyx few setae; eye oval, rather small; fore tibia more
slender, slightly wider apically, with supplemen-
26(24). Metepisternum visible to posterior extremity (Fig. tal tubercles on posterior face ......... Hylocurus
106); antennal club varying from flat to obliquely — Sutures of club very strongly, narrowly procurved,
truncate; tibia and antennal funicle variable ..... the first most often reaching middle of club;
....................................................................... 27 scape compressed, subtriangular, with numerous
— Metepisternum largely covered by elytra, visible long setae; eye elongate, large; fore tibia broad,
only in front (Fig. 107); antennal club strongly sides subparallel, posterior surface devoid of tu-
flattened with sutures on both sides, those on bercles except for teeth on apical margin .... 35
posterior surface not strongly displaced apically;
tibia slender, in most, bearing about three teeth 35(34). Eyes moderately separated beneath, entire; fore
on apical portion; antennal funicle 1- to 5-seg- tibia with all five teeth on distal margin, mucro
mented ........................................................... 66 broad .................................................... Micracis
— Eyes subcontiguous beneath, emarginate; fore tibia
27(26). Lateral margins of prothorax subacutely elevated; with at least one of the five teeth on outer mar-
procoxae widely separated (Ctenophorina) .. 28 gin, mucro more slender ................ Micracisella
— Lateral margins of prothorax rounded; procoxae
subcontiguous ............................................... 29 36(29). Male frons bearing a very large, long, partly double
process which may curve upward and backward
28(27). Anterior area of pronotum transversely rugose; over prothorax, in some, reaching its posterior
pronotum and elytra subglabrous .... Scolytodes margin; pronotum asperate to base in median
— Pronotum uniformly punctured, unarmed; vestiture area, summit on basal third, in most, extending
of pronotum and elytra abundant, consisting of behind its basal margin and over scutellum; body
erect, stout, almost scalelike bristles ............... usually covered by an incrustation (Cactopinina)
...................................................... Pycnarthrum ......................................................... Cactopinus
— Male frons not armed by a large median process;
29(27). Fore tibia with sides parallel, in most, armed only on pronotal summit at or slightly behind middle of
apical margin by small teeth never with process prothorax, basal third devoid of asperities .... 37
 Family 131. Curculionidae · 797

37(36). Antennal club more strongly flattened, with sutures 45(44). Antennal club not septate; raised lateral margin of
on both faces, those on posterior face strongly pronotum extending two-thirds of distance from
procurved and limited to apical half; costal mar- basal margin; elytra glabrous except for a few
gins of elytra slightly ascending posteriorly; subcapitate interstrial bristles ... Cryptocarenus
vestiture scale-like (Cryphalina) .................... 38 — Antennal club with suture 1 partly septate; raised
— Antennal club obliquely truncate or at least with lateral margin extending only one-third of dis-
sutures of posterior face restricted to less than tance from basal to anterior lateral margin; elytra
apical one-fourth; costal margins of elytra de- clothed by rows of strial and interstrial setae ...
scending posteriorly; vestiture hairlike setae ... ................................................... Hypothenemus
....................................................................... 46
46(37). Antennal funicle 2- or 3-segmented; pronotum un-
38(37). Pronotum without a fine, raised lateral line; eye, in armed, punctured over entire surface, lateral line
some, sinuate, never emarginate; costal margins not sharply raised; length 2.0 mm or less
of elytra ascending only slightly posteriorly .... (Crypturgina) ................................................... 47
....................................................................... 39 — Antennal funicle 4- or 5-segmented; pronotum
— Pronotum acutely margined at sides, and with a fine, mostly armed anteriorly by granules or asperi-
raised line at least on basal one-third; eye emar- ties, if unarmed, lateral line sharply raised; length
ginate or entire; costal margins of elytra distinctly mostly over 2.0 mm ........................................ 48
ascending posteriorly .................................... 42
47(46). Antennal funicle 2-segmented, club with 1 obscure
39(38). Antennal funicle 5-segmented; antennal club nar- suture indicated at tip ..................... Crypturgus
row, pointed at tip, sutures straight, not septate; — Antennal funicle 3-segmented, club with 3 sutures
basal half of pronotum without scale-like setae ............................................................. Dolurgus
...................................................... Trypophloeus
— Antennal funicle 4-segmented; antennal club 48(46). Eye completely divided by an emargination; anten-
broadly rounded at tip, sutures curved, partly nal funicle 4-segmented, club without distinct
septate or not septate; basal half of pronotum sutures (Xyloterina) ........................................ 49
with scalelike setae ....................................... 40 — Anterior margin of eye sinuate or emarginate, never
completely divided; antennal funicle 4- or 5-seg-
40(39). Antennal club not septate, sutures indicated by 3 mented, club, in most, with evident sutures .....
strongly procurved rows of setae (Fig. 108) ..... ....................................................................... 50
......................................................... Ernoporicus
— Antennal club with at least part of first suture sep- 49(48). Antennal club with subcorneous basal area
tate, none of sutures indicated by strongly strongly, rather narrowly procurved; protibia of
procurved rows of setae (Fig. 109) ................ 41 female thickened and tuberculate on posterior
face, flattened and finely tuberculate in male;
41(40). Sutures of antennal club straight, the first septate; male head deeply, broadly excavated, the pro-
anterior margin of pronotum slightly produced; thorax sub-quadrate; female frons convex, ante-
pronotum with no indication of a fine raised lat- rior margin of female pronotum rounded ...........
eral margin .................................... Procryphalus ..................................................... Trypodendron
— Antennal club with a strongly oblique septum on — Antennal club with subcorneous basal area broadly
one side, no other sutures indicated; anterior procurved; protibia flattened and devoid of tu-
margin of pronotum broadly rounded; pronotum bercles on posterior face; frons not excavated in
with an indistinct, fine, raised lateral line .......... either sex; anterior margin of prothorax rounded
...................................................... Scolytogenes in both sexes ................................... Xyloterinus

42(38). Antennal club with sutures indicated by rather 50(48). Pronotum either punctate or else finely granulate
strongly recurved rows of setae; third tarsomere over almost entire surface, dorsal profile evenly
broad and emarginate ........................ Cryphalus convex, not strongly declivous anteriorly, ante-
— Sutures of antennal club straight or procurved; third rior margin never armed; tibia rather slender and
tarsomere cylindrical ..................................... 43 armed by few, coarse teeth; declivity unarmed
(Dryocoetina) .................................................. 51
43(42). Eye entire; antennal club large, aseptate, funicle — Pronotum coarsely asperate and strongly declivous
normally 3-segmented, rarely 4-segmented; body anteriorly, in most, punctate at least on posterior
stout, less than 2.3 times longer than wide; body third, in some, anterior margin armed; tibia vari-
shorter than 1.1 mm .......................... Trischidias able; declivity frequently armed by spinous
— Eye emarginate; antennal funicle 5-segmented, processes ...................................................... 55
rarely 4- or 3-segmented; in most, body longer
than 1.1 mm .................................................... 44 51(50). Antennal club compressed or with membranous api-
cal portion extended beyond corneous portion,
44(43). Strial punctures obscure, not impressed; posterior sutures procurved; scutellum very small ...... 52
half of pronotum finely granulate; antennal club — Antennal club subtruncate, sutures transverse or
large, not septate; male and female similar in size recurved; scutellum moderate to large ......... 53
and appearance .......................... Hypocryphalus
— Strial punctures distinct; posterior half of pronotum 52(51). Antennal funicle 4-segmented; club compressed,
not closely granulate, in most, punctate; male sutures strongly arcuate; pronotum granulate on
much smaller than female .............................. 45 anterior half, punctate behind; host Acer ..........
........................................................... Lymantor
798 · Family 131. Curculionidae

— Antennal funicle 5-segmented; club less strongly 59(58). Lateral margins of elytral declivity armed by 4 to 6
compressed, sutures rather broadly procurved; pairs of spinelike denticles; ventrolateral margin
pronotum granulate to base; host Cucurbita ..... of elytral declivity very strongly produced, cir-
.................................................. Dendrocranulus cumscribing an arc much less than one-third of a
circle, its lateral extremities ending a long dis-
53(51). Frons convergently aciculate; elytral declivity tance from largest denticle; sutures 1 and 2 of
evenly convex, extending over at least poste- antennal club weakly bisinuate to strongly
rior one-third of elytra, granules absent; protibia angulate ............................................... Ips (part)
armed on lateral margin by 2-4 socketed teeth; — Lateral margins of elytral declivity armed by 3 pairs
posterior face of antennal club with 2 sutures . of spinelike denticles; ventrolateral margin of
....................................................... Coccotrypes elytral declivity only slightly to moderately pro-
— Frons never convergently aciculate; elytral decliv- duced, circumscribing an arc at least one-third of
ity flattened or impressed, confined to posterior a circle, its lateral extremities ending near third
one-fourth of elytra, granules mostly present; (last and largest) denticle; sutures 1 and 2 of an-
protibia armed on lateral margin by 5 or more sock- tennal club weakly to very strongly, broadly
eted teeth; posterior face of antennal club with- procurved ...................................................... 60
out sutures or with 1 suture .......................... 54
60(59). Sutures on antennal club weakly procurved, almost
54(53). Pronotum 1.4 times longer than wide, anterior mar- straight; strial punctures at least twice as large
gin slightly notched or emarginate; elytral decliv- as those of interstriae, in clearly defined rows;
ity moderately deeply, evenly sulcate ............. spine 3 on elytral declivity cylindrical or conical,
........................................................... Dryoxylon not constricted before apex; body length 2.3-3.6
— Pronotum 1.0-1.2 times longer than wide, anterior mm ............................... Ips (part, latidens group)
margin evenly rounded; elytral declivity evenly — Sutures on antennal club very strongly procurved;
convex to slightly flattened, may have second strial and interstrial punctures subequal in size,
interspace impressed ...................... Dryocoetes not always in clearly definable rows; spine 3 on
declivity subcapitate, distinctly constricted be-
55(50). Meso- and metathoracic tibia rather slender, fore apex; body length 3.5-5.0 mm ...... Pseudips
abruptly narrowed apically, armed by a few rather
widely spaced coarse teeth; males and females 61(55). Antennal club more strongly compressed, corneous
similar in size and general shape (Ipina) ........ 56 area small, near base, its distal margin strongly
— Meso- and metathoracic tibia rather broadly dilated procurved, distal pubescent portion reaching
to a point slightly beyond middle then gradually basal one-fifth at sides; pregula not impressed;
narrowed to apex, and armed by a series of small elytra obliquely truncate behind, declivity
closely set teeth of more or less uniform size and broadly, concavely excavated and acutely margi-
shape; males rare, in most, smaller and radically ned on a complete circle at periphery .............
different in shape (Xyleborina) ....................... 61 ......................................................... Premnobius
— Antennal club thickened basally, corneous area
56(55). Elytral declivity rather narrowly bisulcate, margins larger with its distal margin recurved, pubescent
moderately elevated, rounded and armed by not area not reaching basal third; pregula depressed;
more than 3 teeth; lower margin of declivity elytral declivity convex, not acutely margined
rounded; in most, body shorter than 3.0 mm ..... on upper half .................................................. 62
....................................................................... 57
— Elytral declivity broadly, rather deeply excavated, 62(61). Procoxae widely separated; body stout, elytra less
margins acutely elevated and armed by 3 or more than 1.3 times as long as pronotum ..................
tubercles or teeth; lower margin of declivity with ....................................................... Xylosandrus
an acutely elevated transverse ridge separating — Procoxae contiguous; body elongate, often slen-
declivital excavation from apical margin; body der, elytra at least 1.5 times as long as pronotum
mostly longer than 3.0 mm ............................. 58 ....................................................................... 63

57(56). Female frons deeply, rather narrowly excavated; 63(62). Pronotum wider than long, subquadrate, anterior
male declivity with 2 or 3 pairs of enlarged teeth; margin unarmed .............................................. 64
antennal club compressed, 2 sutures visible on — Pronotum longer than wide, subcircular, anterior
distal third of posterior face ............. Pityogenes margin armed by a series of median serrations .
— Female frons convex; male declivity more narrowly ....................................................................... 65
impressed with 2 or 3 pairs of very small teeth or
granules; antennal club obliquely truncate, with- 64(63). Pronotum asperate to base; declivity steep, bear-
out sutures on posterior face ........ Pityokteines ing several granules or rather large denticles,
strial and interstrial punctures small ..................
58(56). Antennal club obliquely truncate, with sutures re- ................................................... Ambrosiodmus
curved; elytral declivity less strongly excavated, — Pronotum asperate only on anterior half, punctate
the third tooth displaced mesally, not on summit on basal half; declivity more sloping, bearing small
of declivital margin ...................... Orthotomicus tubercles, strial and interstrial punctures larger
— Antennal club flattened, with sutures procurved or .......................................................... Euwallacea
strongly bisinuate; elytral declivity broadly ex-
cavated, armed by 3 to 6 major denticles, all den- 65(63). Scutellum conical; lower margin of declivity, be-
ticles on summit of lateral margin .................. 59 ginning about interspace 7, bearing a series of
pointed tubercles, the one nearest suture (at end
of interspace 2) largest ................... Xyleborinus
 Family 131. Curculionidae · 799

— Scutellum flat; lower margin of declivity acute or 73(66). Antennal funicle 5-segmented, club smaller, less
rounded, unarmed ............................. Xyleborus than twice as long as funicle ..... Gnathotrichus
— Antennal funicle 1- or 2-segmented; club very large,
66(26). Antennal funicle 5-segmented (3-segmented in more than three times as long as funicle ....... 74
Dendroterus) club mostly small, symmetrical; pu-
bescence more abundant; bark or twig beetles 74(73). Antennal funicle 2-segmented; posterior surface of
(Pityophthorina) .............................................. 67 fore tibia tuberculate; elytra emarginate or
— Antennal funicle 1-, 2-, or 5-segmented, club much divaricate at sutural apex .............. Monarthrum
larger, asymmetrical in most; pubescence less — Antennal funicle 1-segmented; posterior surface of
abundant; ambrosia beetles (Corthylina) ....... 73 fore tibia smooth; elytra evenly rounded behind,
without a sutural notch at apex ......... Corthylus
67(66). Basal and lateral margins of prothorax rounded, with-
out a fine raised line; antennal club somewhat
large in size; vestiture shorter and more uniform
in length ......................................................... 68 CLASSIFICATION OF THE NEARCTIC SCOLYTINAE
— Basal and posterior portion of lateral margins of pro-
thorax with an obvious, fine, raised line; anten- 88. Hylesinini Erichson 1836
nal club proportionately smaller; most with
vestiture longer on declivity than on disc .... 69
Hylastina Leconte 1876
68(67). Antennal funicle 3-segmented; club less than twice
as long as funicle; female pronotum without Scierus LeConte 1876, 2 spp., northern and western North America
patches of pilose pubescence; elytral pubes-
in Picea; usually found in the phloem of roots and stumps of
cence abundant ............................ Dendroterus
— Antennal funicle 5-segmented, club at least twice standing dead trees or next to the ground in boles of downed
as long as funicle; female prothorax with a pair of trees.
pilose pubescent areas on middle third of lateral
areas; elytral pubescence sparse ... Pityoborus
Hylurgops LeConte 1876, 6 spp., 2 with subspecies, throughout
69(67). Antennal club devoid of sutures except for one coniferous forests of North America; all species breed in the
strongly oblique septum on anterior half of club phloem of stumps, roots and souring logs. The genus is closely
only; prothorax evenly rounded in dorsal profile, related to Hylastes from which some species are distinguished
summit inconspicuous, asperities fine, transition
with difficulty.
from asperate to punctate area gradual ............
.............................................................. Araptus Hylesinites Germar 1813
— Antennal club with at least two complete sutures Hylastities Hagedorn 1906
indicated at least by setae; prothorax more Myelophites Hagedorn 1906
strongly declivous anteriorly, summit and arrange-
Hylescierites Schedl 1947
ment of asperities variable ............................. 70

70(69). Sutures of antennal club not septate; in most, Hylastes Erichson 1836, 14 spp. in Pinaceae throughout America
pronotal asperities extending behind middle at north of Mexico, H. opacus Erichson 1836 is an adventive from
sides, the transition from asperate to punctate
Europe. All species breed in the phloem of stumps and roots.
area gradual; body moderately to very stout ....
..................................................... Conophthorus
— First and second sutures of antennal club septate; Hylesinina Erichson 1836
pronotal asperities mostly not reaching middle,
the transition from asperate to punctate area usu-
Hylastinus Bedel 1888, 1 sp., H. obscurus (Marsham 1802), native
ally abrupt, summit usually well developed; body
slender to moderately stout .......................... 71 to Palearctic, now found throughout North America. Breeds in
roots of legumes, especially Trifolium species.
71(70). Pronotum and elytra minutely densely punctured;
vestiture very short, mostly dense, almost always
Alniphagus Swaine 1918, 2 spp. in western North America (an
scalelike; antennal club with first segment shorter
than others; greater development of frontal additional species occurs in east Asia). All species breed in phloem
vestiture a male character; hosts Quercus, rarely of Alnus species.
other broadleaf trees ...... Pseudopityophthorus Hylastinoides Spessivtev 1919
— Pronotum and elytra more coarsely, less densely
punctured; vestiture usually longer, less abun-
dant, always hairlike; greater development of fron- Hylesinus Fabricius 1801, 7 spp. throughout America north of
tal vestiture a female character; hosts usually co- Mexico in mostly Fraxinus hosts. Adults and larvae deeply mine
nifers, but also broadleaf trees and shrubs ... 72 the wood in the phloem-cambial area. Adults construct biramous
galleries and larvae mine parallel to the grain of the wood.
72(71). Pregular area greatly enlarged and ornamented by
a beard-like brush of exceedingly long hair ...... Leperisinus Reitter 1913
........................................................ Pityotrichus Apidocephalus Wickham 1916
— Pregular area small, without conspicuous vestiture
..................................................... Pityophthorus
800 · Family 131. Curculionidae

Tomicina Thomson 1859 the phloem-cambial region of hosts. P. liminarus (Harris 1852)
occasionally is a pest of Prunus.
Hylurgopinus Swaine 1918, 1 sp., H. rufipes (Eichhoff 1868) occurs Phloeophthorus Wollaston 1854
east of the Rocky Mountains. This phloeophagous species breeds Dryotomus Chapuis 1869
in large branches and boles of Ulmus, and is a vector of the Dutch Phthorophloeus Rey 1885
elm disease fungus. Elzearius Guillebeau 1893
Eulytocerus Blandford 1897
Pseudohylesinus Swaine 1917, 9 spp., 2 with subspecies, occur in west- Comesiella DelGuercio 1925
ern North America (2 additional species occur in Mexico). They breed Neophleotribus Eggers 1943
in the phloem of limbs, boles and roots of weakened conifers. Dryotomicus Wood 1962

Xylechinus Chapuis 1869, 2 spp. occur in northern and western Phloeosinina Nusslin 1912
North America coincident with their Picea hosts. They are
phloeophagous in small, weakened trees. Dendrosinus Chapuis 1869, 1 sp., D. bourreriae Schwarz 1920, in the
Pruniphagus Murayama 1958 Florida Keys (nine additional species occur in Central and South
Squamosinus Nunberg 1964 America). Adults and larvae feed in the wood of small woody
Xylechinops Browne 1973 plants.

Hylurgus Latreille 1807, 1 sp., H. ligniperda (Fabricius 1787), native Phloeosinus Chapuis 1869, 25 spp., 3 of which occur in the east and
to Europe, was recently found in cut pine stumps in New York the remainder in the west. All species, except P. pini Swaine 1915,
State (Hoebeke 2001). All species are native to Palearctic. attack Cupressaceae and Taxodiaceae. Adults construct longitudinal
galleries under the bark that usually deeply engrave the wood.
Tomicus Latreillle 1802, 1 sp., T. piniperda (L. 1758), native to Pale- Phloeosinites Hagedorn 1906
arctic, was first found in North America in 1992. It is now re-
corded from the Lake States, Maine, Maryland, New Hampshire, Chramesus LeConte 1868, 9 spp. are found north of Mexico. These
New York, Pennsylvania, Vermont, West Virginia, Ontario and small beetles are phloeophagous in twigs and small branches of
Quebec. Adults feed in shoots of Pinus and breed in boles of hardwood trees and shrubs.
weakened or downed trees. Rhopalopleurus Chapuis 1869
Blastophagus Eichhoff 1864 Thaumasinulus Reitter 1913
Myelophilus Eichhoff 1878 Prochramesus Wood 1956

Dendroctonus Erichson 1836, 13 spp. found throughout America Hypoborina Nusslin 1911
north of Mexico. Most species breed in the boles of conifers and
some are capable of killing healthy hosts. Species in this genus are Chaetophloeus LeConte 1876, 9 spp., eight in western North
among the most economically important bark beetles. America and one in the Florida Keys and adjacent islands. All
species attack branches and twigs. Long larval mines radiate from
Bothrosternina Blandford 1896 the parental gallery, deeply engraving the xylem and phloem.
Renocis Casey 1886
Cnesinus LeConte 1868, 1 sp., C. strigicollis LeConte 1868, in south- Pseudocryphalus Swaine 1917
east United States and Mexico (an additional 100 species occur
from Mexico to Argentina). Twigs and small woody stems are Liparthrum Wollaston 1854, 2 spp., one in Arizona and one in
selected for attack. Adults bore through the bark and into the Mississippi and Indiana. These phloeophagous species are very
wood, normally reaching the pith. Larvae feed in the center of small and attack small twigs of woody plants.
twigs extending the parental gallery. Erineosinus Blackman 1920
Nemophilus Chapuis 1869 Phloeochilus Schedl 1953
Phloeotrypetus Wood 1960
Pagiocerus Eichhoff 1868, 1 sp., P. frontalis (Fabricius 1801) occurs Dacryophthous Schedl 1971
north of South America from North Carolina to Mexico (addi- Trypanophellos Bright 1982
tional species occur in South America.). This species infests large
seeds of trees and other plants, especially corn. Polygraphina Chapuis 1869

Phloeotribina Chapuis 1869 Polygraphus Erichson 1836, 3 spp. of the 60 worldwide species
occur in North America. They are phloeophagous in recently bro-
Phloeotribus Latreille 1796, 9 spp. occur north of Mexico; 2 in the ken, cut or fallen Picea.
west and 7 in the east, especially in the southeast. Adults breed in Lepisomus Kirby 1837
 Family 131. Curculionidae · 801

Spongotarsus Hagedorn 1908 Scolytodes Ferrari 1867, 1 sp., S. schwarzi (Hopkins 1902) infests
Pseudopolygraphus Seitner 1911 Ficus in south Florida. Approximately 100 spp. occur in Central
Ozophagus Eggers 1919 and South America.
Nipponopolygraphus Nobuchi 1981 Hexacolus Eichhoff 1868
Ctenophrus Chapuis 1869
Carphoborus Eichhoff 1864, 9 spp. in the 48 states and one addi- Prionosceles Blandford 1897
tional species in northern Canada and Alaska. All are Epomadius Blandford 1897
phloeophagous in small or broken branches of Pinaceae. Erinophlius Hopkins 1902
Estenoborus Reitter 1913 Hylocurosoma Eggers 1940
Hexacolinus Schedl 1963
Carphobius Blackman 1943; 1 sp., C. arizonicus Blackman 1943, in
Arizona, extends north from Central America. Two additional Micracina LeConte 1876
species occur in Central America. They are phloeophagous in small
broken branches of conifers. Pseudothysanoes Blackman 1920, 19 spp. throughout the United
States, one of which extends into Canada; most inhabit arid areas
89. Scolytini Latreille 1807 in the western states, three species are found in the east and
southeast (approximately 60 additional species occur in Central
Scolytina Latreille 1807 America). Within this genus a wide variety of hosts are attacked.
Several western species breed in the phloem of dying mistletoe
Cnemonyx Eichhoff 1868, 2 spp. of this Neotropical genus are (Phoradendron), other species occur in the twigs of hardwood trees.
found in the Florida Keys. They are phloeophagous in woody This is a very diverse genus, and several species groups were pre-
hosts. viously treated as distinct genera. Species keying out to couplet 31
Ceratolepis Chapuis 1869 in this section’s key were previously recognized as the genus
Loganius Chapuis 1869 Cryptocleptus. Species with the antennal scape short and broadly
Minulus Eggers 1912 expanded are placed in the subgenus Aphanocleptus, and those
Coptodryas Schedl 1948 with an elongate and slender antennal scape are placed in the
Coptosomus Schedl 1952 subgenus Psuedothysanoes.
Cryptocleptes Blackman 1920
Scolytus Geoffroy 1762, 20 spp. found throughout America north Chalcohyus Blackman 1943
of Mexico. Native western species are found in conifers, while Bostrichips Schedla 1951
most eastern species are in hardwoods. Three Palearctic species are Gretschkinia Sokanovskii 1959
established in North America, most notably, S. multistriatus Aphanocleptus Wood 1960
(Marsham 1802), which transmits the Dutch elm disease fungus. Cryptulocleptus Wood 1967
All are phloeophagous and construct characteristic galleries under Neoglostatus Schedl 1978
the bark.
Ekkoptogaster Herbst 1793 Stenoclyptus Blackman 1943, 1 sp. in U.S., S. sulcatus (Bruck 1936).
Coptogaster Illiger 1807 Two species in the genus, one in California and one in Mexico.
Eccoptogaster Gyllenhal 1813 The genus is closely related to Pseudothysanoes. They are
Scolytochelus Reitter 1913 phloeophagous in small branches of woody plants.
Ruguloscolytus Butovitsch 1929
Archaeoscolytus Butovitsch 1929 Thysanoes LeConte 1876, 7 spp. across the southern United States,
Spinuloscolytus Butovitsch 1929 1 species extends north to Illinois and Pennsylvania. Apparently
Tubuloscolytus Butovitsch 1929 they are xylophagous in small branches of trees.
Pygmaeoscolytus Butovitsch 1929
Pinetoscolytus Butovitsch 1929 Hylocurus Eichhoff 1872, 15 spp. north of Mexico, most of which
Confusoscolytus Tsai and Huang 1962 occur in the southeast (more than 40 additional species occur in
Central and South America). The rudis group needs further study;
Ctenophorina Chapuis 1869 Atkinson (1989) suggests the synonymy of some species. All
species are xylophagous in small branches.
Pycnarthrum Eichhoff 1878, 1 sp., P. hispidum (Ferrari 1867), in- Micracisoides Blackman 1920
fests Ficus limbs and boles in south Florida and Texas.
Nemobius Chapuis 1869 Micracisella Blackman 1928, 5 spp. in eastern and southern United
Monebius Hopkins 1914 States. These small (1.0-2.5 mm) beetles breed in the pith of
Nomebius Navas 1915 damaged, small twigs.
Pseudomicracis Blackman 1920
802 · Family 131. Curculionidae

Micracis LeConte 1868, 4 spp. in the United States, 1 extends to on Picea (Pseudips concinnus (Mannerheim 1852)) and Pinus (Pseudips
Canada, 2 are known only from Arizona. They are xylophagous mexicanus (Hopkins 1905)).
in twigs.
Dryocoetina Lindemann 1876
Cactopinina Chamberlin 1939
Dendrocranulus Schedl 1937, 3 spp. in southern and western United
Cactopinus Schwarz 1899, 5 spp. in southwestern United States, States. All species infest stems of Cucurbitaceae. The genus is
additional species occur in Mexico. The unique, paired epistomal closely related to the Old World Xylocleptes Ferrari.
male horns distinguish this genus. They are phloeophagous in
woody plants, but more commonly feed subepidermally in Cereus Lymantor Lovendal 1889, 1 sp. in eastern United States and Canada
and related cacti. and 1 species in Alaska. These beetles are phloeophagous in small,
Cactopinorus Bright 1967 dry, often dead, branches of Acer and, rarely, other hosts.

Ipina Bedel 1888 Dryocoetes Eichhoff 1864, 7 spp. in United States and Canada.
They are phloeophagous in the boles of mostly conifers, except
Pityogenes Bedel 1888, 7 spp. across the United States and Canada. D. betulae Hopkins 1915, which infests the bole of Betula.
One species, P. bidentatus (Herbst 1784), is native to the Palearctic. Anodius Motschulsky 1860
The North American species of this primarily Eurasian genus are Dryocoetinus Balachowsky 1949
phloeophagous in branches, limbs and boles of Pinus.
Eggersia Lebedev 1926 Dryoxylon Bright and Rabaglia 1999, 1 sp., D. onoharaensum
Pityoceragenes Balachowsky 1947 (Murayama 1934), native to Japan, originally described as a
Xyleborus, is established in southeastern United States. Little is
Pityokteines Fuchs 1911, 6 spp. in North America, one of which, known about the biology, but it appears to feed in the xylem
P. sparsus (LeConte 1868) occurs in the east. They often construct (Bright and Rabaglia 1999). Normark et al. (1999) discussed the
star-shaped galleries in the phloem of limbs and boles of dying genetic affinities of this genus and other Dryocoetini to Xyleborini.
trees. This genus is closely related to Orthotomicus. Various Pinaceae
serve as hosts. Coccotrypes Eichhoff 1878, 9 spp. are known from United States,
Othotomides Wood 1951 mostly Florida and California. This genus contains many species,
mostly from southeast Asia and Africa, and species found in
Orthotomicus Ferrari 1867, 1 sp. found across North America, most other areas, including the United States, have arrived through
Orthotomicus caelatus (Eichhoff 1868), is phloeophagous in Pinus, commerce (Wood 1986). Females mate with dwarfed siblings
Picea and Larix (about 10 species are known from the Palearctic). before they emerge to seek a new host. They most often infest
Neotomicus Fuchs 1911 large seeds; however, a few species are phloeophagous. Wood
(1986) stated that this genus is “in a state of taxonomic chaos”.
Ips DeGeer 1775, 23 spp. plus subspecies are currently recognized Jordal et al. (2000) showed the genetic relatedness of the genus to
from across North America. Some species placed in synonomy by Xyleborini.
Wood (1982) are recognized as valid species (Lanier 1987, Lanier Poecilips Schaufuss 1897
et al.1991). Species in this relatively large genus have been put into Cryphaloides Formanek 1908
various species groups by several workers (Hopping 1963, Lanier Thamnurgides Hopkins 1915
1970a, 1970b, 1972, Wood 1982, Cognato and Sperling 2000). Spermatoplex Hopkins 1915
Cognato and Vogler (2001) recently revised Ips as monophyletic Dendrurgus Eggers 1923
with the removal of the latidens group and their tentative place-
ment in Orthotomicus. In addition, they also named four sub- Crypturgina LeConte 1876
genera for monophyletic groups of Ips species. This well known
and important genus is phloeophagous in Pinus and Picea. Most Dolurgus Eichhoff 1868, 1 sp. is known from western North
breed in dying trees and slash, but some may attack the boles and America. Dolurgus pumilus (Mannerheim 1843) occurs from Alaska
tops of healthy trees. Characteristic egg galleries engrave the to California where it breeds in dying Picea. It utilizes the entrance
phloem-cambial area. holes of larger bark beetles, and its galleries are often wholly in
Cumatomicus Ferrari 1867 the bark.
Cyrtotomicus Ferrari 1868
Crypturgus Erichson 1836, 3 spp. occur in America north of Mexico,
Pseudips Cognato 2000, 2 spp. in North America and 1 species in one of which, C. pusillus (Gyllenhal 1813), is native to Europe
Asia. Cognato (2000) used molecular, morphological and behav- and Asia. They utilize the entrance holes of other beetles to gain
ioral characters to separate these species from Ips. The two North access to the phloem in the boles of conifers.
American species occur in the west where they are phloeophagous
 Family 131. Curculionidae · 803

Xyloterina Lindemann 1876 Anisandrus Ferrari 1867

Anaertus Duges 1887
Trypodendron Stephens 1830, 5 spp. in North America, additional Progenius Blandford 1896
species occur in Europe and Asia. These are monogamous am- Heteroborips Reitter 1913
brosia beetles that breed in either conifers or hardwoods. Xyleborips Reitter 1913
Trypodendron lineatum (Olivier 1795), which occurs across North Boroxylon Hopkins 1915
America and into northern Europe and Asia, is often a pest of Notoxyleborus Schedl 1934
conifer logs in processing yards.
Xyloterus Erichson 1836 Xylosandrus Reitter 1913, 4 spp. in eastern North America, 3 of
which are native to Asia. The three exotic species are becoming
Xyloterinus Swaine 1918, 1 sp., Xyloterinus politus (Say 1826), is very common, and occasionally aggressively attack apparently
recognized in the genus, which is found throughout eastern North healthy, small trees. Xylosandrus compactus (Eichhoff 1875) often
America. This monogamous ambrosia beetle is commonly found attacks healthy, vigorous twigs of living trees. All species cultivate
attacking weakened hardwood trees. ambrosia fungi and are consanguineously polygynous.
Apoxyleborus Wood 1980
Xyleborina LeConte, 1876
Xyleborinus Reitter 1913, 3 spp. occur in America north of Mexico,
Premnobius Eichhoff 1878, 1 sp., P. cavipennis Eichhoff 1878, from 2 are exotic. Xyleborinus saxeseni (Ratzburg 1837), native to Eu-
Africa is found in Florida. This genus is unique within the rope, is found across the United States; X. alni (Niisima), from
Xyleborini. Browne (1961) treated it as a distinct tribe, and Europe and Asia, has recently been found on the west coast of
Normark et al. (1999), using DNA, showed a separate origin North America (Mudge et al. 2001). The genus was often treated
from Xyleborini and a closer relationship to Ipini. Males of these as a subgenus or synonym of Xyleborus, but it is morphologically
ambrosia beetles are flightless and mate with siblings (consan- distinct. Their biology is similar to Xyleborus, attacking limbs and
guineous polygyny) before the females leave the brood gallery. boles of weakened trees.
Premnophilus Browne 1962
Cryphalina Lindemann 1876
Ambrosiodmus Hopkins 1915, 7 spp. occur in the eastern United
States, mostly in the southeast. They are consanguineously po- Trypophloeus Fairmaire 1868, 4 spp. in northern and western North
lygynous in a wide variety of hosts. Most attacks occur in the America. These small, less than 2 mm, beetles are monogamous
lower bole and stumps of trees. and phloeophagous in the bark of thin-barked limbs and boles
Phloeotrogus Motschulsky 1863 of Alnus, Salix and Populus.
Brownia Nunberg 1963 Glyptoderes Eichhoff 1878

Euwallacea Hopkins 1915, 1 sp., E. validus (Eichhoff 1875), native Procryphalus Hopkins 1915, 2 spp. in western North America, one
to Asia, is now established in the eastern United States. It is a additional species in Asia. Biology is similar to Trypophloeus.
consanguineously polygynous ambrosia beetle that breeds in the
stumps and boles of hardwoods and conifers. Ernoporicus Berger 1917, 1 sp., E. kanawhae Hopkins 1915, known
only from the type series taken in flight in West Virginia.
Xyleborus Eichhoff 1864, (Vandenberg et al. 2000, key to eastern Eocryphalus Kurenzov 1941
United States species); 17 spp. are identified from America north of Ernopocerus Balachowsky 1949
Mexico, 5 of which are native to Europe and Asia. Most United
States species are found in the east. More than 500 species are de- Scolytogenes Eichhoff 1878, 1 sp., S. knabi (Hopkins 1915), occurs
scribed from the neotropics, Africa and Asia. Representatives of this in vines in south Florida. Many other species are found in sub-
large and important genus attack almost all parts of woody plants. tropical and tropical areas of the world.
Most of these ambrosia beetles attack declining trees, but some may Lepicerus Eichhoff 1878
attack apparently healthy plants. Flightless, haploid males mate with Cryphalomorhpus Schaufuss 1891
sibling or parental females within the brood galleries before emer- Letznerella Reitter 1913
gence. Extreme inbreeding and partial parthenogenesis may be the Hypothenoides Hopkins 1915
cause of the many morphological races and species. In addition, this Neocryphalus Eggers 1922
mating system has allowed for new founder populations to be easily Negritus Eggers 1923
distributed through commerce (Atkinson et al. 1990). The generic Cylindrotomicus Eggers 1936
and tribal limits of these rapidly radiating species need taxonomic Lepicerinus Hinton 1936
revision. Jordal et al. (2000) and Normark et al. (1999) showed ge- Cryphalophilus Schedl 1970
netic relatedness to Dryocoetini and Wood (1986) suggests a rela- Xylocryptus Schedl 1975
tionship with Xyloterini.
804 · Family 131. Curculionidae

Hypocryphalus Hopkins 1915, 1 sp., H. mangiferae (Stebbing 1914), Pityophthorina Eichhoff 1878
native to Asia, occurs in mango, Mangifera, in south Florida. They
are phloeophagous in branches of their host. (This group has been treated as a subytribe of Corthylina by
Dacryphalus Hopkins 1915 Wood and Bright (1992).)

Cryphalus Erichson 1836, 3 spp. occur in conifers in northern and Dendroterus Blandford 1904, 2 spp. in United States, one in Texas
western North America. They are generally less than 2 mm and in Jatropha and one in California in Bursera. They are phloeophagous
infest declining branches and small trees. Adults construct cave- in the bark of declining branches.
type galleries in the phloem. Several hundred nominate species Plesiophthorus Schedl 1940
occur in Asia to Australia, and a worldwide taxonomic revision is Xylochilus Schedl 1956
needed (Wood 1986).
Pseudocryphalus Ferrari 1868 Araptus Eichhoff 1872, 1 sp., A. dentifrons Wood 1974, occurs in
Taenioglyptes Bedel 1888 south Florida (Atkinson and Peck 1994) and possibly Texas. This
Cryptarthrum Blandford 1896 Neotropical species breeds in the pith of vines. Araptus politus
Allarthrum Hagedorn 1912 (Blandford 1904) has been intercepted in large seeds in the port
Ericryphalus Hopkins 1915 of Miami, but it is not known to be established.
Piperius Hopkins 1915 Neodryocoetes Eggers 1933
Ernocryphalus Murayama 1958 Thamnophthorus Schedl 1938
Acryphalus Tsai and Li 1963 Neopityophthorus Schedl 1938
Jugocryphalus Tsai and Li 1963 Sphenoceros Schedl 1939
Hypertensus Hagedorn 1950
Cryptocarenus Eggers 1937, 2 spp. are found in south Texas and Brachydendrulus Schedl 1951
Florida and extend through Central and South America. Males Gnathocranus Schedl 1951
are flightless in these consanguineous polygynous pith borers of Gnathoborus Schedl 1970
small twigs.
Tachyderes Blackman 1943 Conophthorus Hopkins 1915, 8 spp. are currently recognized from
America north of Mexico, 2 spp. occur in the east and 6 in the
Hypothenemus Westwood 1836, 21 spp. have been recorded from west. All species breed in the cones of Pinus.
the United States, many of which are native to Asia or Africa.
Most United States species occur in the southern half of the Pityoborus Blackman 1922, 2 spp. in United States, one in south-
country. These small, less than 2 mm, beetles infest twigs, vines, east and one in southwest. They are phloeophagous in dying
pith, seeds and other plant material. They are consanguineously branches of Pinus. Their galleries in the cambium deeply score the
polygynous and have been widely distributed through commerce. xylem.
Over 200 species have been assigned to this genus, and species
identification is often difficult. Pityotrichus Wood 1962, 2 spp. in southwest United States (Ari-
Stephanoderes Eichhoff 1872 zona and New Mexico). These species are distinguished from
Homoeocryphalus Lindemann 1876 closely related Pityophthorus by the unique pregula referenced in
Triarmocerus Eichhoff 1878 the key. They are monogamous and feed in the phloem of small
Adiaeretus Hagedorn 1909 branches.
Stylotentus Schedl 1939 Pityophilus Blackman 1928
Chondronoderes Schedl 1940
Archeophalus Schedl 1941 Pseudopityophthorus Swaine 1918, 11 spp. across America north of
Pachynoderus Schedl 1941 Mexico. All species breed in branches or boles of Quercus, except
Lepiceroides Schedl 1957 P. fagi Blackman 1931 which is found in Fagus.
Ernophloeus Nunberg 1958 Xenophthorus Wood and Yin 1986
Epsips Beeson 1941
Macrocryphalus Nobuchi 1981 Pityophthorus Eichhoff 1864, 104 spp. are recognized north of
Mexico, more than 200 additional species occur in Central and
Trischidias Hopkins 1915, 5 spp. occur in the southeastern United South America and more than 50 in Europe, Asia and Africa.
States. These very small, less than 1 mm, beetles are relatively rare. This large and diverse genus is found throughout the United
One species feeds on fungus pustules under the bark of man- States in many different hosts. Representatives may be found
grove, and others are phloeophagous in injured, often fungus- breeding in twigs, seedlings, boles or pith. Most are
infested twigs. heterosanguinously polygynous and some are monogamous.
Trigonogenius Hagedorn 1912
Hagedornus Lucus 1920
 Family 131. Curculionidae · 805

Myeloborus Blackman 1928 Platypodinae are an enigmatic group that have been recognized
Gnathophorus Schedl 1935 either as a distinct family or a subfamily within Curculionidae.
Conophthocranulus Schedl 1935 There are 4 genera in North America based on the recent division
Breviophthorus Schedl 1938 of the genus Platypus into a variety of smaller genera (Wood
Pityophthoroides Blackman 1942 1993). Traditionally they have been closely allied with Scolytinae,
Cladoborus Sawamoto 1942 but Lyal (1995) could not find support for a monophyletic group
Neomips Schedl 1954 comprised only of scolytines and platypodines nor could he find
Ctenyophthorus Schedl 1955 support for them having a separate ancestry from Curculionidae.
Gnathophthorus Wood 1962 Similarly, Thompson (1992) chose to give Platypodidae family
Hypopityophthorus Bright 1981 level status while at the same time considering Scolytinae as a
subfamily within Curculionidae. A review of their phylogenetic
Corthylina LeConte 1876 position is given by Kuschel et al. (2000).
Platypodinae are easily recognized by the lack of a rostrum,
Gnathotrichus Eichhoff 1869, 7 spp. north of Mexico. G. materiarius presence of pregular sutures, pregular sclerite distinct, located
(Fitch 1858) occurs in Pinus throughout eastern North America, between median gular suture and labial articulation, at least one
the remaining species are in the west in oaks and conifers. They pair of tibiae with denticles or stout socketed setae along the
are monogamous ambrosia beetles breeding in dying or fallen dorsal (outer) margin, tarsus with article 1 as long as articles 2-5
trees or logs. Some species are pests in wood processing yards, combined, pronotum usually with a lateral constriction near the
especially in the Pacific Northwest. middle and the antennal club without sutures (Fig. 131).
Gnathotrichoides Blackman 1931 Where known, adults and larvae infest the wood of dead or
Ancyloderes Blackman 1938 recently cut or dying trees. Larvae mine galleries deep into the
Paraxyleborus Hoffman 1942 wood which become stained black by ambrosia fungi which grow
Prognathotrichus Bright 1972 on the walls of the tunnels and serve as the larval food (Bright
1993).
Monarthrum Kirsch 1866, 5 spp. north of Mexico, 2 spp. through-
out the east and 3 spp. in the west. More than 100 additional KEY TO THE NEARCTIC GENERA OF PLATYPODINAE
species are found in Central and South America. These ambrosia
beetles attack logs and boles of dying hardwoods, especially oaks. 1. Metasternum and metepisternum near hind coxa
weakly or not impressed for reception of femur,
Corthylomimus Ferrari 1867
anterior margin of impressed area not continu-
Cosmocorynus Ferrari 1867 ously carinate or with a row of small spines, sur-
Pterocyclon Eichhoff 1869 face of impressed area with at least some setae
Anchonocerus Eichhoff 1878 .................................................. Treptoplatypus
— Metasternum and metepisternum near hind coxa
Phthorius Eichhoff 1878
impressed for reception of femur, anterior margin
Trypocranus Eichhoff 1878 of impressed area either continuously carinate
Eupteroxylon Eggers 1936 or with a series of small spines, surface of im-
pressed area glabrous ...................................... 2
Corthylus Erichson 1836, 3 spp. in United States, one of which
2(1). Male with ventrite 3, 4 or 5 simple, not armed with
occurs in Canada; all are found east of the Rocky Mountains. spines .............................................. Euplatypus
Approximately 100 additional species occur in Central and South — Male with ventrite 3, 4 or 5 with a pair of widely
America. These ambrosia beetles breed in a variety of locations separated coarse spines .................................. 3
on a tree. Corthylus papulans Eichhoff 1869, in Florida, breeds in
3(2). Male with ventrite 3 with a pair of spines; female
small branches; C. punctatissimus (Zimmermann 1868) breeds in with mycetangia pores moderate in size ...........
sapling trees, especially Acer, near ground level and C. columbianus ....................................................... Myoplatypus
Hopkins 1895 breeds in the xylem of living trees, usually Acer, — Male with ventrite 4 with a pair of spines; female
with mycetangia pores unusually large in size .
which survive after the brood emerges.
....................................................... Oxoplatypus
Morizus Ferrari 1867
Pseudocorthylus Ferrari 1867
Corthylomimus Schedl 1972

XVIII. Platypodinae Shuckard 1840

by Robert S. Anderson
FIGURE 110.131. Platypodinae. 110. Myoplatypus flavicornis
(Fabricius), lateral habitus.
806 · Family 131. Curculionidae

CLASSIFICATION OF THE NEARCTIC PLATYPODINAE ANDERSON, R. S. 1988b. The Curculionidae of the Queen
Charlotte Islands, British Columbia (Coleoptera). Canadian
90. Platypodini Shuckard 1840 Journal of Zoology, 66: 2406-2414.
ANDERSON, R. S. 1989a. A revision of the subfamily
Treptoplatypus Schedl 1972, 2 spp., T. abietis (Wood 1958) and T. RhynchaeninaeintheNewWorld(Coleoptera:Curculionidae).
wilsoni (Swaine 1916), northwestern United States and British Transactions of the American Entomological Society, 115:
Columbia. Species are associated with timber of conifers (Bright 207-312.
1993). ANDERSON, R. S. 1989b. New synonymy in North American
Stephanocleonus Motschulsky. Coleopterists Bulletin, 43: 93.
Myoplatypus Wood 1993, 1 sp., M. flavicornis (Fabricius 1776), south- ANDERSON, R. S. 1991. A new species of Plocetes from the
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UnitedStates(Curculionidae;Curculioninae;Tychiini).Florida
Oxoplatypus Wood 1993, 1 sp., O. quadridentatus (Olivier 1795), Entomologist, 74: 105-110.
southeastern United States. This species is associated with vari- ANDERSON, R. S. 1993a. The Curculionoidea of southern
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[excluding Curculionidae; Scolytinae, Platypodinae]). Insecta
Euplatypus Wood 1993, 3 spp., E. parallelus (Fabricius 1801), E. Mundi, 6: 193-248.
compositus (Say 1824) and E. pini (Hopkins 1905), southern United ANDERSON, R. S. 1993b. Weevils and plants: Phylogenetic
States; one species adventive. versus ecological mediation of evolution of host plant asso-
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