ter is the longitudinal groove on the prosternum
that receives the 3-segmented beak (Fig. l25A).
I I
Figure 133 A, a minute pirate bug, Orius insidiosus
(Anthocoridae); B, bed bug, Cimex lectularius
(Cimicidae); C and D, damsel bugs (Nabidae): C'
Nabicula subcoleoptralc,' D, common damsel bug,
Nabis americoferus. ',
Common Species
Nabicula subcoleoptrard (Kirby) (Fig. l33C)-7'
9 mm; shiny black; minute wings, usually fully
winged form rare; resembles ant; antennae, beak'
legs, and margins of abdomen yellow; easlern l/Z
U.S. south to KS and NJ, southern Canada.
Nqbis americoferus Carayon (Fig. l33D). Common
damsel bug-6-9 mm; dull gray or gray-yellow; head
and pronotum with dark stripe; throughout U'S' N'
alternatus Parshley, western damsel bug, similar,
wings speckled brown, reddish brown spots on hind
tibiae, west of Mississippi R.
Family Reduviidae-
Assassin Bugs
Members of this family are small to large bugs
that range from stout to extremely sr .lder and
elongate. The important identificatio t charac-
The first segment of the beak forms a rounded,
downward curve helping to give the head an
elongated appearance. The abdomen of some
species is wider than the wings and flattened
dorsally. Assassin bugs are predators of other
insects and many will bite humans. Several
southwestern species in the genus Triatoma
transmit trypanosome (protozoan) diseases to
rodents and man (Chagas disease in humans).
Common Species
Apiomerus crassipes (Fabricius) (FiC. l34A) 12-
l9 mm; black; pronotum, scutellum, and abdomen
broad and with dark red fringe; head, thorax, and
legs with dense hairs; most of North America l'spe-
cially West).
Arilus cristalus (Linnaeus). Wheel bug-28-36 mm;
dark brown with thick grayish pubescence; prono-
tum with semi-circular crest containing 8-12 stout
tubercles; NY to IL and south to FL.
Emesayø brevipennis (Say) (Fig. l348)-33'37 mm¡
extremely slender and long-legged; grayish or silvery
covering of fine hairs; front tarsi 3-segmented; most
of U.S. (in barns, old buildings).
Mela no les tes p icipes (Herrich-Schäffer)-l 5-20 mm;
black; apical l/4 of tibiae
irregularly dilated; front
wings often very short ); eastern l/2 U.S., south'
(19
west to C^. M. abdominalis (Herrich-Schäffer),
similar, red abdomen
Rasahus biguttatus (Say)-16-20 mm; dark reddish
brown; most of wing dark yellow with large yellow
spot in middle of black membrane; southern U.S.
west to CA, sometimes Midwest. R. thoracicusStäl,
similar, amber, amber spot in middle of black wing
membrane, Far West.
Reduvius personatus (Linnaeus) (h-ig. ¡34C).
Masked hunter-17-23 mm; dark brown to blackish;
promincnt vertical hairs; eastern U.S. west co KS
(often at lights, in houses; immatures thickly cov'
èred with dust¡.
Repipta /øurus (Fabricius) (Fig. l34D)-ll-13 mm;
slender; orange-red or yellow; 2 spines on head; pos'
terior L/2 pronotum with 4 longitudirral stripes, 2
long black spines; PA to FL, west to TX.
Order Hernipt,:rra 145
 e
o
(Ð

ÇJ

Ê!.

Figure134 Assassin bugs (Reduviidae). A, Apiomerus crassipes; B, Emesaya brevipennis;

C, masked hunter, Redvvius p.'.tonatus; D, Repipta taurus; E, spined assassin bug, Sinea
diadema; F, bloodsucking conen, tse, Trialoma sanguisuga.

Sinea dtadenra (Fabricius) (Fig. l34E). ìpined as- Family Phymatidae-

sassin bug-12-14 nrnr; dark brou n to drrll reddish
brown; pronotum with prominent sharp spines; each
abdominal segment with a lateral spot; nl()st oi U.S
S. spinipes (Fabricius), similar, pronoturn with tu-
bercles but not spines, east of Rocky Mts.
Triatoma sanguisuga (Leconte) (Fig. l3aF). Blood-
sucking conenose-16-21 mm; mixed reddish and
dark brown to blackish; corium with reddish yellow
area near base and apex; lateral areas of broad ab-
domen with alternating red and black patches; FL to
TX, rrorth to NJ, IL, KS (often indoors;severe bite).
T. protracta (Uhler), western bloodsucking con-
enose, smaller, black, UT, AZ, CA.
Ambush Bugs
These stout-bodied bugs are i3 mm or less in
lengtl'r, the raptorial fro¡lt femora are greatly
enlarged, and the abdomen distirrctly úìclens
posteriorly. Ambush bugs are commonly yel-
low or greenish yellow with brown or black
markings and individuals often \rýait on similar-
ly colored flowers (e.g., goldenrod) for an in-
sect to land. The bug then quickly seizes the
prey (typically a fly, bee, or wasp) and sucks
out its blood. Phymatids do not bite humans.

146 Subclass Pterygota

 Common Species
Corythucha ciliata (Say) (Fie. l36A). Sycamore laci
bug-4 mm; uniformly milky white except for dark
mark on thorax and center of each wing; east of
Rocky Mts. (on sycamore, ash, hickory, mulberry).

Corvthucha pruni Osborn and Drake-4 mm; whit-

¡rrr litrr uiownish black crossbar near front wing
apex; pronotal hood low; eastern l/2U '5., UT' OR
Figure 135 An ambush bul, Phymala pennsylva- (on wild cherry).
nlca (Phymatidae). fm, femur.

Common Species
Phymata fasciata (Gray)--9-12 mm; pale yellow to
greenish yellow; scattered dark markings; wide
brown band across middle of abdomen;most of U 'S'

Phymata pennsylvanicø Handlirsch (Fig' 135)-

8.5-9.5 mm; yellow with brown or black areas,
sometimes greenish when alive; front wing mem-
brane brown; 4th antennal segment longer than 2nd
and 3rd combined (cr); 4th abdominal segment
abruptly expanded (cr); eastern U.S. P. americano
Melin, similar, 4th antennal segment shorter than
2nd and 3rd segments combined (o' abdomen
evenly expanded to 4th scgment (g), 'hroughout
U.S.

Family Tingidae- Lace, Bugs

Lace bugs are small (usually lcss than 6 mm
long) insects with the front wings and expanded
thorax sculptured into a lacelike ¡rattern of
cells. Most adults are grayish white and the
nymphs are blackish. Lace bugs are plant feed'
ers and often found on leaves of trees and
shrubs; they tend to be host specific. One spe-
cies is used in control of a weed in certain parts
of the world. The largest and most common ge-
nus in North America is Corytftuclra which is
characterized by a prominent pronotal hood
that extends over the head and greatly ex- Figure136 A and B, lace bugs (Tingidae): A, syc'
panded, lacy, pronotaì margins that bear small, amore lace bug, Corythucha ciliatu; B, Melon'
sharp spines (Fie. l36A). orhopala clavata. C, a piesmatid or ash-gray leaf
bug, Piesma cinereo (Piesmatidae); D, a flat bugr -':
(Aradidae)
A radus øcøf¡¡s

Order Hemlptera
 Family Piesmatidae-
Piesmatids or Ash.Gray
Leaf Bugs
Members of this family are small (about
4 mm long) and the cell network in the front
wings resembles that of certain lace bugs
(Tingidae), The pronotum has five longitudinal
ridges. These insects feed on leaves of trees and
weeds. All 7 species are in the genus Diesma.
C<lmlnon S¡recics
l)¡e\ntu t'incre¿t (Sar ) (Fi_q. l.-16( ) - 1.5,:t ,s m;.n; cJuli
gra]'or ¡cllori;--ìrcl antcnnal scLtrì1enI i,.lìgrh = to
lst and 2nd cttntbined; most ol U.S. (t,n pigx,cecl,
other rvccds).
Family Aradidae- Flat Bugs
These insects are generally 3-l
I mm long,
flat, and brown to black in color. There are no
ocelli and the wings do not cover the entire ab_
domen. Flat bugs occur on or under loose bark
ol dead rrees and feed on the liquids of fungi
and decaying wood (rarely living wood).
Common Species
Aradus acutus Say (Fig. l3óD)-7_10 mm; blackish
brovvn; 2nd antennal segment clubbed; white spot on
side ol each abclominal segment; most of U.S. (on
oak),
Aradus robustus Uhler-5-7 mm; brown to gray;
head and pronotum same length; 2nd anrennaiseg_
ment thick a¡rd covered with short stout spines; east-
ern U.S. (on red and black oak).
Neuroctenus simplex (Uhler)-.1.5_5.0 mm; dark
reddish brown or blackish; tip of 4th anrennal seg_
ment yellow; tarsi yellow; lront wing membrane
whitish; narrow transverse ridge behind anterior
marg.in ol abdominai segments 4_6; eastern U.S. (on
oak, beech, u,eeds).
Family M iridae- plant Bugs
These very common bugs are generally less than
l0 mm long and the front wing has a cuneus
and one or two closed cells in the membrane
(Fig, l268), Ocelliare absenr. Most species feed
148 Subclass pterygota
on plant juices and many are serious pests (e.g.,
lygus bugs in the genus lygus). Some species
are predaceous on small insects. Fleahoppers
2- (genus Hølticus) are jumping mirids with en-
larged hind femora and short, evenly thickened
front wings (Fie. l37A). Numerous wingless
species are remarkable mimics of ants. This
family contains about 1,700 species in North .
America and is the largest in the suborder Het-
eroptera. Keys to genera arc given in Slater and í
Baranowski (1978).
Colnnrr¡n Species
Ådelphot'oris rapidus (Say). Rapicl plant bug_7- 6l
8 mnr; dark brown; 2 dark spots on póstelior t72 of
pronorum; sides of front wing dull yellow_white; ql
eastern U.S. ,4. lineolatus (Coeze), alfalfa plant bug,
8-9 mm, green or greenish yellow, faint cl,ouded ai_
very eas on front wings, on alfalfa and sweet clover, ü{
throughour U.S. ¡tl
Halticus bractatus (Say) (Fie. l37A). Garden
fleahopper-l.S-2.5 mm; btack; lst aniennal seg-
menJ and parr of 2nd and 3rd olten pale; front winls
t
of .9 .short, thickened, with or *it'nouí *"*Urunr,
with silvery flat hairs; fronr wings of cr normal
length; hind legs enlarged; jumps;iastern l/2V,5.
(often on white clover in lawns).
t.
Ilnacora molina (Uhler)-S.5 mm; black; base of
pronotum and front wings deep green; eastern and
t
central U.S. (on goldenrod in damp areas). (
Leptopterna dolabraÍa (Linnaeus) (Fig. t378). (
Meadow plant bug-7-9 mm; pale greeñish whiie
with 2 broad longitudinal bãnds ãn pronotum
(cr, Q) or reddish brown to orangish yejlow (c'); I
wings often short ( Q ); northern l/2U.5.
(_
Lygus
.lineolarr (palisot de Beauvois) (Fig. l37C), (
Tarnished planr bug-5-6 mm; yellowish'brãwn with
some dull reddish, brown, and black mottling; y_
¡haped yellow marking on scutellum; easterñ l,/2 (
U.S.
t'--
(
Plagiognathus politus Uhler-3.5-4.0 mm; black
with whitish hairs; tibiae and tips of femora pale;
later in summer not as dark, yellowish shade on tho-
rax; eastern U.S. (on ragweed, gold"nrod, other
weeds).
Potcilocu¡tstts li¡tt'¡ttuç (Fabricius) I ig. llTE)
irourlinccj plant trug-7,0-7.,s nrrnl gt3c ì or vcllo*'-
ish; heacl nray be ¡cllo*'ish red;.1 broad, black longi-
tudrnal stripes on pronotum and I ront $'ings; eastern
l/2 U.S., soulhern Canada.
Pseudatotnoscelis serialus (Reuter). Cotton
lleahopper-3-4 mm; pale yellowrsh green; covered
with brownish black specks and minute black hairs;
South and Southwest (on cotton, other plants).
Slaterocoris stygicus (Say)-4.5 mm; shiny black;
broadly oval; eastern l,/2 North America, MT, CA
(on goldenrod).
Slenodema vicinum (Provancher) (Fig. l37F)-
7-8 rnrn: dull yellorv or stra\v cc'lor wilh greenish
toncs; ilr'ìtL'nnae reddish, l st anter.na. segment thick
and h;¡trr,; pronotum with.l-l dirrk longitudinal
stri¡res; rnost of U.S
.stenotus ltinotatus (Fabricius)-6 rnm; yell wish
orange; pronotum and front wiltgs with 2 broad
black stripes sometimes changing to spots on poste-
rtor l/2 pronotum; northern and central U'S'

Figure 137 Plant bugs (Miridae). A, garden fleahopper, Halticus bractatus B, meadow
plant bug, Leptopterna dolabrata; C, tarnished plant bug, Lygus lineolaris,'D, clouded plant
bug, Neurocolpus nubilus; E, fourlined plant bug, Poecilocapsus lineatus; F, Stenodema
vtctnum.

Order Hemiptera
 (".,

Ìì'@
Family Lygaeidae-Seed Bugs species have more than five veins in the front
Seed bugs are brownish or sometimes brightly wing membrane. Most lygaeids feed on mature Ð
seeds, typically injecting saliva and sucking out
colored insects that vary from 2-18 mm in
length. They are recognized by the 4-segmented
beak and antennae, the presence of ocelli, and
the dissolved contents of the seed. Some species
are sap feeders (e.g., chinch bug) or predators
t
four or five distinct veins in the membrane of (e.g., Geocorrs species prey on other insects). Ð
Various species are agricultural pests. Seed
the front wing. Some species resemble the Mir-
idae but the latter have a cuneus and no ocelli.
Lygaeids may resemble certain species of Aly-
bugs, with over 250 species in North America,
are the second largest family in the suboider
.6
didae, Coreidae, or Rhopalidae, but the latter Heteroptera, $,
u
6,
EJ

çÐ

t}
L
ç::

s\ort-winged form
,,ü
'tü
U
('

t't

Figure 138 Seed br,gs (Lygaeidae). A, chinch bug, B/rssrzs leucopterus; B, Geocoris
punctipes;C,small milkweedbu¡ Lygaeuskalmii; D,Nyslus niger;E,largemilkweedbug,
Oncopeltus Íascialus, F, Ortholont s scolopa-r.

150 Subclass Pterygota

Common Species
B/lssus leucopterus (Say) (Fie. l38A). Chinch bug-
3-4 mm; dull black and white with thick gray pubes-
cence; winged and wingless forms; lront wings white
with large black spot on corium; Midwest (in corn
and grain fields).
Geocoris punctipes (Say) (Fie. l38B)-3-4 mm;
gravish or dull )'ellowish; top ol head smooth and
shiny, rt ith a ¡niddorsal grove; scutellunl with 2 pale
stnpcs; strulhern 2,/3 U.S. G. ltullctlus (Say), large
bigcyed bLrg, sinrìlar, top oi head roughened, most
ol U.S G. pallens Stål, western bigeyed bug, pale
yellowish green, small black spots on head and tho-
rax, scutellum black and without a middorsal ridge,
western I /2 North America.
Ligyrocoris diffusus Uhler-5-7 mm; head and front
of pronotum gray or black; front wings dull reddish
brown; posterior part of pronotum with 4 faint
darker stripes; pale spot near inner corner of cori-
um; most ol U.S. except Southeast (on composit
flowers).
Lygaeus kalmii SLàl (Fig. l38C). Small milkweed
bug-ll-13 mm; red and black; red spot at base of
head; irregular red "X" formed by wings; iront
wing membrane with 2 white spots or none (subspe-
cies in eastern 1/2 U.S.) or I large white spot (sub-
species in western I /2 U.S.); on milkweed.
Melanopleurus belfragei Stål-10 mm; head, prono-
tum, scutellum and front wing membrane black;
clavus and corium orange-red; TX to CA.
Myodocha serripes Olivier-8.0-9.5 mm; head shiny
black, extended posteriòrly into a distinct long neck;
pronotum gray; wings dark brown, margin pale; Iegs
pale yellow, dark bands on femora; eastern l/2
U.S., southeastern Canada.
Neacoryphus bicrucis (Say)-7.0-9.5 mm; body
black above; posterior part of pronotum and corium
bright red; southern U.S.
Nysius r¡ger Baker (Fig. l38D)-3-4 mm; dull
yellow-gray; 2 irregular longitudinal rows of black
spots on corium; membrane clear; most of North
America.
Oncopeltus fasciutus (Dallas) (Fig. I r 3E). l-arge
milk*'eed bug--10-l8 mm;dark tr<)wn t r bla:k; top
ol hcail, sides ot'l)ronotum, and .l broati crossbancls
on ll'ings are orangc; east of Ror:ky N{r. (olr milk-
weed).
Ortholomus scolopax (Say) (Fig. l38F)-5-6 mm; '
dull gray-brown; tip of corium dull reddish; short
silvery or gray hairs on body; most of U.S., southern
Canada.
Paromius longulus (Dallas)-6-7 mm; pale tan; head
projecting prominently; southern U.S.
Family Berytidae-St¡¡t Bugs
These slender bugs are 3-10 mm long and have
unusually long legs and antennae. The first an-
tennal segment is much longer than the Jthers.
Stilt bugs resemble the shape of Hydrometridae
and certain Reduviidae; however, stilt bugs do
not occur on water or have thickened front legs,
and the beak is 4-segmented. Berytids feed on
plants (a few may be partly predaceous on other
insects) and occur on dense vegetation or on the
ground.
Common Species
Jalysus spinosus (Say)(Fig. l39A). Spi¡red stilt
bug-6-9 mm; dull reddish brown or yellowish
brown; scutellum with angled spine; eastern l/2
U.S.
Family Largidae- Largid Bugs
These medium- to large-sized bugs are often red
and black. They have a 4-segmented beak, no
ocelli, and seven or eight branched veins plus
two large cells in the front wing membrane.
Some have short wings and resemble alrùs. Lar-
gids are plant feeders and generally southern in
distribution.
Common Species
Largus succinctus (Linnaeus) (Fig' l39B)-13'
17 mm; black or dark yellowish brown with short
gray hairs; orange-red or orange-yellow pronotal,
wing, and abdominal margins, and also base of
femora; southern U.S. west to CO anrJ AZ, north to
NY.
Order Hemiptera 151 1 :
 u
of pronotum with dark
femora red; posterior area
Ø
brown patch; front wings dark brown with dull
yellow margins; FL and southeastern U.S. t
e
Family Coreidae-Coreid Bugs
Members of this family are 7-40 mm long, @
dark-colored, and elongate but thick-bodied in-
sects. The front wing membrane contains many LJ
veins. Scent gland openings are present on the
thorax between the middle and hind coxae
(Fig. l28A). Because some species have dilated,
leaflike areas on the antennae and/or the hind
u
legs, the family name leaffooted bugs has been @
used. They are also known as the squash bug
family. Most species feed on plants and some
are agricultural pests.
c
@

,-,
Figure 139 A, spined stilt bug, Jal_ysus spinosus
(Berytidae); B, a largid bug, Largus succincÍus (Lar-
gidae); C, cotton stainer, Dysdercus tuturellus (Pyr-
\J
rhocoridae).

t
Family Pyrrhocoridae-
Red Bugs and Stainers
Pyrrhocorids are medium- to large-sized bugs
that are olten marked with red, yellow, brown,
and white. Many pyrrhocorids resemble the
largid bugs br¡t the lormer's body lacks hairs,
the beak extends to the abdomen, and the sixth
sternite ol the female is not split. AII seven (
North American species are in the genus Dys-
dercus and occur in the southernmost states. 1'

Common Species {
Dysdercus suÍurellus (Herrich-Schäf fer) (Fig. l39C). Figure 140 Coreid bugs (Coreidae). A, Acsn-
Cotton stainer-ll-17 mm; head, base of lst anten- thocephala femorata; B, squash bug,'Anasa tristis; f
nal segment, front ol pronotum, scutellum and C, Lept o gl oss us oppos i tus.

1.,,

152 Subclass Pterygota

Common Species
Acanthocephala femorata (Fabricius) (Fig'
25-28 mm; reddish brown or yellowish brown; thick
yellowish hairs; small tubercles on pronotum; hind
femora greatty thickened and curved (cr), leaflike
expansions of hind tibiae nearly reach tip of tibiae;
southern U.S.
Acanthocephala terminalis (Dallas)-20-22 mm;
brown; lst 3 antennal segments dark brown or dark
reddish brown in contrast to orange-yellow 4th seg-
ment; Ieaflike expansions of hind tibiae reach about
2/3 length of tibiae; eastern l/2V.5'
Anasa tristis (De Geer) (Fie' laOB). Squash bug-
l3-18 mm; dull brownish yellow; body with dark
gray punctures; head with 3 yellow stripes; sides of
each abdominal segment partly yellow; most of
North America (on squash, cucumber, pumpkin)'
Common Species
la0A)- Arhyssus lateralis (Say)-5-7 mm; color variable¡
reddish brown to dull yellow; head and pronotum
red, pale yellowish brown, or combination of both;
abdomen commonly yellow; most of U.S., southern
Canada.
Leptocoris trivittatus (Say) (Fig. l4lA)' Boxelder l:'
bug-11-14 mm; black; dark reddish pink stripe .
down center of pronotum and edges of prot- ttum
and corium; most of U.S. (on boxelders, congregates
in masses in fall).

Euthochtha galeator (Fabricius)-14'5-17.0 mm;

dull brown; stout-bodied like,4. /rtsf,s,'4th antennal
segment dark brown; antennalike tubercles on body
with a spine on their outer sides; hind "emora with
large teeth or spines ventrally; castern U j.

Leploglossus o¡tprt.sittts (Sal ) (Fig. 1'10('')- I 8-

20 rnrn; brou'n rt hil I bands on

ithout distillct rr
wings; leallike expansion on hind tibiae ras inclenta-
tion on outer margin; eastern U S north 'vest to MN,
southwesl ro A'2. L. occidenlahs Heidemann, simi-
lar, hind tibia expansion elongate ancl uith inclenta-
tions, western 2/3 U.S.

Leptoglossus phytlopus (Linnaeqs). Leaffooted

bug-18-20 mm; brown with distinct white cross-
band on front wing; leaflike expansion on hind
tibiae; southeastern U'S. north to NY and MO ¿'
clypea!is Heidemann, similar, spine projects upward
at tip of hcad, western 2/3 U.S.

Figure l4l A, boxelder bug, Leptocoris tr.iÿittatus

Family Rhopalidae-
Rhopalid Bugs
Rhopalids resemble small species of Coreidae
but either have very small lips around the scent
gland openings or lack openings. The ocelli are
distinctly raised. Rhopalids are
called scentless plant bugs and are classified
Corizidae in some texts.
(R-lropalidae); B and C, alvdid bues (Alydidae): B'
Alydus eurinus; C, Protenor belfragei.
Family Alydidae-AlYdid Bugs
sometimes These yèllowish brown or black bugs resemble
as the Coreidae but have broad, prominent heads
and more slender bodies. They often Droduce
strong odors from thoracic scent glands. Aly-

Fiv'
Order Hemiptera 153i:ìil'

dids are sometimcs called broadheacl :d bugs
and are classilicd as Coriscid¿tc in s<lrlte texts.
Nymphs of some spccies (genus '4 lydus) resem-
ble ants; a lew adults mimic wasps. Alydids are
plant feeders and occur on foliage and flowers
including grasses,
of, weeds
Common Species
Alydus eurinus (Say) (Fig. l4lB)-ll-15 mm;
blackish; sides of each abdominal segment with
small yellow spot; throughout U.S. ,4 pilosulus
Herrich-Schäfl'er, sinrilar, light brown, pronotal
margins white, anterior corners of pronotum sharp,
throughout U.S. but chiefly southern.
Protenor bclfragei Haglund (Fig. lalC)-12-15 mm;
shiny brown; slender and parallel sided; antennae
Figure 142 Stink bugs (Pentatomidae).
B, Say stink bug, Chlorochroa sayi;
D, Homaemus bijugis; E, Oebalus pugnax; F, spined soldier bug, Pod¡sus maculiventris.
154 Subclass Pterygota
{,1
€t
and rrftcn Lrcak and legs yellorv-r,:C; body wjth dark
puncrurcs; n(lrthern u.S. lronl \{D tc Fi.()cky Mts. Êr
Family Pentatomidae--S tink Elugs
0
qD
Stink bugs are comrnon, broad, shield-shaped
bugs with 5-segmented antennae. 'lhr: large scu-
tellum is distinct and in the subfantily Scutel-
.0
lerinae (shield bugs) it extends to the tip of the
abdomen (Fig. la2D). The shield bugs are con-
sidered a separate family, Scutelleridae, in
some texts. Most stink bugs are plant feeders, U
some are predators, and others leed on both
plants and animals. Pentatomids produce odors t
from thoracic glands for self-defense.
€Ð
@
q¡
€
A, rough stink bug, Brochymena quadripustulata;
C, onespot stink bug, Euschistus variolarius;
Common Species lel stripes on dorsum; west of lvlississippi R.' tL. H.
parvulus Cermar, similar, 4-6 nrm, paler "ellow,
Acroslernutn hilare (Say). Creen stink southern | /2 U .S .
bug-12.0-17.5 mm; bright green; elongate body;
head and pronotum with narrow yellow, orange or
Mormidea tugens (Fabricius)-6-7 mm; dull bronze;
reddish border; Iateral pronotal margins narrow an-
scutellum with whitish margins; east of Rocky Mts.
teriorly in straight line rather than curving; most of
U.S.
Murgantia histrionica (Hahn). Harlequin bug-
Broch¡'menu quadripustulctta (Fabricius) l0-li mm; variegated orange and black markings;
chiefly southern U.S. (on cabbage, other crucifers).
(Fig. la2A), Rough stink bug-14-18 mm; grayish
or ãark reddish brown; body marked rvith dark and (Fa mm;
Oebalus pugnax
light areas and resembles tree bark; anterior corners dull yellow; elongat Pro-
of pronotum bluntly pointed; throughout U.S' B' ject iorward; east o thern
suliata Van Duzee, similar, southwestern U.S.' CA (on rice, wheat, sorg :
(in orchards).
Podisus maculivenlris (Say) (Fig. l42F). Spined
Chlorochroa sayi Stål (Fig. l42B)-Say stink soldier bug-6-8 mm; brown; often pink toward tip
bug-15-17 mm; bright green; white spots or tiny of corium; prothorax pointed laterally; base of ab-
raised points on dorsum, 3 white or pale orange domen with spine projecting between hind coxae; 2
spots across base of scutellum; western 2/3 U'S' dark spots near tip of front wing membranes; most
C. tigala (Say), Conchuela, sinrilar, often dark olive of U.S. (predaceous).
greeñ, lacks 3 white or pale orange spots at base of
scutellum. Thyanta accerua McAtee. Redshouldered stink
Cosmopepla himaculqla (Thomas)-5-7 mm; shiny t,ug-S-tO mm; pale green; pronotum with anterior
.oin.r, pointed and lateral rnargins narrowly edged
black; .vellowrsh or bright red stripes along lateral
and anlerior margins of pronotutn and middorsum with dull red; east of RockY Mts.
of pronotum (anterior I /2); tip of scutellum with red
dot on each side; most of U.S
Family Thyreocor¡dae-
Euschisrus variolarius (Palisot de Beauvois) Negro Bugs
(Fig. la2C). Onespot stink bug-ll-i4 mm; hind
tibiae with distinct groove for complete length;edges
These shiny black, broadly oval bugs ;.re gen'
of anterior abdominal sternities without spots; large erally 3-6 mm long. The huge scutellum covers
black spot on genitat capsule at tip of abdomen (d ); the abdomen giving the appearance of small
chiefly northern l/2 tJ.S. E. servus (Say), brown beetles. The front tibiae are not flattened for
stink bug, similar, abdomen with black spots on ven- digging as are those of the similar burrower
tral sidJnear margins but lacks row ol midventral
black spots, most ol U.S., especially South.
Uugì (CvOnidae). Negro bugs occur in weedy
fields. The name Corimelaenidae is used for thei'
Holcostethus timbolarius Stàl-7-8 mm; dark gray- family in some texts.
ish yetlow; dense dark punctures; lateral margin of
pronotum, basal l/2 of lateral margin of corium, Common Species
änd tip of scutellum are whitish; lateral margin of
black àbdomen uniformly pale yellow; most of U'S' Common genera: wi:h groove
along lateral mar s black) and
H. abbreviatus Uhler, similar, alternating light and (no colored lat-
Corimelaena
dark patches on lateral margins of abdomen, west-
ern 1/2 U.S. eral margin).

Corimelaena Negro bug-;rl t

Homaemus bijugis Uhler (Fig. l42D)-ó-8 mm;con- of corium with
vex scutellum nearly reaches tip of abdomen; light to 2.5-3.5 mm;
pale stripe tiP of corium
dark yellowish tan; head black, lateral margins red-
rounded; mo nada.
dish tan; black punctures form diverging and paral-

Order Hemiptera 1
Oalgupha atra Am1-ot and Servillc (Fig. la3A)-
4 5-6.0 mn-r; shiny black; scutellurn cvenll' artd
broadly ror¡rrdcrl ¡rostcriorly; nrost ol' U.S
Family Cydnidae- Burrower Bugs
Members of this lamily are usually 7 mm or less
in length, black or reddish brown, and resemble
small stink bugs (Pentatomidae). Cydnids dif-
fer from stink bugs by having strongly spined
tibiae. These bugs burrow into soil by using
their front legs and occur around plant roots
and under boards and rocks. Burrower bugs are
most likely to be encountered at lights.
Common Species
Pangaeus bilineatus (Say) (Fig. l438)-6-8 mm;
black or reddish black; pronotum with groove paral-
leling anterior margin to form a collar; corium and
clavus with minute cracks; eastern U.S. southwest to
cA.
Sehirus cinctus (Palisot de Beauvois)-4-7 mm;
shiny black, bluish black or dark reddish brown; lat-
eral margins of body with white lir.e; throughout
U.S., southern Canada (on mints, nettles).
Iìigurc 1.13 ,,\, a n,:l¡ro hug, Gart:t,¡tltu nt o (Tb,y-
rcocoritlae); iJ, a blrrrouer bug, i)urtgui'i; htlitt-
eala.s (Cydnidae),
156 Subclass Pterygota
SUtsORDER HOMOPTERA23
e
The lront wings of the Homoptera are uniform-
ly clear and membranous (Figs. 147; 153) or
qt
colored and slightly thickened (Fig. 149). They
are commonly held at an angle (rootlike) oVer {t
the body when the insect is resting. The ventral (Ð
beak originates from the posterior part of the
head (Figs. 1444, C). Homopterans feed by
sucking plant sap. Numerous species injure
b
crops by feeding and many transmit plant dis- (.rl
eases. A few species are used as sources of dyes,
shellac, and waxes. The Homoptera are divided ü
into two groups (suborders of some authors):
the Auchenorrhyncha (active fliers and jump- €r
ers) includes the cicadas, treehoppers, Ieaf-
hoppers, froghoppers or spittlebugs, and about üt
I I families of planthoppers; the Sternor-
rhyncha (most of which are relatively inactive) eÅ
include the psyllids or jumping plant lice,
whiteflies, three or four aphid families, and l5 f,
or I 6 families of scale insects in the U S.
.
Most homopterans are collected by
sweeping vegetation. Trees and shrubs may be
shaken or beaten followed by sweeping under-
neath the plant. The entire contents of the net {- j
may be killed and sorted later or individual
specimens may be collected from the net with
an aspirator. Less active or inactive forms are
trapped in a killing jar or scraped or picked off
the host. Cicadas in trees may be collected by
using a longhandled net or dislodged to a lower
branch by using a long pole or throwing a rock.
Homoptera are generally pinned or
mounted on points. Soft-boclied irnmatures,
aphicìs, and sometimes psyllids and whjteflies
are prescrveci in 8090 alcohol or nrr¡unled on
slìdes. ilost plants with scalc:s rnrry be: coller:ted,
drieci, ancl pinnecl, or thr irsecl.r; ma) be
scrapecl oi'l' and placed in alcchcl Vlale scales
are obiained primarily by rearing irnmatures.
To identify rnost scale insects to farnilies and
species, females must be mounted ,ln micro-
23. Homoptera: homo, same or vnilorm; pteto,
wlngs.
 Antennae arising from sides of head
scope slides after clearing in potassium hydrox-
beneath eyes (Fig. l44C).. . . . . . . . I
ide and staining

3a(2a') Three ocelli present (Fig. l44B)i.

KEY TO FAMILIES
OF SUBORDER HOMOPTERA
This key is modified lrom that in Borror,
Delong, and Triplehorn (1976). Scale insects
are keyed only to superfamily; the paper by
Howell and Williams (1976) should be con-
sulted for a key to the families.
1a Antennae very short and bristlelike;
tarsi 3-segmented ''
front femora much larger than mid'
dle femora; membranous front
wings; large insects
. . . . . (p. 760't Cicadidae
i
Two (rarely 3) ocelli present; front
and middle femora similar in size;
f ront wings sometimes thickened;
small insects ......4
..., "2
4a(3b) Pronotum extends backward over
abdomen and may form sPines or .

lb Antennae usually long and filiform; other processes (Fig. 148). .

tarsi l- or2-segmented . . .' .' .' . 16
. . . . . (p. 16l) Membrscìdae l.i:.,
ir ¡:'
4b Pronotum does not extend bael:ward :'ili'
2a(1a) Antennae arising from front of head over abdomen (Fig. 149r. .
between eyes (Fig. l44B) or in front
ofeYes(Fig.144A) """'3 '!i
5a(4b) Hind tibiae wilh I or nlore rows of
small spines d<¡rln its 'entire length
(l"is. l45A). . . . (p. l'5.\\ Ciculellidae

Hind tibiae with I or 2 large sPines

and usuallY several at apex
([ig. 1458). . . . . (p. 162') Cerîepídae " .

6a(2b) Hind tibiae with a large, rnovable

spur at aPex (Fig. l45C)
. . . . (p. 164) DelPhacídae

6b Hind tibiae without large, movable

an- spur at apex .

Figure 144 Beak and antennae positions in sub-

orãer Homoptera. A, antennae arise in front of eyes;
B, antennae arise between eyes; C, antennae arise
beneath eyes. an, antenna; bk, beak.

Order HemiPtera
 ü

9a(8a) Front wings longer than body, at rest tî,0

held almost vertically at sides of
body, triangular anal area (clavus) at
,
ii,e
least partly granular, mânÿ cross.
:..O
I

veins in costal area (Fig. l51D); often

mothlike in form. . . (p. f 66) Flatídae ':'O
Front wings variable in length and
.,@
position, triangular alral ¿rre¡r (,cl:rvus) íl'
not granular, usually rvithoul nu-
merous crossveills in c'ostal area; q,
formvariable.... ......10
@
l0a(9b) Apex of front wings more densely
veined and set off from rest of wing
by crossveins; Florida to lÝlississippi
....Tropiduchidøe

l0b Apex of front wings not es described

Figure 145 Hind legs. A, rows of spines on tibia of
a leafhopper (Cicadellidae); B, single or grouped
spines on tibia of'a spittlebug (Cercopidae); C, large
spur on apex ol delphacid tibia (Delphacidae). sp,
spur; tb, tibia.
,\\
above; widelydistributed . . . .. .. 11
lla(f0b) Front wings broadly oval, longer
than body, at rest held almost verti-
cally at sides of body (Fig. 151C),
and costal area net-veined; somewhat

7a(6b) Anal area of hind wings with a i::l Ti: :: .':,'J,ää i ;:::ii":;#:,
network of many crossveins
(Fig, l50B) . . . . . (p. 164) Fulgoridae 11b Front wings variable in size and
shape, often shorter than abdomen,
Anal area of hind wings without a and costal area not net-veined; hind
network of crossveins., . ., . . . . . . E tibiae with spines on sides and apex;
often stout and frequently beetle-
like . . (p. 165) Issídae
8a(7b) Hind tarsi with 2 spines on apex of
second segmenl, apex rounded . . . .9

Hind tarsi with a row of spines on L

apex of second segmenl, apex
notchedortruncated ....12

158 Subclass Pterygota

 t i-
: ''iÌi r
!,
12a(Eb) Terminal segment of beak not more l6b Tarsi l-segmented and with I clalv;',
than I l/2 times as long as wide; female wingless, often legless' and
often slender or fragile forms with body covered with hard shell or rvaxy
bright coloration . . (p. 16. ' Derhidae secrelions (Fie, 155); nt:tle vvith 1 pair
of n'ings a¡rrl no ire ak (Iiig. n55E) . . .
l2b I erminal seglnent ¡t bc k al lcas( . " , . (p . l1(l) |>u¡re rJ',ttttil.r l)c,ccoidea
twice as long as vride; I orm vari-
able.. ...13
t7a(1óa) Antennae 5- to l0-se¡;rnr:nted (usually
l0); front wings oflen thicker than
l3a(l2b) Front wings overlap at apex hind wings; jumping insects
(Fig. l50C); bodY somewhat flat- . . . (p. '.166'l PsYllidae
tened . ...(P.165)Achilidae
17b Antennae 3- to 7-segmented; wings
13b Front wings do not overlaP at aPex;
bodynotflaftened,.....l4 :::: :: î::'::r "i':T:'::':::ii

14a(l3b) Head strongly elongated at front lEa(17b) \ilings opaque whitish and covered
(t'ig. l5lB), or front with 2'3 raised with a whitish powder; hind wings
nearly as large as front wings; no cor'
'l
1 iir : : T :ä1i ä ,'Li;; ;o; ":,; ; ";,
o; nicles. .(p.167)AleYriidae

l4b Head not strongly elongated at front rEb Wings clear and without whitish
although sometimes slightlY ex- powder; hind wings much smaller
tended, front of head without ele- than front wings; cornicles usually
vated ridges or only l; median ocellus present(Fig. f53) .......19
usuallypresent ...'......15
¡l
l9a(fEb) Front wings with 4-5 (rarely 6) veins
fSa(l4b) Abdominal ferga 6'E are below darkened stigma, veins reach
chevron-shaped, sometimes sunk be' wing margin (Fig. 153); cornicles
low rest of lerga' and with wax-se- usually present (Fig. l!i3); antennae
creting pores; western U.S. (genus generally6-segmented ....20
Oectidius\ Kinnaridae
l9b Front wings with 3 veins below
15b Abdominal terga 6't are rectangular; stigma, veins reach wing margin; cor'
widely distributed . ' (p. 1ó5) Cixiidae nicles absenl; antennae 3'5 seg'
mented ...21

f6a(1b) Tarsi 2-segmented and with 2 claws;

4 wings or non€. . .17 20a(19a) Cornicles typically preser't and
conspicuous (Fig. 153); M vein in
front wing branched (Fig. 153); no : .'.
waxy or woolly secretions.
. . (P.1ó7) APhídidøe

Order HemiPtera
 20b Cornicles minute or absent; M vein in
fronl wing not branched (Fig. l4óA);
many wax glands and waxy or woolly
secretions . . . . (p. 168) pemphigidae
Cu, Cu
Cu,
Cu,
Cu,
Figure 146 Front wings. A, woolly and gallmaking
fnli{s (Pemphigidae); B, pine and spruce aphids
(Adelgidae); C, phylloxerans (phylloxeridae).
2la(l9b) Wings held rooflike at rest; veins Cu,
and Cu, in front wing do not join at
bases (Fig. 1468); wingless females
covered with threads and masses of
lvax; on conifers. . (p. 169) Adelgidae
ztb Wings held horizontally at rest; veins
Cu' and Cu, in front wing join in a
common stalk at bases (Fig. l46c)i
wingless females not covered with
threads and masses of wax, al mosl
covered with powder
. . . . . (p. 110\ Pht,lloxeridae
160 Subclass Pterygota
Family Cicadidae- Cicadas¡
Cicadas are medium to large (16-50 mm long),
robust insects with four membranous wings.
The Iarge beak is easily seen and the antennae
are short and bristlelike. Cicadas are usually
dull or dark with lighter colored markings.
Males of each species produce loud, sometimes
shrill, characteristic buzzing sounds generally
from popping in and out riblike bands associ-
ated with a large resonating air sac; the bands
and sac are located in each of two ventral or
Iateral cavities at the base of the abdomen.
Each cavity also contains an auditory organ
(tympanum) and may be partly covered by a
plate. Some species stridulate or make clicking
noises with their wings.
Cicadas are primarily arboreal in the
eastern U.S.; many frequent forbs and grasses,
in the southwestern states where fewer trees
grow. Females deposit eggs in twigs of trees and
shrubs by slitting the bark and wood, usually ,
causing the twig tips to die. The nymphs drop
to the ground, enter the soil, and feed on plant
roots. The hemimetabolous development takes
4-17 years depending on the species. The ma_
ture nymphs dig out of the soil, climb onto an
object such as a tree, and molt leaving promi-
nent skins attached to the tree. (t
The dogday group of cicadas are large,
blackish species that occur from July to Sep-
tember. The periodical cicadas of the eastern
U.S. (genus Magicicada) occur from May to
early July, have reddish eyes and wing veins,
and l3- or l7-year life cycles. There are numer_
ous broods of each of the three l3-year or three
l7 -year Magicicada species. The broods emerge U
in different years, have different geographical
ranges, and may consist of one to three species
of each life-cycle type, The large broods emerge
only every few years. The name ,,locusts" has
been commonly misused for these cicadas be_
mbers that periodical-
40,000 under a tree),
the biblical locusts or
migratory grasshoppers (Orthoptera).

Figure 147 Linne's cicada, Tibicen linnei (Cicad-
idae).
Common Species
Magicicada septendecim (Linnaeus), Periodical ci-
cada or Linnaeus' 17-year cicada-2l-33 mm; black
and reddish; eyes reddish and sides of thorax red-
dish; ventral part of abdomen with reddish brown or
yellow markings; eastern U.S. west to NB and
southwest to TX (song is l-3 second buzz that drops
in pitch at end and sounds like "Phaaaroah"; begins
at dawn in May and June).
Magicicada lredeci¡n Walsh and Riley. Riley's l3-
year cicada-like M. seplendecim but range is south-
ern U.S. north to IA, west to OK.
Okanagana bella Davis-24 mm; shiny blue-black
with dark orange markings; western U.S., south-
western Canada.
Ptatypedia areolata (Uhler). Orchard cicada-19-
20 mm; dark bronze; amber and yellow markings;
greenish yellow crossband between pro- and meso-
thorax; western U.S., southwest Canada (common
in orchards),
Plat,vpedia ntinorllhler. Mirror cicada - 7 mr¡; Iike
P. araolui¿t but Ìa*:ks palc baii,ì ('n th')ì l\; \\ csterÌr
U S (song. is brie I , sharp r' in-s L licks r peatcrl 4-5
tinrcs; in trces on grassy slo¡res altcl alorlg :reeks).
Tibicen canicularis (Harris). Dogcla¡ crcacla-
32 mm; greelt and black; anterior part ol pron()tunl
(collar) green or brown; wing margins greenish; wing
length usually < 40 mm; eastern and midwestern
U.S. (song is even, rvhining buzz without vibrato and
lasting I minute or less).
Tibicen /innei (Smith and Grossbeck) (Fig. 147).
Linne's cicada-like T. canicularis but wing length
usually ) 40 mm; eastein and midwestern U.S.
(song a shrill l5-25 second vibrato; from trees in
afternoon).
Tibicen pruinosa (Say)-43 -47 mm; green and black;
wings and legs greenish; mid- and southwestern U.S.
(song is "7a-wie"; often with T, linneí),
.
Family Membracidae- "i,il
Tieehóppers ;¡
Treehoppers are Ieaping insects usually less :,i:;
than 12 mm long, and are characterized by ttre irìÏ
large pronotum that projects over the head and .
extends backward over the abdomen. The in'
sects may appear humpbacked, thornlike, or ,i
have various spines and horns as part of the
pronotum. Treehoppers feed primarily on
stems of trees and shrubs although some occùr
and feed on weeds, grasses, and crops. Nymphs
have a less-expanded pronotum and flften
branched spines on the dorsum of the abdo-
men. Many nymphs and some adults are tended
by ants for the sweet anal secretion treehoppeis .'
produce.
Common Species
Campylenchia latipes Say (Fig. l48A)-
6-8 mm; reddish brown; without spots; anterior pro-
notal horn with prominent ventral keel and 2 lateral . .
ridges; pronotum hairy and densely punctured; lr:'
throughout North America (on golcienrod, a:-'alfa¡' i ,
clover, grasses, others). 1l
Enchenopa binolata (Say). Twomal'ked t:^ehop-
per-5-ti mm; dark brown; 2 yellow r;pots on mid'
dorsal line of pronotum; anterior horn pro.iects up
and forivarrl, is slightly expaniled at ti¡r, rrnd has 3
rrlge s aÌong le ngth; eastern I j.S.
Plul,\'cotit t,ittuta (l:abricrus)--;0 rnln; rlull olive
green, blue-green or bronze; sttip':s,lr r,:d-orange
dots; line punctures; middle horn long, sh<trt, or ab-
scnt; 2 short lateral horns; most of U.S, (on oaks)'
Order Hemiptera
 F ly Cercop idae
spi t t lebugs or rog oppers
Adults are small (less than 13 mm), very com-
mon hopping insects that are shades of gray or
brown. Spittlebugs are disringuished from the
similar leafhoppers by the one or two stout
spines on the hind tibiae rather than the rows of
C
small spines (Fig. 1a5B). Spittlebugs feed on ql
weeds, grasses, shrubs, and a few trees. The
name is derived from the inactive nymphs that e.
surround themselves with a conspicuous mass
of white spittlelike froth which provides a moist
habitat.

Common Species
Aphrophora permutata Uhler-8-10 mm; mottled
brown; eyes twice as wide as long; spzrce between
ocellrrs and ¡rosterior margin of heacl =, or ) di- €t
ar-nctcr ol occllus; westcrn U.S eal;t ro lr,fl (,rn dan-
dclion, dock, leltLrce, otitcrs). qÐ

Aplrrophora quadrirtotota Say-7.0-8.1; mnr; pale; û

red ocelli; head and pronotum rvith slil¡htly raised
middorsal line; anterior margin of front wing with 2 F,
large, clear spots ringed with da¡k brown; North u_
Figure 148 Treehoppers (Membracidae). A, Cam- America east of Rocky Mts.
pylenchia latipes; B, buffalo rreehopper, Sflc- Ç",i
loc'ephala bubalus.
Aphrophora saratogensis (Fitch). Saratoga spittle-
Stictocephala bubalus (Fabricius) (Fig. laEB). Buf-
lalo treehopper-9-l0 mm; bright green or yellow-
bug-9-11 mm; brown; light middorsal stripe onl
head and pronotum; front wings with tan and silvery
u
mottled crossbands; North America east of Roóky ,;t] r
ish; no colored bands; sternum yellowish; pronotum
Mts. (on pines, especially red pines in Great
\-,,
with 2 stout side horns projecting outrvard; front Lakes
sta tes). !
wings not covered by pronotum; rhroughout North
America (weeds, legumes, trees [especially elm, ap-
plel).
Clasloptera proteus Fitch. Dogwood spittlebug-
4-5 mm; black with 2 yellow crossbands on head and r t-_
Slictocephula diceros (Say)-9-10 mm; dark bro*,n;
I on thorax and each front wing; scutellum yellow;
brown spots;2 sharp lateral horns; posterior pro-
throughout U.S., southern Canada. U
notal process wirh 2 white bands; front wings not
covered by pronotum; eastern U,S. (elderberry,
shrubs ol wet areas). "Q
e
Stictocephala festina (Say)-5 mm; bright green; no
pronotal horns; most of U,S., sourhern Canada (on together; blackish, curved marking at tip of frontì ti(_
wing; most of North America.
alfalfa, other forage, weeds, fruit trees).
(-
ermis (Fabricius)-7_9 mm; uni_
reddish; no pronoral horns; tip of
throughour U.S. (especialÌy on
lover).
C
(

162 Subclass Pterygota

qJ

\ Draeculucephala mollipes (Say) (Fig t49Eì). Water-
:\
ri
llt1
i-'
4-i
4i
;:
I
ii
1l lr
ü
tï
I tt
Figure 149 A, diamondbacked spittlebug, Lepy-
ronio quadranp.ularis (Cercopidae); B and C, leal-
hoppers (Cicadellidae): B, watcrcress sharpshooter,
Draeculacephula mollipes, C, painted leatlopper,
Endria ini¡nit'a.
Family Cicadel lidae- Leaf hoppers
This very Iarge family (nearly 3,000 North
American species) consists of small, jumping
insects chiefly under l3 mm in lengt,h' Leafhop-
pers are similar to froghoppers but have one or
more rows of small spines on the hind tibiae.
Many are brightly colored and some produce
weak sounds. Cicadellids occur on most types
of plants and in most habitats. Some are limited
to one host, whereas others are lound on many
plant specics. Many are plant pests due to feed-
ing on pìant sap, destroying cells, plugging ves-
sels, ancl transmitting viruses. Some species mi-
grate to southern states in the iall and norlhern
states in the spring.
Common Species
Aceratagal!ia sanguinolenta (Provancher), Clover
leafhopper-2.5-3.5 mm; light brown or gray with
dark markings; dark spot above each ocellus; longi-
tudinal light brown stripe each sidc oi'pale middor-
sal line; lront wings pale brown with white on poste-
rior area; North America east of Rocky Mts. (on
legumes, other crops).
Circulifer Ienellus (Baker). Beet leafhopper-3.0-
3.5 mm; greenish yellow to dark brown; front wings
with dark markings or broken bands; western U.S'
east to IL, southwestern Canada (on beets, sugar
beets, tomatoes, others).
cress sharpshooter-6-8 mm;ycl)ovu ¡rrtd glet:n; head
exlcncieci and pointed anteriorlv in rlorsalantl lateral
views; thror.rghout U.S., souihern Call¡tda.
Empoasca fabae (Harris). I)otalo leafhopper-
3 mn.r; pale io irridescent green; 6 ol' 8'lhite spots on
anterior rnargin of pronotum; small white "H"
marking on scutellum; front wings with crossveins
only near tip and without a b¡oad cr:ll that wraps
around apex; I apical cell in hind wing; North Amer'
ica east of Rocky Mts. (on potato, alfalfa, clover,
beans, apple).
Endria inimica (Say) (Fig. 149C). Painted
leafhopper-4 mm; grayish yellow; 2 black spots on
top of head, anterior margin of pronotum, and base
of scutellum; most of North America (on lawns,
meadows).
Exitianus exitiosus (Uhler). Cray lawn leafho. per-
3.5-5.0 mm; color variable; sometimes brownish;
dorsum of head grayish white and sometimes tinged
orange-yellow; red ocelli with large, round, black
spots on margins; irregular black spots on-pro-
notum; clear front wings with dark veins; long'
broad cell wraps around wing tip; throughout U.S.,
southern Canada (on grasses, wheat, legumes).
(Förster)-8-9 mm; toP of
band on anterior margin;
ings red with longitudinal
S. southwest to TX, south-
eastern Canada (on ornamental shrulls, e.9., For'
sythia).
Idiocerus patlidus Fitch-6.0-6.5 mm; white with
green tinge; without markings; eyes reddish brown;
throughout U.S., southern Canada (one of dozens
Order Hemiptera 163
ol Idioceru.s spp. on ivillows, poplar, aspen, cotton-
wood).

Prtraphlc¡tsitts ¡rroretus (Say)-_s.:-0.0 mm; dark

brorvn; lrcÎi,ilr corcrcd rvith ntinr¡tc spots; top ol
hcad l,'.1 lLrngcr at nticicllc rhan nc\r to e),cs; head at
e
lcast r iclth ol pt onotrnìì; lnost ol North Anrcrica.
o
Family Delphacidae- Ð
Delphacid Planthoppers
This is the largcst planthopper lamily. Its ø
members are small, lairly comrnon, and charac-
terized by a large apical spur on the hind ribia. u
Common Species
Stobaera Iricarinata (Say) (Fig. l50A)-4 mm; face
and front wings banded with whitrsh and dark mark-
ings; mesonotum pale or brownish yellow; Iegs white
and black; apical spur on hind tìbia rhick and convex
on sides with tecth on hind margin; mosr ol U,S.,
southcrn (-lnacla

Family Derbidae-
Derbid Planthoppers
The terminal segrnent of the bcak is about as
long as wide in this lamily. Some species resem-
ble small moths or caddisflies and the wings of,
many are broader toward the tips. Qtiocerus
spp. have branched antennae and elongated
heads that are bluntly but not slrarply angled.

Common Species
Apoche degeerii (Kirby) (Fie. l5lA)-l I mm to
wing tips; pink; veins of front wings dark pink;
lower margins ol head yellow and crossed with biack
lines; head sharply angled; southern and midwestern
U.S. (on deciduous trees).

Family Fulgoridae- Figure 150 Planthoppers. A, Stobaerø

Fulgorid Planthoppers
Some of the largest planthoppers in the U.S.
(up to 25 mm wingspan) occur in this lamily
which is recognized by the many crossveins in
the anal areas ol the hind wings.
(Delphacidae); B, Cyrpoptus belfragei (Fulgoridae);
C, Synecdoche impunctatø (Achilidae)i D, Oliqrus
humilis (Cixiidae); E, Bruchomorpha oculata (ls-
sidae).

164 Subclass Pterygota

Common Species
Cy'rpoptus bel.frugei Stål (Fig. l50B)-8-9 mm;
grayish bro*,n with reddish ringe; head broad, flar-
tened, and rounded anteriorly; wings with small
blackish patches; smoky bar near tips of wings and
clear areas along anrerior margins; southern U.S.
north to oH.
Common Species
Oliarus humilis (Say) (Fig. l50D)-5 mm; dark
brown; hind tibiae with 2-3 conspicuous spines; me-
sonotum with 5 longitudinal ridges; apical l/3 of
front wings with smoky or black patch; eastern U.S.,
southeastern Canada.

Family lssidae-
Family Dictyophar¡dae- lssid Planthoppers
Dictyopharid Planthoppers
Members in the genus Scolops have the head Most species are dark-colored and somewhat
produced forward into a long, slender, beaklike broad; some have a weevillike snout and short
structure (Fig. l5lB). The head is not produced wings.
forward in the genus Ph¡'lloscelis,' instead there
are two to three raised ridges on the head and
the lront f'emora are broad and flattened.

Common Species
Scolops pallidus Uhler-6-7 mm; yellowish and
gray; southwestern u.S,

l5l8)-6 mm; brownish

Scolops sulcipes (Say) (Fie.
gray; front wings with many netlike crossveins; east-
ern U.S. (meadows).

Family Achilidae-
Achilid Planthoppers r'

The apices of the front wings overlap in species

of this family. In the genus Catonia, the lateral
compartments of the pronotum are shorter
than the eyes.

Common Species
Synecdoche impunctata (Fitch) (Fig. l50C)-
5 mm; face with black spots separated by white
crossband; front wings uniformly brownish yellow;
eastern U.S southwest to OK, southeastern Canada.

Family Cixiidae-
Cixiid Planthoppers
The head is not usually extended forward in this Figure l5l Planthoppers . A, Apache degeerii (Dev
family. The genus Cixius has two to three bidae); B, Scolops sulcipes (Dictyopharidae); C,
spines on the hind tibiae and three longitudinal Acanalonia bivittata (Acanaloniidae); D, Metcolfa
ridges on the mesonotum. pruinosa (Flatidae).

Order Herni¡rterar 165

Common Species
Bruchomorpha oculata Newman (Fig. l50E)-
3 mm; black;distinct, short snout; front wings either
reach base of abdomen (short wing form) or extend
slightly beyond tip of abdomen; eastern Norrh
America.
Family Acanaloniidae-
Acanaloni id Planthoppers
These greenish insects have broadly oval lront
wings held almost vertically at rest. Acanaloni-
idae are similar to the Flatidae bur lack granules
in the triangular anal area (clavus) ol the front
wrng,
Common Species
Acanalonia bivittata (Say) (Fig. l5lC)-7 mm ro
wing tips; green, yellowish green, or brownish yel-
low; brown head is broad, rounded, and not conical;
2 longitudinal dark stripes from eyes to en,r of front
wings; eastern North America.
Family Flatidae-
Flatid Planthoppers
The front wings of ntembers ol'this lamily have
a unilorm rorv of crossveins aìong the anterior
margins and granules in the triangular anal area
(clavus).
Common Species
Anormenis septentrionalru (Spinola)-9-l I mm; dull
green; fronr wings without dots; front wings trun-
cate and not rounded at tip; eastern U.S. west to
Rocky Mts
Metcal.fa pruinosa (Say) (Fig. I5lD)-8-9 nlm; dark
gray abovc; undernc.ath paler; whitish powder on
body; lronr rvings with dark dots; easrern North
America.
Family Psyllidae-
Jumping Plant Lice or Psyllids
These small (2-5 mm) jumping insects resemble
tiny cicadas although many psyllids have long
antennae. The front wings are usually mem-
166 Subclass Pterygota
branous but may be thickened and the four
wings are held rooflike over the body. Many
nymphs produce strands and masses of white
wax; nymphs in the genus Pachypsylla cause
numerous galls to form on hackberry leaves. A
few species are pests of crops.
Common Species
Paratrioza cockerelli (Sulc). Potato psyllid, tomato
psyllid-1.4 mm; black; white crossband on lst ab- /:':
dominal segment; white "Y" marking on last seg-
ment; western U.S. except WA and OR, southwest-
\J
ern Canada (on potato, tomato, wild Solanaceae).
Psylla floccosa Patch. Cottony alder psyllid- 6l
2-4 mm; pale yellowish green or brownish; nymphs
yellow and green; northern l/2 of U.S. (cottony wax .@
masses on alder in spring and early summer; Iate
summer masses are woolly alder aphids lPemphi-
gidael; brown nymphs with dark abdominal cross-
€
bands under waxy masses in CA and NV are P. alni
qmericana Crawford). \-
Ps1'lla p.r,ricola Försrer (Fig. l5:!A). pear 6
psylla- l 5-1.0 mm to rving tips; light r:ran¡¡e, red-
dish l¡rou,n or brown to black; longitLLdinal srripes
(usuall-y 4) on mesothorax; black sl)or. Íìt tip of anal
arerì on lronr wing; cross stripes on abd,cmen; east-
crn Li S.. Irar West, southwestern Canad¿1.
1i"
í-_
(
Figure 152 A, pear psylla, Psy//a pyricola (Psyl-
lidae); B, greenhouse whitefly, Trialeurodes vapor-
a rio ru m (Aleyrodidae).
L
(,
(
('
Family Aleyrodidae-Whitef lies Aphids form a large family whose species
Adult whiteflies are small (l-3 mm), winged in- are known for their variation (polymorphism)
sects that are covered with white, powdery wax in shape, size, and winged condition, and their
and resemble tiny moths. Larvae (nymphs) complex life cycle. Typically, small numbers of
have a modified type of hemimetabolous devel- eggs overwinter and hatch in the spring into
opment: (l) the first instar is active; (2) the wingless females. These females reproduce.par-
following instars are legless, produce waxy se- thenogenetically (without fertilization) arrd give
cretions over their bodies, and the wings devel- birth to young rather than laying eggs. Several'
op internally; and (3) the next to last instar generations of wingless females may occur until
(pupa) is inactive. Whiteflies are primarily sub- winged females are produced which migrate to
tropical and tropical. In the U.S. several species another species of host plant to continue repro-
are important greenhouse and citrus pests. Be- ducing. Late in the season winged forms return
sides feeding on sap, whiteflies secrete a sugary to the original species of food plant, a gen-
anal material (honeydew) on plants that attracts eration of males and females is produced, mat-
a sooty fungus, ing occurs, and eggs are depositecl for overwin-
tering.
Common Species
Dialeurodes cilri (Ashmead). Citrus whitefly-
2 mm; whitish powder on body and wings; wings
without darkened area in middle; FL and other Culf
coast states, CA (on citrus, orna.mental \

Tri o I eu rod e,ç vo po ra ri o ru nr (\\'es ili'ood ) i Fi g. I 5 2B)

Creenhouse * hitcl-ly-l .5 mnl: *'hitisì pou cìe r on
body ancl wings; nymphs pale gre,:n, llrt and oval,
long filaments extend from b,odies; throughout
North American (in greenhouses), southern 1/2 oî
U.S. (outdoors on fruit, vegetables, flowers).

Family Aphididae-
Aphids or Plantlice
Aphids are small (l-6 mm), winged or wingless,
pear-shaped insects that are found clustered on
stems or leaves of plants. Most species have a
pair ol tubes (cornicles) near the posterior end
of the abdomen (Fie. 153). When an aphid is
disturbed the cornicles secrete alarm phero-
mones which stimr¡late delensive behavior
(e.g., dropping from the plant) of nearby
aphids. Aphids produce a sugary anal secre-
tion, honeydew, often seen as a shiny coating
on leaves. Ants and other insects feed on hon-
eydew and ants often tend aphids. Many species
are serious plant pests: they weaken a plant by Figure 153 Winged and wingless bean aphids,
Aphis labae (Aphididae). c, cornicle.
sap removal and may cause it to wilt, they cause
leaves to curl and turn yellow, and some trans-
mit virus diseases of plants.

Order Hemiptera 'lg7

Common Spccies
,lc'vrtÌtosi¡tlton /)t\unt (Ilarris) I,:a apl iLl
-l 0-
-l 5 nlnt; prrlc grcen; rips ol ribilLc. rarsi, trìd,ior-
nicìcs blackish; rvingcd forms rvrrh rcllorvish hritrrn
head and rhor¿ìx; throughour Norrh Arncrica (on
pcas, alfalla, clovcr, orher legumes)
Aphi.s fabae Scopolì (Fig. 153). Bean aphid-1.5-
2.5 ¡rm; dark olive grcen ro black; rips ol'ribiac, ab-
donrcn, cornicics, a¡lil all ol tarsi black; caucial a¡r-
pendage bushy; throughout North America (on
beans, beets, dock)
Aphis goss¡pii Glover Corton aphìd, melon aphid-
l-3 mm; tan, brown, green, or black; head tl¡bercles
between antenrlae not prominent; several dark
dashes on dorsum ol'abdomen; cornicles black;
thror,rghoul Ll S., southern Canada íon melons, cu-
cumber, cotton, citrus, vegetables, ornamentals,
pigweed, dock, other weeds).
Brevicoryne l¡rassicae (Linnaeus). Cabbage aphid-
1.5-2.5 mm; grayish green; striped; dusty appearing
from wax covering; throughout North America (on
cabbage, cauli llower, broccoli).
Hyalopterus pruni (Ceof froy). Mealy plum aphid-
l-2 mm; green; covered with whitish powder; 3lon-
gitudinal darker green ìines; winged iorm with head
and thorax brown; throughout U.S , southern Can-
ada (on plum, other stone fruits).
ìvlacrosiphoniella sanborni (Gillette) Chrysanthe-
munt aphid-1,5-2.0 mm; amber brown ro black;
shiny; corniules black and most of length rvith scale-
like markings: most ol U S., sourhern Canada (on
chrysanthemums, in greenhouses).
Macrosiphum avenoe (Fabricius) English grain
aphid-1.5-2.5 mm; grass-green; somerimes yellow
or pink; head with brown marking; darker dorsal
blotch on abdomen of wingless form; darker median
line in winged form; throughout North America (on
grasses, cereals).
Macrosiphum rosae (Linnaeus), Rose aphid-
2.0-3.5 mm; grcen or pinkish; winged form wirh
black anrennae, head, thorax and cornicles;
throughout U.S., sourhern Canada (on roses).
Myzus persicae (Sulzer). Creen peach aphid-
1.5-2.5 mm; wingless lorm pale to dark green with
stripes in summer and pinkish in lall; winged form
dark brown wirh dark dorsal patch on yellowish ab-
domen; 2 prominent, converging head tubercles be-
168 Subclass Pterygota
t\\ecr'ì antcnnac in all f'orms; throu¡5hour: U.S.,
sorrrlrcr n C:rnada (on, fr uit, vcge rrtrles, shrubs, flow-
ef\)
R
I
ltttlt¡tIitçiltltt.t ttt t¡taicli.ç (Iritch¡. iìonl lt:aI aphid-
.l 2 5 ¡it¡n; blLrish green; slen<icr; rhroughorrt U.S,,
fl
sor¡thcr n Canada (on corn, sorghrrnrs). 0
Sc'hizaphi.s grantinum (Rondani). tìreenbug- {:
1.2-2.0 rnnr; pale green; winglcss fornl with dark
green rnedian stripe; winged fornr with brownish ¡1 '
yellow head and black cornicles; cornicles narrower
than hind tibiae; no reddish blotches at bases of cor-
nicles; throughout North America, chielly in South ü
and Midwest (wheat, other grains, grasses, alfalfa).
€¿
Therioaphis tttaculaÍa (Buckton). Spotted alfalfa
aphid-1.5-2.0 mm; pale yellow; 6 + íows of black
spots on dorsum; smoky areas along wing veins; €
most of U.S. except New England.
6i
(-
Family Pemphigidae
( = Eriosomatidae)-Woolly and
Gallmaking Aphids L
The cornicles are small or absent in this family {,
and most members have wax glands that pro-
duce large amounts of woolly or waxy material
that covers part or all of the body. Mouthparts
are absent in the males and females of the bisex- IJ
ual generation. The family is classified as a sub-
lamily (Pemphiginae) of the aphids (Aphidi-
dae) by some taxonomists.
Most species alternate betlveen host
plants and galls are produced on trees and
shrubs by many species. The vagabond aphid
gall oi poplar and cottonwoods, caused by
Mordwilkoja vogabundo (Walsh), appears as
an irregular, folded and twisted leaf
(Fig. l5aB). The leaf petiole gall of poplar is an
oval gall with a transverse opening on the leaf
stem; it is caused by Pemphigus populitranEver-
srzs Riley (Fie. l5aC).
Common Species
Eriosoma lanigerum (Hausmann) (Fig. l54A).
Woolly apple aphid-0.5-2.5 mm; reddish or pur-
plish; bluish white woolly masses on body; through-
out North America (on roots and bark of apple,
pear, hawthorn).
 Figure 154 A-C, woolly an{ gallmaking aphids (Pemphigidae): A, woolly apple aphid'
Eiiosoma lanigerum; B, vagaÙond aphid gall of poplar and cottonwood caused by Mord-
witkoja vagabunda; C, leaf petiole gall of poplar caused by Pemphigus populitransversus,
o, gail on spru.e caused by eastern spruce gall aphid, Adelees abietis (Adelgidae).

Prociphilus lessellatus (Fitch). Wootly alder Common Species

aphid-0.5-2.5 mm; woolly masses cover body; no
cornicles; east of Rocky Mts, (on alders and maples
from middle summer to early fall; early summer
masses are cottony alder psyllid [Psyllidae])'
Adelges abietis (Linnaeus) (Fig. l5aD)' Eastern
spruce gall aphid-1.5-2.0 mm; brownish yellow;
northeastern and northcentral U.S., southeastern
Canada (forms large, brown or gray pineapple-
shaped galls primarily on twigs of Norway and
spruce).

Family Adelgidae (= Chermidae) Adetges cooleyi (Cillette). Cooley spruce gall

Pine and Spruce Aphids
The winged or wingless members of this family
lack cornicles, veins Cu' and Cu, are separated
at their bases in the front wing (Fig. l468), and
wingless lemales are often covered with waxy
filaments. These species live and feed only on
needles, twigs, or in galls of conilers. Generally
a different kind of conifer acts as an alternate
host during part of the year.
aphid-1.5 mm; reddish brown; white, woolly wax;
similar to A. abietis; northern U.S. from Pacific to
Atlantic coasts, range of white spruce in Canada
(forms green or reddish brown galls on blue, Engel-
man, Stika, and white spruces; wax oll Douilas fir).
aphid-
Adetges piceae (Ratzeburg). Balsam woolly
I mm; black; convex; covered with white, curly wax
threads; northeastern U.S., Appalachian Mts., OR'
WA, British Columbia, southeastern Canada (on
firs but not Douglas firs).
Order Hemiptera

Pineus .itroöl (Hartrg). Pine bark aphid-l-2 mm;
dark brown; covered with white, fluffy wax; forms
spots and patches on rrunks and limbs; most of
U.S., southern Canada (on Austrian, Scotch, and
white pines).
Family Phylloxeridae-
Phylloxerans
Phylloxerans are tiny, winged or wingless in-
sects that have 3-segmented antennae, lack cor-
nicles, and have the bases of veins Cu, and Cu¿
joined in a common sralk (Fig. l46C). The
wings are held horizontally; wingless forms
sometimes have a whitish powder on them but
never waxy threads. The grape phylloxera,
Daktulosphoira vitifoliae (Fitch), has been a se-
rious pest of European grapes grown in the
western U.S. but causes little damage to native
grapes in the eastern states. The tiny (0.3-
1.2 mm), wingless, yellowish or greenish brown
phylloxerans form galls about half the size of a
pea on leaves of wild and sometimes cultivated
grapes in the eastern U.S. The Pacific coast
populations cause root galls and eventually kill
the European grapes. Crafting European
grapes to resistant rootstocks of native grapes
has alleviated most of the problem caused by
phylloxera.
the grape \
Most of the leaf and leaf stem galls on
hickory trees are produced by species in the ge-
nus Phylloxera. The disk-shaped to lobular
galls are recognized by the presence of ,. n open-
ing on tìrc upper ot lorvcr sLlrla:,.'
SUPERFAMILY COCCOIDEA-
Scale lnsects
The families in this group consist of small
(0.5-8.0 mm), winged and wingless insects that
are often unlike the other Homoptera in ap-
pearance. Adult females are always wingless,
very sluggish or fixed in position, and usually
are covered by a hard scale or waxy secretion,
170 Subclass Pterygota
f,,
@
or have a tough integument. Some females lack
legs and have reduced antennae, and often the
abdominal segmentation is indistinct. Adult
t
males have no beak, one pair of well-developed ü
wings (rarely wingless) on the mesothorax, a
pair of filaments (halteres) on the metathorax, o
and sometimes an elongate process at the tip of
the abdomen (Fig. l55E). The absencl of 0
mouthparts and the presence of the abdominal
process help separate males flrom the similarly €
appearing gnats (Diptera).
The active, first instar nymphs (crawlers) \r
have legs and antennae and disperse themselves
on the host plant. After the first molt the legs
a,
and antennae are generally lost and most spe-
cies become sedentary. A waxy or scalelike cov-
6'
ering may be secreted as the nymph develops @
toward the adult stage. Adult females repro-
duce under their permanent scale or wax cover- ü
ing. The wings of males develop in the pupalike
resting stage of the last instar; adult males are \-
tiny, active, flying insects.
Fam i ly Margarodidae- Margarodid
Scales or Giant Coccids and
Ground Pearls U
Females are often large and brightly colored.
Yellowish green, beadlike wax cysts are formed ü
by members of the genus Margarodes and are
I
called ground pearls; they occur on roots of
many plants in the warm areas of the U.S. M.
/airrgi Jakrrbski and M. meridic¡nøl¡,ç Morrison
arc pcsts rrl llrrlns in i"he soulhcnl {J,li
.
Clorumcn Spccies
Icer¡a purcha,ti Maskell. Cottonvcushion scale-
l0-15 nrm with egg sac (9); reddish brown; large,
white, t'ìuted cotrony egg sac; SC and Gulf coast
states to CA (on citrus, other trees).
Fami ly Ortheziidae- Ensigrr Scales
Members in this family live on plant leaves,
stems, and roots. Females are usually covered
with white, waxy plates and at times carry a
long egg sac that protrudes posteriorly from the
abdomen. The egg sac of all other scale insects
is attached to a substrate.
Common Species
Orlhe:tu lnri.qni.s Brorr.'ne (Fi¡r 1554) (ìrce nhousc
ortllezra 8 g rrlnt ro tip ol \\'lt). lgg sa, dul i:.reen;
white and longirudinally ribbed wa\! egg sac e\-
tends ) 2 times bodr- length; Lhror.rghout \orth
Anlerica (onramental planrs in greenirou;es)
Family Kerriidae- Lac Scales
Most species are subtropical and tropical. The
Indian lac insect of India and southeast Asia,
Laccifer lacca (Kerr), produces wax or lac used
in making shellac and varnish. Six species (ge'
nus Tachurdiella) occur in southwestern tJ.S.
on desert plants; the lac of some s¡pecies is high-
ly pigmcnted
F arni ly Coccidae --. S0 f t {i cale s
Flrnllcs ¡rt'oval and usually ccrnve)( (btrt some-
tir¡es llattened) with a hard, smooth cuticle, or
the cuticle is covered with wax. Some species
are turtle-shaped (Fig. l55B). Certain s¡ecies
are plant pests and several Asian species pro-
duce wax used in candles and medicines.

'li1'1 '.

Figure 155 Scale insects. A, greenhouse orthezia, Orthezia insignis (Ortheziidae); B, ter-
rapin scale, Lecanium nigrofasciaturn (Coccidae); C, a gall-like scale, Kermes sp.
(Kermesidae); D, oystershell scale, Lepidosaphes utmi (Diaspididae); E, San Jose scale,
Quadraspidiotus perniciosus (Diaspididae). a, mature female; b, immature female; c,
im-
mature male; d, mature male.
Order Hemiptera
Common Species
Ceroplastes ceriferus (Anderson)-5-20 mm ( Ç );
spherical, black body with hornlike posterior tube;
covered with thick, whire, doughlike wax; through-
out U.S. (in greenhouses), sourhern l/3 U.S. (sub-
tropical ornamentals, citrus).
Coccus hesperidunt Linnaeus. Brown solt scale-
3-5 mnr (?); flat, oval body; broç'n or marblcd
brown, sometintcs palc l,ello*'or greenish; through-
out U S (rt grucnhoLrscs), soulhcrn li2 lJ S (orrt-
<loors).
Lecanium corni Bouché. European
lruit lecanium-
2.5-5.0 mm (ç); brown when marure; oval; very
convex; throughout North America.
Lecanium nigro/asciaturn Pergande (Fig. l55B).
Terrapin scale-2.5-3.5 mm ( ?); reddish ro dark
brown; very convex; squarish areas around slightly
crinkled margin outlined by black lines giving turtte-
shell appearance; eastern U.S., southwest to NM.
Pulvinaria innumerabilis (Rathvon). Cottony maple
scale-6 mm ( ç); brown or reddish brown; secretes
large, white, cottony mass of wax which elevates
posterior part ol body at angle; most of North
America (on maple, other deciduous trees).
Saissetia coffeae (Walker). Hemispherical scale-
3 mm (9); shiny brown; very convex; elliptical;
smooth; throughout U.S. (on ferns and other plants
in greenhouses and homes, citrus). \
Family Aclerdidae
Most members ol this family feed near the
bases of perennial grasses and sedges. All spe-
cies from North Anlerica are in the genus Acler-
da.
Family Kermesidae-
Gall-like Scales
Members of the genus Kermes (Fig. l55C) re-
semble tiny spherical galls or berries and are
found almost exclusively on oak lea :s and
twlgs.
172 Subclass Pterygota
Fam i ly Asterolecaniidae-
riÌ
e
Pit Scales
Some species in this small family produce gall-
0
like pits in the bark of their hosts (commonly o
oak) and others occur on leaýes. Most species
are in the genus Asterolecqnium; matvre fe- o
males are covered with a pale green, glassy se-
.@
cretion. A. puÍeanum Russell, which occurs on
holly and southcrn buckthorn prirnarily in the Í,.
Atlantic coast states, is slightÌy convex and
sometimes has wax threads at the mar,gins.
\l
€.,
Family Dactylopiidae-
Cochineal lnsects €,
Females have reddish, rotund bodies covered
with a white, waxy filamentous secretion. The e¡
cochineal insect, Dactylopius coccus Costa, oc-
curs on prickly pear cactus in Mexico and a few ö
other counrries (not the U.S.) and still retains
some value for its red cochineal dye. D. opun-
',c
fiøe (Cockerell), the cactus mealybug, and
D. confusus (Cockerell) occur on Opuntia and L
other cacti in the southern third of the U.S. and
are occasionally encountered in the warm des- t/
erts of Colorado, Utah, and Wyoming.
ü
Family Diaspididae-
Armored Scales
The bodies of immature males and females and
adult females are covered with a scale com-
posed of cast skins and a waxy secretion.
Diaspididae is the largest family of scale insects
(over 300 species in the U.S.) and contains
numerous, important pests of deciduous fruit
and shade trees, citrus trees, and ornamental
shrubs.
Common Species
,4onidiello aurantii (Maskell). Calrfornia red scale-
:;calc 1.7-2 0 rnm (Ç); red body givt:s transparent
scalt: a rc,.ldisir crrlor; nearly circular rvith lst cast
skìn a¡r¡rcarilrt ni¡rpJelikc ncar {_.L.lrror; Cz\ to s¡uth-
oaslcrn i. lì a¡lrl i'l- (chiell)' ¡tr ctl, u!; ¡e¿,rl'¡ jden-
tical to rl()tc )cllou¡ish vcllorv:;cal,:, A titrinia
f Corìuille¡rl)
Aspidiotus nerii Bouché, Oleander scale-scale
l-2 mm (Ç ); light gray; circular; llat; yellowish
brown cast skin near center; throughout U S. (on
English ivy, olcander, olive, citrus, subtropical or-
namentals)
Aulaca.spis ro.sdc (Bouché). Rose scale-scale 1.5-
2.0 mm (Ç ); white or grayish rihite, circular, flat,
body reddish ( Ç ); scale rvhite, narrow, 3 ridges (c¡);
throughottl LJ S, sotrthcrn C'anacja (cllr roses' brrslt
lruits)
l'urlutu (lrilch) Scrrrlv scale scale
(.-ltiortusltts
2.5-30 rnnr (Ç¡¡ scale 0 7-0 9 rlm (cr); gra¡"ish
whire; oystcrshell-shapcd and thìn ( Ç ) or $ hite,
elongate-oval rvith 3 ridges (cr); throughout U.S'
(apples, pears, other trees).
Chionaspis pinifoliae (Fitch). Pine needle scale-
scale 2.5-3.0 mm (ç); scale I mm (cr); snowy white;
oystershell-shaped elongate-oval; throughout North
America (on nec:dles of pine, other conifers).
Lepidosaphes ulmi (Linnaeus) (Fig. l55D). Oys-
tershell scale-scale 2.0-3.5 mm ( Ç ); scale
1.0-1.5 mm (c¡); gray or brownish, elongate oyster-
shell-shaped; very convex; throughout U.S', south-
ern Canada.
Pseudaulacaspis pentagona (Targioni-Tozzetti).
White peach scale-1.7-2.8 mm (Ç ); opaque white;
(ç without ridges
ast lbt skin often
ant thern I /2 U'S.
her al trees, lilac)'
Family Pseudococcidae-
Mealybugs
Most members of this relatively large family
(over 300 species in the U.S.) have a whitish
covering of mealy or powdery wax. The margin
is olten adorned with several waxy filaments.
The female's body is elongate-oval and the legs
arc normally well developed. These insects
range lrom the tropics to the arctic circlc but
arc predorninantly in warm climates; some are
major pests in greenhouses and on citrus. The
tamarisk manna scale of the Micldle East se-
cretes copìous amounts of honeydew to form
layers with the tree leaves. The edible sugary
product is believed to have produced the manna
in the biblical account of the Children of Israel.
Common Species
Planococcus cllri (Risso) (Fig. 1564). Citrus mealy-
bug-3 mm (ç); white; median stripe darker; anal
lilaments at most l/4body length; citrus regions (on
citrus, avocados, cotton, ornamentals), freque::tly in
greenhouses.
Pseudococcus longispinus (Targioni-Tozzetti)
(Fig. l568). Longtailed mealybug-3 mm ( 9);
white; body margin with t6 or 17 long, waxy fila-
ments, the 2 filaments at tip of abdomen as long as
or longer than body; throughout North America
(greenhouses, citrus, other subtropical plants).

Quadraspidiotus perniciosus (Comstock)

(Þig. I Sse). San Jose scale-scale I .5-2.0 mm Ç );
(
scale 0.8-1.0 mm (cr); gray; nearly circular with lst
cast skin appearing nipplelike at center (Ç) or ot'al
with skin at one end (cr); bodies bright yellorv;
throughout U.S., southern Canada.

Llnaspis euonymi (Comstock). Euonymus scale-

scale 2.0 mm (9); scale l'5 mm (cr); dark grayish
brown to blackish, yellowish cast skin attached,
broadly pear-shaped, convex (9); white' 3 ridges
(o); most of U.S., southern Canada (on euonymus Figure 156 Mealybugs (Pseudococcidae). A' citrus'
mealybug, Planococcus citri; B, longtailed mealy'
[chiefly in greenhouses]). bug, Pseud ococcus lo ng is Pin us.

Order Herniçrtera 173


Pseudococcus maritimus (Ehrhorn). Grape mealy-
bug-3-5 mm (ç); white; powdery wax covering is
very thin; lateral wax filaments short and slender;
slender anal filaments | / 4-l /2 body length;
throughout U.S. (in greenhouses), sourhern l/3 U.S.
straying north to MI (grapes, citrus, peaches, pears,
walnuts, ornamentals).
Family Eriococcidae
These insects are similar to Pseudococcidae but
the body usually is bare or only lightly covered
with wax.
Common Species
Gossypariø spuria (Modeer). European elm scale-
6-10 mm (9); reddish brown; oval body su rounded
by white cottony fringe; most of lJ.S. (on lr aves and
Iimbs ol elrns),
CENERAL RFFERENCES ON
HETEROPl'ERA
References ro smaller families appear in tjlarer and
Baranowski (1978)
Blatchley, W.S. 1926. Heteroptera or True Bugs of
Eastern North America, w¡[h Especia!
Reference to the Faunas of Indiana and
Florida Nature Publ. Co.,
II l6 pp.
Britton, W.Ë. e/ al. 1923. The Hemiprera or Suck-
ing Insects ol Connecricut Guide ro the In-
sects of Connecticut, Part IV. Conn'St. Ceol.
and Nat Hist Surv. Bull. No 34. 80? pp.
DeCoursey, R,M l97l Keys to the Families and
Subfamilics of rhe Nymphs of North
Amcrican Henriptera-Heteroptera. Proc. Ent.
Soc Wash, 13:413-428.
Drake, C J , and Ruhofi, F.A. 1965, Lacebugs of
the World: A Catalogue
Tingidae) Bull. U S Narl Mus. 243 634pp.
F'roeschner, R.C l94l-1961 Contributions to a
Synopsis of rhe Hemiptera of Missouri. pts.
I -V. Amer. M idl. Nar.
27:591-609; 3l:638-683; 42:t23-tB8;
67:208-240.
Herring, J L., and Ashlock, P.D. 1971. A Key to
the Nymphs of the Families of Flenriptera
(Heteroptera) of America North of Mexico.
Fla. Ent 54:207-212.
Hungerford, H.B. 1920. The Biology arrd Ecology
of Aquatic and Semiaquatic Hemiptera. Kans.
Univ, Sci Bull. I l:l-328.
1948 The Corixidae ol rhe
Hernisphere (Henriptera), Univ. Kan. Sci
Bull 32:l-¡ì27
j9 llclliptera, In l:rcslt H'atcr Btolog+,
--- '-W l(iT []dmundson (ed.). \Lilcv, N,\, pp 958_
912.
174 Subclass Pterygota
I
Knight, H.H, 194t. The plant Bugs or Miridae of,
Illinois. Ill. Nar. Hist. Surv. Bull. 22:l-234.
l9ó8. Taxonomic Review: Miridae of the e
Nevada Test Site and the Western United
States. Brigham Young Univ. Sci. Bull., Biol.
Ser.9:l-282.
o
Parshley, H.M. 1925. A Biblidgraphy of the North
A mericon Hemiptera-Heteroptera. Smith Col-
lege, Northampton, Mass. 252 pp.
o
Pennak, R.W. 1953. Fresh-Lltater Invertebrates of û
the United Stqtes. Ronald Press, N.Y. 769 pp.
Readio, P.A. 1927. Studies on the Biology of the
Reduviidae of America North of Mexico.
ñ
Univ. Kan. Sci. Bull. 17:l-29t.
Slater, J.A. 1964. A Catalogue oÍ the Lygaeidae ol
the World. Vols. I and II. Univ. Connecticut,
ü
Storrs, Conn. I,668 pp.
Slater, J.A. and Baranowski, R.M. 1978. How to ',,,Ø
Know the True Bugs (Hemiptera:
optera) Wm. C. Brown Company Publishers,
Dubuque, Iowa.
Heter-
g
Usinge r, R I-
I956. Aquatìc Hcrni¡>tc.ra. ln Aquaric
lns¿¿¡; ç¡ Culi.fornio, R L [Jsingr:r (ed.). Éà
Univ, California ['rc.ss, IJerkr:lr:y Irp 182-
22fl
Van Duzee, E P. 1917. Catalogr-re of'the Hemiptera
ü¡
ol' America Norrh of Mexico, Iixceptilrg the t.
Aphididae, Coccidae, and Alcurodidae. Univ.
California Pubs. Tech. Bull. Ent. 2.902pp.
GENERAL REFERENCES ON TIO¡IOPTERA
t-'
Indianapolis. Annand, P.N. 1928. A Contribution Toward a tsì
Monograph of the Adelginae (Phylloxeridae)
of North America. Stanford Univ. Pub. Biol.
Sci.6:l-14ó. ü
Beirne, B.P. 1956. Leafhoppers (Homoptera: Cica-
dellidae) of Canada and Alaska. Can. Ent. f l ,
Suppl.2:l-180.
Britton, W.E. ef ol 1923. The Hemiptera or Suck-
ing Insects of Connecticut. Conn. St. Ceol.
and Nat. Hist. Surv. Bull. 34. 807 pp. -
Crawford, D.L. 1914. Monograph of the Jumping (r-
(Hemiprera: Plant Lice or Psyllidae of the New World.
U.S. Natl. Mus. Bull. 85. l8ó pp.
DeLong, D.M. 1948. The Leafhoppers, or ;
Cicadellidae, of Illinois (Eurymelinac-
26:122-146; Balcluthinae). Ill. Nat, Hist. Surv. Bull. L\J
24:91-376.
Doering, K, 1930. Synopsis of North American Cer.
copidae. J. Kan. Ent. Soc. 3:53-64,81-108.
Ferris, C.F. I937-t955. Atlas of the Scale Insects of
North America. Vol. l-7. Stanford Univ.
Press, Stanford, Calif.
Hottes, F.C., and Frison, T.H. 1931. The plant
Lice, or Aphididae, of Illinois. Ill. Nat. Hist. (
Surv. Bull. 19:121447.
Western Howell, J.O., and Williams, M.L. 1976. An An-
norated Key to the Families of Scale Insects
(,
(Homoptera;Coccoidea) of Amt:rica, North
ol Mexico, Based on Characteristics of the
Adult Female. Ann. Ent. Sioc. Amer.
69:l8l-189.
U
 , .: j

I ii' 4..
Kennedy, J.S., and Stroyan, H.L.C. 1959. Biology Oman, P.W. 1949. The Nearctic Leafhnppe i1',
of Aphids. Ann. Rev. Ent. 4: 139-160. (Homoptera:Cicadellidae), a Cenèr i,'
MacCillivray, A.D. l92l . The Coccidae. Scarab, Classification and Check List, Ent. Soc. :

Urbana, lll. 502 pp. Wash. Mem. No. 3. 253 pp.

McKenzie, H L. 1967. The Nlealybugs of Calitor-
ni¿ Univ. California Press, Berkeley. 525 pp
Metcalf, Z.P 1923. Fulgoridae ol Eastern North
America, J. Elisha lv'litchell Sci.
38: I 39-230.
1945 A Bibliography of the
(Auchenoruhyncha). Dept. Zool. & Ent., No.
Carolina St. Coll., Raleigh, N.C. Vol' I -2.
1954-19ó3. Ceneral Catalogue oÍ the
Homoptera. No. Carolina St. Coll., Raleigh'
N.C. Fasc. IV. Fulgoroidea. Parts I l -18'
Fasc. VIl. Cercopoidea. Parts l-4' Fasc. VIII.
Osborn, H. 1938. The Fulgoridar: of Ohio. Ohio
Biol. Surv. Bull. 35:283-349.
1940. The Membracidae of Ohio. Ohio
Soc Biol, Surv. Bull. 37:5 l-101.
Palmer, M.A. 1952. Aphids of the Rocky Mountain
Homoptera Region. Thomas Say Foundation Vol. 5.
Hirschfeld Press, Denver. 452 PP.
Smith, C.F. 1972. Bibliography of the Aphididae of I 1,
the World. N.C. Agric. Expt. Sta. Tech. Bull, ¡¡'
216.717 pp. i..
Wade, V. 1966. General Catalogue of the Homop- ì '
iera. Species Index of the Membracoidea and r., :

Cicadoidea. Parts l-2. Fossil Homoptera (Homoptera:Auchenor'i'

1962-1967. General Catotogue of the rhyncha). A Supplement to Fascicle l- i ''

Hornoplera. USDA, ARS Fasc. VI. Mêmbracidae of the Gencral Catalogue of the ' .

Cicadelloidea. Parts l- 17 Hemiptera. N C. Agric. Exç,¡. 5¡". Pap' No.

Moore, T E 1966. l-he Cicadas ol \{ichigan 2160 40pp
(Homoptera:Cicltdidlci Pap \lich Acad \\,illiarns, M.l_, and KoszrartL', N{. lft72. Mor-
Scr :75-96 phology anrJ Systcrralics c't the Ct¡ccidae of
*ith Notc' o. 'fh,:ir
5I
Morrison, H, and Renk, A V l95 r A Sclected \/ìrgirria, Biology
Bibliography' of the Coccoidea USDA Agric (l-lomoptcra:Coct:oidcrt) \'lr.
Polytech. lnst,
Res. Serv lvf isc. Pub No. 7-ì4 222 pp First and Sr Univ Res. Div Bull.74.2l5 pp'
supplement (Morrison, H, and E N4orrison;
l9ó5; Ibid. 987). Second suppletnent (Russell,
L.M.; M. Kosztarab; and M.P. Kosztarab;
1970; lbid. l28l).

ORDER COLEOPTERA24
Beetles

Beetles comprise the largest order of insects
with over 300,000 species formirig about 40 per-
cent of the known insects. Beetles have
thickened front wings (elytra) that are soft,
leathery, or hard and brittle' The elytra usually
meet in a straight Iine along the middle of the
back and cover most or all of the longer,
folded, membranous hind wings (Fig' l57A).
Some beetles have short elytra andlor hind
wings or are wingless. Most beetles use only the
hind wings for flight; the elytra are raised to
allow for the hind wing movement. The prono-
tum is generally distinct whereas the mesono-
tum and the metanotum are hidden beneath the
elytra. The mouthparts of adults and most lar-
vae are adapted for chewing. Some predatory
larvae are able to suck body f,luids ol prey, Lar-
vae have a holometabolous type of develop-
ment.
The appearance, life history, and habitats
of beetles are so diverse that only general
statements can be made. Their sizes varv from
Iess than I mm to over 200 mm in length and up
to 75 mm in width (e.g., tropical rhinoceros and
goliath beetles). Body shapes vary from round
to very slender, and flattened to stout or cylin-
drical. Some adults have long snouts' horns, or
tubercles on the head, extremely long antennae'
greatly enlarged mandibles, or other diverse
features. The body and elytral surface may be
smooth or rough, brightly colored or dull.
24. Coleoptera: coleo, sheath; plera, \\iîÉ\s

Order Coleoptera
 ,j

Many beetles produce sounds by rubbing body

parts together (e.9,., f'emur ol' rhe lrind lcg
against thc ntargin ol' the elytron; head rubbed
against the front rnargin oi the prorhorax) or by
expelling air lrom rhe spiracÌes. Some members
of the Lampyridae, Phengodidae, Drilidae, and
Elateridae emit light (biolumenescence) from
special organs. Certain beetles in the families
Staphylinidae, Pselaphidae, Scydmaenidae,
and othcrs live in rhc lìests ol ants and tcrntites.
Beetles are primarily plant leeders anci
scavengers, although a significant nunlber are
predators of insects and other small inverre-
brates and a few are parasites. The plant-
feeding habits (fruit, seed, root, stem, and
wood borers; leal feeders) ol adults and larvae
have made many beetles serious pests of
agricultural crops and forests. Some beetles
also feed on museum specimens, clothing, and
stored food. A lew species transmit bacterial
and lungal diseases to plants. Life cycles vary
from several weeks in warm climates to lour or
more years in cool climates. Wood- and root-
boring beetlcs usually have longer life cycles
than leal-leeding species. Eggs are usually
rlcpositccl ilt ¿ur arca suitable lor larr,al clevelop-
nlcnt. The iarvac, somctimes calle d grubs,
typrcally ntolr rhree to live times. The immobile
pupa is soft, pale, and resembles rhe adult.
Beetles may be collected froni almost any
habitat by sweeping, beating, sifting, using
Berlese funnels and light traps, etc. Pit traps
baited with carrion such as old meat, a dead
mouse, or a lish head will attract scavenger
beetles, Molasses or brown sugar syrup placed
in a pit trap or in a container suspended from a
low tree branch will draw in many kinds ol
beetles. Wood-boring beetles are attracted t.o a
screen-covered container of turpentine sus-
pended in a tree. Adults are pinned through the
base ol the right elytron. Small beetles are
mounted on paper points. Larvae are preserved
in 7090 alcohol. Degreasing large members of
the families Carabidae and Scarabaeidae is
'¿,
imrne¡sed in ethcr. xylenc, or benzeue, ¿rnd the
solvcnt rc-ne r','ccl until it is no longer discc'lored. (r
Species: North America, 28,OOC; world, Ð
> 300,000. Families: North America, about
r 13. Ð
ç)
KEY TO
COMMON FAMILIES (.
OF COLEOPTERA {")
Familiarity with the external morphology of
beetles (Fig. 157) is important in identification.
Çi
For example, determining that the hind co4ae {u
divide the posterior margin of the first ab-
dominal sternite (Fig. l57B) will help separate a (t
very common family, Carabidae (and also the
suborder Adephaga), from the remaining ü
beetles (Fig. 158). The difference between open
and closed coxal cavities must be recognized
(Fig. l6a). The number, shape, and size of tar-
L
sal segments is important; a 5-5-4 tarsal for-
t'
mula means that the front and middle tarsi have
five segments and the hind tarsus has four. If L
the tarsus appears to be 4-segmenti:d and the
third segment is bilobed ("U "-shaped), the {}
tarsus usually is 5-segmented with the extra
tiny segment hidden inside the third segment
(-,
(Fig. l62D). I
la Head rarely extended into a snout; {,
antennaevariable,.,...,.2
i'-
Front of head usually exfended into a
snout (Fig. 2f3A) or broad and short
muzzle (Fig. 2flB); antennae arise .,:
along snout, often clubbed and el-
bowed (Fig. l60G), occasionally fili-
ti
form or moniliform (Figs. ló04, B);
if no snout or muzzle then: body is
less than 9 mm long, cylindrical, an.
(*
tennae elbowed and usually with a
large club, and front tibia with row
{

sometimes practicedto remove surface grease of teeth or ¡ts tip extended into a
that forms alter pinning. These specimens are stout spur (Fig. 2f 2) .
l¡

176 Subclass Pterygota

L'
 mxp
lap (suborder Adephaga). ab, abdomen; abs, abdominal
sternite; an, antenna; cpe, compoun,C eye; cx, coxa;
cxc, coxal cavity; el, elytra (front wing) with
grooves; fm, femur; ge, gena; gs, gular suture; hw,
cpe
hind wing; lap, labial palpus; lb, labrum; md, man-
dible; mss, mesosternum; mts, metasternum; mx,
prn maxilla; mxp, maxillary palpus; nps, notopleural
scl suture; prn, pronotum; Frs, prosternum; scl, scutel-
lum; sp, spiracle; tb, tibia; tr, trochanter; trs, trans-.
fm
vcrse suture; ts, tarsus.

2a(la) Hind coxae flattened and widened '

ab inÍo large plates fhat coyer most of

sp abdomen (Fig. 1698); squatic; oval; '

5mmlongorless .......i.. .:ti

. . (p.lel) HaþIiclae Í,
i,
2b Hind coxae not greatly expanded; "r.l

habitat, size, and shape variable. . . . -l , ,l

....'.J

ge
3a(21¡) Posterior margin of first abdominal
nps
sternite divided by hind coxse
prs (Fig. 1578); antennae almost always
filiform (Fig. f60A); tarsi usually
5-5-5; hind lrochanters large and off'
set toward center of body (Fig. 1578) l

(Suborder Adephaga) , .. , .4

3b Posterior margin of first abdominal

sternite not divided by hin.ì coxae
and extending across abdomen
without interruption (Fig. 15E);
antennae and tarsi variable; hind:1:
trochanters small an¡l not offset .1,,
(Fig. 15E) (Suborder Polyphaga). . . t
,,,,l

; :iÏ 't! :"

4a(3a) Hind legs flattened and fringed with.i i,

Figure 157 General structure of a ground beetle,
Harpalus caliginosus (Carabidae). A, dorsal view;
B, ventral view showing the posterior margin of the
first abdominal sternite divided by the hind coxae
long hairs (Fig. 170C); aquatlc, also if;;'
found on land (some fly to lights). t, 5 fråfi
Í;i I
Hind legs generally not flattened or
fringed with long hairs; usually ter-
restrial

Order Coleoptera
 Transverse suture anterior to hind
coxae (Fig. l57B); longitudinal g
grooves on pronofum present or ab-
senl; s¡ze variable ,. . .. ., .7 ö

7a(ób) Head including eyes as wide or wider

o
than pronotum (Fig. l67X mandibles .@
long and sickle-shaped; clypeus
enlarged laterally beyond basés of '€
antennae; elytra not grooved or '{¿
Figure 158 Ventral view of a beetle in the suborder punctured longitudinally (Subfsmily
Polyphaga showing the undivided posrerior margin Cicindelinae).:.....
of the first abdominal srernite. l, iirst abdominal . (p. lE8) Carabidae (in part)
ü
sternite; cx, coxa; fm, femur; tr, lrochanter.

Head including eyes generally

ð
narrower than pronofum (Fig. tfg);r
û mandibles usually not long and
sickle-shaped; clypeus not enlarged,
as above; elyfra often grooved or
Figure 159 General body shapes. A, spherical; B, punctured longitudinally (Fig. l6EB)
broadly oval; C, oval; D, elongare. (p. 1E8) Carabidøe (in part) ,
,

5a(4a) Compound eyes divided info upper Ea(3b) Tarsal segmenlation S-5-5, 4-4-4,
and lower halves (l-ig. lTlB); black 3-4-4, or 3-3-3. 9 í,
and oval; anlennae vcry short; swim '"J
in circleson water'ssurface i. . . . . . 8b Tarsalsegmentation5-5-4. ..,...49
(p. 192) Gyrinidae U
Compound eyes not divided; color 9a(Ea) Tarsal segmentation actually or
brown, black, or greenish, often with apparently 4-4-4, 3-4-4,or 3-3;3 . . t0
pale yellow margins on elytra and
pronolum; oval; antennae long and Tarsal segmentation 5-5-5. . . . . .. ll
filiform; do not swim in "ircles on
waler's su rface
' ' (p' l9l\ [t'tiçcidqe ll)a(9a) [:rrsal segment:itiorr a,;tuaìll] or
rupp:rrentl¡ {-'l-4,o¡:i,l-,1" .,....62

óa(4b) No fransr,erse suture anlerio,.(o hind l0h -f*rsalscgrnerration,ì-3-3

coxae; 3 or more lonliitudirral
grooves on pronotum (Fii¡. ló(¡);
5.5-7.5 mm long lla(9b) Antennae tameilare (Fig. ló0D) or
. . . . . (p. l$E) Rhysodidae flabettate(Fig.lóOE) ....t2

178 Subclass Pterygota

 r4a(12b) Last 3-4 plafes of lamellate antennae
flattened and can be closed into a ball
(Fig. 160D); tibiae often flattened
and toothed or scalloped
. . . . . (p. 198) Scarabaeìdae (in pafil

14b Last 3-4 plates of lamellate antenn¡e

rounded or flattened but cannot be
closedinto a ball (Fig. 160D) . ... 15
'ì
15a(l4b) Last tarsal segment nearly as ' ong
as olher tarsal segments combined :

(Fie.1628) ......16
15b Last tarsal segment simiilar in length
toothersegments,......l7 .:

Figure 160 Antennae. A, filiform; B, moniliform;

C, serrate; D, lamellate; E, flabellate; F, 3-
Lenglh I'E mm; elytra cover enlire
segmented club; C, elbowed and clubbed; H, lóa(15a)
clavate. abdomen; silvery, gray, brown or
blackish; aquatic or shoreline be-lles ; r ¡
. . (p.202)Dryopìdae (in pal) il i ,

llb Antennae not lamellaf e or

flabellate . . lE l6b Length l5-35 mm; ell'tr¿r leave about
.l abdominal segnrerrts exposed
{Fig. l73A); blacl. ,cl oranl¡e and
l2a(l la) Antennae flabellate l3 black; lerreslrial tret'tllr:s a,ssociated
with carrion
l2b Antennae lamellate. . l4 . . (p. 193) Silph,idae (in part)

l3a( l2a) Stout-bodied; hairy on ventral side;
front tibiae wide, flat, and scalloped
or toothed; western U.S. .
. . . . . (p. l9E) Scarabaeidae fin parl)
l3l) Not as above; firsl 4larsal seg-
ments with paired lobes beneath
(Fig. l62A), hairy projection bc-
tween tarsal claws; widely distributed
. (p.203)Rhipiceridae
l7a(l5b) Pronotum with deep median groove;
elytra with conspicuous longitudinal
grooves (Fig. f76C); head with short
horn on top that curves forward;
mandibles not unusually long; shin-
ing black; 30-40 mm long . .
. (p. f97) Passalidae
l7b Pronotum without median groove;
elytra generally smooth; head usually
without horn; mandibles long, half
the body length or longer in some
males; brown or black; E-40 mm long
. . (p. 197) Lucanldag

Order Coleoptera 179

l8a(l lb) Maxillary palpi long, slender, 22b Middle coxae touching or nearly so;
and equaling or exceeding length of el1'fra u su al ly' para llel-sided,
antennae (F'ig. l72A); ''ind legs somctimes poslerior harlf slightly
curvcd and flatlcned I'or s r irnming nidcr: colrlrvariuble. . .. ..,.,.. 23

1'''l lll :l' i'ìï";l;lj' i;il:ii i',,;;,;;,,;,

2.-Ìrrtl.lh I llr:rtrl gt'ner:rllr corr,le¡lr:d bl'
I ttb Mrrrillirrr ¡ltl¡ri rnrrclr slrorlt'r lluttt ¡rrorrolrrrn irr¡rl nol r i,iihlc l'rr¡nn:¡bove f,it
¿ullrrìn:rc; lrilttl lcgs rtol rnotlilicrl ior t I ig, llt3A); e ¡rislcrnurn of mela-
srrilnrtring; lrnlcntt:¡c r:tri:tlll,.' . . . . l9 t l¡¡rr¿rx with ncarly straright inner
6¡
rnargin (!'ig. lólA); many with yel- (-t
lowish band on ventral side of ab-
l9a(ltlb) l'-l¡ tra usuall¡ covcr less lhi¡lr half the clomen qt
lenglh ol' abdomen (t'ig. l7.l); el] lra . (p.206) Lampyridae
occasionall¡ pu besccn I
. . . . . (p. 194) Staph¡,linidae 23b I'lcad slightly or nol concealed by
pronotum and visible from above;
l9b l.l¡ f ra corer rnorc than hall' or all ol' epislernum of metathorax with "S"
abclome n; ely tral prrbesccncc prcsent curved inner margin (t'ig. lólB); no
orirbsenl ,..,.,.20 yellowish venlral abdominal band . .
Cantharídae

20a(l9b) F,l¡tra leave l-2 abdominal segments eps

el eps
erposed; tip of abdomen sharpl¡'

i':': ::' il'i liÍl; ;)ìi'!,1)'X,'),1;":,

i
mts
20b Notasabovc. .....'\....21
Figure l6l Lateroventral views of the metathorax
to compare the inner margin of the episterna. A;
2la(20b) Abdomcn u'ith 7-8 r'enlral segments firelly (Lampyridae); B, soldier beetle (Can-
visiblc ...,22 tharidae). cx, coxa; el, elytron; eps, episternum; mts,
metaslernum.
2rb Allcl¡lnrcrt n ilh lcss lltan 7 rcntral
scgnrcntsrisible ..,.....2{ 21a(2ltt) Abdomen with 6 venlral segments
visible. . 25

22a(2la) Middle coxae separated from each 24b

othcr; el¡ tra usuall¡ rvith netlike
Abdomen with 5 venfral segments
visible. . .
sculpturing and posterior half
noticeably wider (t ig. 184); of len

::'il:'Ì::':':i: (; ,ô;i 1,,,¡io, 25a(21a) Flattened, oval or broadly oval,

generally blackish beetles
(Fig. l73B); antennae with 3- or 4.
segmented pubescent club; usually

180 Subclass Pterygota

 {)\rr l{} mnr l()n!1: :lrlra ;pur;r'l) r¡r ),lll )ir,l with ¡rl,rlr t' r'r:,frlrir¡atiOn of
rol ¡i,rbcsceltl. . f g: tu res 29
. . (p. l9.l) ,>'ilpltitt øe (in part)

25b Usuall¡ c-r'lindrical; anlennac \ ari- 2t)a(¿llb) ()re¡ ¡,0¡t or all r¡f head con.:aled
able; lcngth usuall¡ l0 n'rrn t¡r lcss; from above b¡ pronofurn . .. .. . .30
elytra pubescent. . ......26
29b Over half or all of heatl visible from
above .
2óa(25b) Middle coxae conical (Fig. l63A) and
prominent; hind coxae Prom¡nenf
and exlend ventrally below abdomen; 30a(29a) Antennae long and usually at least
elylra much broader posteriorll' pro- half the body length, filiform, with
ducing a stout, wedge-shaPed aP- or without basal process on some
outersegmenls (Fig. l80A). . . . . . 31
':::ili ''li.li''i; ;, ,) r,,i,,ii",
30b Antennae short and usually less thsn ,1 ,
26b Middle coxae rounded (Fig. l638) half the body length, shape variable,
and nol prominent; hind coxae nobasalprocesses. ......32
somewhal flattened and not Promi'
nently extending below abdomen;
elytra sometimes slightly broader 3la(30a) Anlennae wif h basal process on
posteriorly but generally with an
elongafed and narrowed appearance
'::i:::i 1':l,i'hliH,Ì,';";;;;;;,
. . . (p. 210\ Cleridae (in part)
3lb Antennae without basal processes . . I :
2'la(24b) Antennae both elbowed and clubbed . . 1p.20E) Ptín¡dae
(l.ig, l60G); elllra short and
squared-off at ends leaving 2 abdom-
32a(30b) Body elongate, elongale-oval, or
' :' :' : :il:: i': :: ::'äi li¿*, Åil,?,;":, cylindrical(F¡g.f59D) ...33
21b Antennae, elytra, and abdominal 32b Body broadly oval and convex to
segments notin above combination; nearly spherical (Figs. 1594, B) . . 36 ,

if antennae are clubbed they are not

alsoelbowed .....28
33a(32a) Antennae very short, clavate or
lamellate with most segments
2Ea(27b) Dense covering of while, gray' or broader than long (Fig. 1808). . . . . .
yellowish hairs on light brown or . . (p.202) Dryopìdøe (in Pal) ,

dark orange body; 3-segmenled

antennal club; second and third tar' 33b Antennae not as sbove . . .34
sal segments lobed; body elongate
-.'
:i :': :i". : :'::'::-ä1 il ;) ; ; ;,:,;;*

Order Coleoptera
34a(33b) Pronotum and elytra sculpfured with
curved ridges giving a slightly
wrinkled appearance (Fig. t80C);
rows of punctures on elytra
....(p.203)Elmidae

34b Pronotum and elyf ra not highly

sculpturedorwrinkled ...35
K
/Å" 6r
35a(34b) Pronotum often covers and encloses
all of head from above; at least last 3
(-
segments of antennae lengthened,
often expanded and pectinate or ser- fr. a
rate; hind coxae wif h grooves for 162 Tarsi modifications. A, paired ventral
Figure êð
receiving femora (Fig. l63D) lobes; B, last tarsal segment nearly as long as oåhers
(p.20E) Anobiidae (in parf) combined; C, ventral lobes; D, third segment
bilobed and fourth hidden inside. &ð,

35b Pronotum does nol enclose buf €¡

usually covers all of head from 37a(29b) Tarsi with I or more ventral lobes
above; antennae nol lengthened or
expanded distally; pronotum with
(Fie. 162C) t,
Æ:
tubercles, hooks, or rasplike teeth 37b Tarsi without ventral lobes, although ,
@
nearfronf..... segments may be bilobed or dilated
(p. 209) Bostrichidae
(Fig. 162D) with hair pads beneath. .

36a(32b) At least last 3 segments of qntennae

lengthened, often expanded and pec-
38a(37a) Only fourth larsal segment lobed
tinale or serrale; fourlh tarsal seg- ventrally; hind femora sometimes en-
menl nol bilobed, tarsi decreasing in larged for jumping; 2-4 mnr long . . .
size distally; hind coxae with grooves (p.201) Helodidae (in part)
for holding femora (Fig. l63D); head
enclosed and concealed by pro- 3Eb Two or more tarsal segments or only I
notum; body oval .
third tarsal segment with prominent
. (p.20E) Anobiidae (in part)
ventral lobes; hind femora not \J
,ae not enlarged; 3-24mm long . .. .....39
36b Last 3 segmenfs of ante¡r
lengthened, antennae weakl) serrate;
t'orrrlh lÍìrsal segnt,rr bilol,ed tnd 3r)a(.ì,'lir) i :irstrl l;¡:gr¡cnts i,4,ì,/itli prominent
larger tlurn third; no ltroo\c, in lrind ,.,.".,
coxae; hcad often coll(:clrled b¡ ']
shelflike anterior expansion ol' pro-
: :,., :,.ìi,. r, n, r,,,,".¡,): ii¿ ; r;;;
notum; body oval to s¡rherical . . . . .
39b Tarsal segment 3 or segrnents 1-3,
. (p. 201) Helodidae (in part)
2-3, or 2-4 with prominent ventral
lobes.. ...40
182 Subclass Pterygota
40a(J9b) ['oslerior corners of pronotum 43a(4fb) Front coxae lie at right angles to
usuallt pointed (Fie. lE2); prothorax main axisof body' ,,.,.,.U
looscl¡ joined to mesothorax;
posterior margin of prosternum 43b Front coxae do not lie at right angles
elongated into spinelike projection to main axis of body. . . . . 45
between front coxae (Fig. lE2A). , . .
. . (p. 204) Elateridae
44a(43a) Tarsi slender, first segment very
40b Pronotum, profhorax, and short; pronotum often s€Parated
posternum not as above from base of elytra except at po¡nt of
. . (P.201) Dsscillidse attachment (F¡g. IEEB); elytra never
truncale. . .
(p. 210) Trogosltidss ;: t,

4la(37b) Front coxae conical, projecting

prominently from coxal cavitY 44b First 3 tarsal segments about same
(Fig. 163A). .....42 size with brush of hair beneath'
fourth segment smaller and without ,

41b Front coxae round (Fig. 1638), oval, brush of hair; pronotum usually not :

or lying at right angles to main axis separated from base of elytra; elytla
of body (Fig. l63C), and usually pro' often truncate and showing tip of ab.
jectingonlyslightly ......43 domen
. (P.212) Nítíd'tlldae
"
ùc fln 45a(43b) Posterior margin of prosternum
elongated into spinelike projection
4Rc between front coxae (Fig. fE2A). . . . .

. (p.2031 Buprestldae :

Figure 163 A-C, front coxae: A, conelike; B, .,.

rounded; C, transverse; D, grooved hind coxa' 45b Posterior margin of prosternum not
elongafedasabove ......46
42a(4la) Body oval, broadening posteriorlY
(Fig. l79D); antennae se rate; dor-
sunr rarely hairy. not sr:i, )'; aqrtatic .lóa(45h) ,\ntennue rvillt lr brupt 2-s,egmented
h ee tlcs (raiel¡' J) clt¡b; ltr:ad na rrovl,s behind
(p. ¿02) I'. ePhenidae ey{'s ., ....(p,.I09)LYctidae

42lt llody oval or broadl¡' or al, nt¡l
broadening posteriorly (Fig. lE5);
anlennae clubbed; dorsum offen
noticeably hairy or scaly; terrestrial
beefles
(p. 207) Dermestidae
,l6h Not with abovr: concrbination of
features . ,
47a(46b) Body very flattened and sides parallel
or body oval; middle coxal cavities
open (Fig. f64D)
. 1p. 2f 3) Cucujidae rin Part)

Order Coleoptera 183

 r(Ð
47b Body {)val, elongale-oval, or 49b Front coxal cavif ics open behind
c¡ lindrical with parallel sides; middle (Fig. l64B) ',......52 Ð
coxal cavities closed (F ig. l6aC) . . 4E

50a(49a) Tarsal claws pecfinate (Iiig. lósAX

c
cxc
*: i:ii:ili ;- e
i; ;;;;;;;":, @
50b Tarsal claws not pectinate; body
usually nof pubescent. . . . . . . ... 51 t
mss
B
e)
ls 5la(5()l;) \nfennac {ilil'orr¡l rrilh lersl sr:gment
üD
Ir. lr¡ng us ¡lrccctlirrl' I rlr,l seErrents
t '¡rrtllilre'tl ; n('\( irr Ilrs{ i¡t.:ral sr:gment
tlilatcd anrl spongl lrcncatl¡
(p.220) Lagriidae
CD
Figure 164 Coxal caviries. A, fronr cavities closed 5lb Antennae filiform, moniliform, or'
behind by the sides ol the prorhorax; B, f'ronr slightly clubbed but last segment
cavities open behind (surrounded by the mesoster- about same size as other segmenls;
num); C, middle caviries closed behind (not touched
next to last tarsal segment not dilated
by the lateral sclerires ol the mesothorax); D, midctle
cavities open behind (touched by the lateral sclerires and spongy beneath
of the ntesolhorax) cxc, coxal caviry; Is, lareral 1p. 2f8) Tenebrionídae
sclerites; mss, mesosternum; mts, metasternì.rrn; prs,
prostefn um .

52a(49b) Lateral margins of pronotum sharp

4Ea(41b'l Antennae w'ith 3-segmented club orflattened. .....53
(rarel), 4-5 segmented) (t-ig. l60F);
body broadly' oval to elongate-oval; 52b Lateral margins of pronotum
pronolum much broader fhan head, rounded . . 56
anterior corners slightly pointed. . . .
. (p. 214) Erotylidae (in parr)
53a(52a) Body elongate and extremely flat . . .
4Eb Anlennae with 4-segmenled club; . (p. 213) Cucujidae (in part)
body elongate and c¡'lindrical with
parallel sides; pronolum equal to or 53b Body not extremely flatlened . . . . 54
slightly wider lhan head, anterior
corners rounded
. (p. lE4) Languriidae 54a(53b) Body humpbacked, as high as or
higher than wide (l'ig. 2048); head
bent downward (Fig. Z04B)i tip of
49a(tìb) l'ronl coxal cavilies closetl behi n d
([ ig. ló44) ...s0 ::i: :: : i :' i." .^,f ':;lfi) ; ;o:,i i i o, U

184 Subclass Pterygota

54b Wilhout above combination ol 58b At least ouler portion or all of
lea I u res s5 antennaefilifornl .,59

55a(54b) Antennae filiform; 2 dents or 59a(5Eb) Head not abruptly narrowed be.
depressions near posterior margin of hind eyes; pronotum widest at middle
pronotum; first segment of hind tarsi and narrows toward front and back
much longer thân anÿ other segment; (Fig. 20lA).
3-20 mm . (P. 22fl) SulPingidøe
. . . . , (p. 222\ Ntelandr¡'ir re (in part)
5,)h tlt atl ahruptll nnrrowcd behind eyes
55h ,'\nlen¡rae nilh lliomincnf to ['rlrnl a sìcnrl er rrct'k (lìills. 205;
3-segtnentecl clulll no denls or 20fi).. ...60
tlepressions near posteriot margin ol
pronolumi first segmenl oI hincl tarsi
nof grcatl¡ lengthened; l-5 mm . . . ' ó0a(59b ) [,ach tarsal clarv split. to base or
. . . . (p. 2l3l CrYPtoPhagidae toothed (Figs. ló58, C); abdomen
with 6 ventral segments
. . . (p. 1123) Meloidse
56a(52b) Two dents or depressions near
posterior margin of Pronotttnr 60b Tarsal claws not split or toothed;
(Fig. 203C); first segment of hind abdomen usually with 5 ventral
tarsi tnttch longer than any other seg- segments. 6l
ment
, . . . . (p. 222) Melandryidse (in part)

56b Nolasabove. ...:.'....57

57a(56b) Next to last segment of hind tarsi
expanded and heart-shaPed or
bilobed, often with dense brush of
hairs ventrally; pronotum widest in
Iront and narrower than elYtra at
back; elongated body
(P.222'l Oedemeridae
57b Not with above combination
t'eaturcs
""""5t1
5Ea(57b) Anlennae serrate, I labellate'
pectinate, or plumose (Figs. l60C' E;
2024) . (P. 221) PYrochroidae
ABC
Figure 165 Tarsal claws. A, pectinate
(Alteculidae); B, split to base (Meloidae); C, toothed
(Meloidae).
6la(ó0b) Length 7-12 mmi eyes usually with
slight notch on margin; hinri coxae
'::: : i:' i':::::
: ï;: åii'!üi,oi,
of
6lb Leng(h under 6 tnm; eyes without a
notch; hind coxae sep:rrated; often
with pronotal horn extending over
head (Fig. 20óA)
. . (p.22Ð Anthicidae

Order ColeoPtera
 62a(10a) Tibia widened and armed with row of
prominent flattened spines on outer
margin (Fig. lEOD); mandibles great_
ly' flaf lened and project forward; first
and f ourth tarsal segments longer
:lii :ï::: lll,1'lå;, ; :,) ì, oi)¡oà)
62b Not wilh above combination of
feafures.
63a(62b) Terminal l-3 segments of
distinctly clubbed, or if
clavale the last 2-3 segments
clubbed ...,....64
63b Anlennae variable but not clubbed, if
clavate then the last 2-3 segments not
distinctl¡,clubbed .......ó9
64a(63a) Antennae elbowed and with a
distinct, large, solid club (Fig. l60G);
elongate and cylindrical body
, (p. 233) Scotytidae (in parr)
64b Antennae not elbowed alf hough l- to
3-segmented terminal club present;
bodyshapevariable .....65
65a(64b) Body broadly ovat and highly convex
il'T : IilÌ :::ii,i:'J, ;; ;;;;":,,;;",
65b Body oval to elongate and
cylindrical; color variable, often
brightlypafterned.,....,66
66a(65b) Head concealed from above; first
larsal segments short; horn
sometimes present on head or pro-
186 Subclass Pterygota
notum; pubescence consists o,f erect
|i'::''l::::i: iii :ii: \iTT',,;;;
6ób Head not concealed; tarsal segments
variable in length; no horn;
pubescence presenl or absenf but
hairs not erect; elongate-ovat or
oval . .
.....63
67a(66b) Antennae clavate (segments 7-ll
antennae enlarged), last 2-3 segments with
anlennae loose club; anterior corners of pro.
are notum not prolonged; tarsal segmen.
tation 4-4-4 or 3-4-4; usually brown
':.'.'::i ',;;;,;;;;;;;"'"
.. :ö.;;
61b Antennae nol clavate buf last 3
segments abruptly clubbecl; anterior
corners and margin of pronotum
may extend forward to partly conceal
head (Figs. 1974, BX farsal segmen-
tat¡on actually or apparently 4-4-4;
often red or yellow markings . . . . 6E
ü
6Ea(67b) Anterior corners of pfono- \-l
tum extended distinctly forward L-,
(Figs. 1974, B); pronotum usually
with 2 longitudinal grooves,
(Fig. 1974) or 2 pits near posterior
'L
margin connecting to longitudinal l:l
grooves extending up to half the
length of pronotum; tarsul segmen-,
tation actually 4-4-4 but sometimes
u
appearing 3-3-3 due to small third ü
segment . ..
. . . . (p. 217) Endomychidøe (in paf)
3 68b Anterior corners of pronotum very
slightly or not extended forward;
pronotum withouf longitudinal
grooves or pits with connecting
grooves although lateral margins
 may be very distinct and groovc'like; 72a(l0b) Elytra short, covering about half of
larsal scgmenlation rctuall¡ 5-5-5 httt abdomen (l'ig. l75A); l.arsi rarely 2-
appearing 4-4-4 due to srnall fourlh
segment
(p. 2f4) ErotYlidae ( np
i:::T:: ô ó¿;ï,;i"p;;;;,
72b Elytra cover abdomen; tarsi never
2-segmented, apparently 3-3-3 but
69a(63b) Anlennae usually clavate; body oval actually 4-4-4
and greatly convex to give a pill
shape (Fig. t79C); head nearly or
completely concealed 73a(72b) Oval to elongate; 2 longitudinal
. . (P' 202) BYrrhidøe grooves on pronotum (Fig. l97A) or
2 Pits near Posterior margin con'
nected to longitudinal grooves ex'Ì
"
69b Anfennae sometimes clavate but
usually filiform, serrate, pectinate' tending uP to half the length uf the
or flabellate; if body oval then nol pronotum; anterior corners of Pro'
greatly convex; head sometimes con- notum extended and partly enclosing
cealed .'..70 nead (FiB. f97A)
....(p.2f7) :;-lr,i:
ìi4$,
;i!'.1''
70a(69b) Body elongafe and parallel-sided or 73h Broadly oval ¡r':;

narrowing posteriorlY; antennae convex abov

'1.

usually much more than half bodY without longitudinal grooves or pits
length (Fig. 207) (rarely shorter than with connecting grooves; anterior
half body length) and inserted into corners of pronotum not extended ..
the notched area of eyes (Fig. 2074); (P.215) Coccinellidae
commonly 3-50 mm long ..
. . . . . (P. 225) Cer'ømbYcidae
74a(rb) No snout or broad muzzle presenti .. -

70b Body usually oval or elongate oval, if antennae elbowed and clubbed ' I

elongate ¡t does not narrow (Fig. l60G); body elongated and

posteriorly; antennae shorter than cylindrical (Fig. 212')i most are
half body length; eyes not notched; l-3mmlong......
usually less than 12 mm leng. . . ' . 7l . . . . . (p. 2331 Scoll'tida¿(in part)

7"tb Snr¡ul (Figs. 2l l'C; :!13; :',14) or

7la(70b) lÌ<¡dt oraì rlt egg.-;lrape , llroitdest llr'arl r'¡lrt¿zlt' ¡rlt:sent (li'-itl. 21lB);
postcriorl¡; ht'ati 'lx¡rlr rrletl¡nto ÎD:enna(.r'ìriir ill',: tiItl lo elongate;
short, broad snotlli elvtra ex¡rosc lip l--l5nrnr ........75
of abdomcn (Fig. 21,1) ' ' ' '
. ' (P.232t Bruchidae

7lb Body not egg-shaped or broadest

posteriorly; no short, broad snout;
tip of abdomen nol usuallY exPosed
. . . . (P. 221\ ChrYsomelidae

Order Coleoptera 187

 75a(74b) Broad, short muzzle present
(t'ig. 2l I [ì); antennae clubbed but
not elborved, rarely unclubbed; el),fra
oflen moltled with while, brown. or
black pubescence . .
. (p.232)
75b Short to long snoul presenf
(Figs. 2llci 2l3i 214)i antennae
clubbed and almosl always elbowed,
or somefimes filiform or moniliform;
:'l::1 ::': ::::':::: llllll'lll ;;
7óa(75b) Snoul
gcne rurll¡ curr ccl
214); anrennae clubbed arrd armost
: li i l' :' : : ::i,ii5;iåi,î),,,, i.,,à))
76b Snout exlends straight forward
(Fig. 2l lC); antennae filiform or
moniliform (Fig. 2llC) . .
. . . (p. 232) Brentidae
SUBORDER ADEPHAGA
Family Rhysodidae
Rhysodid beetles are 5.5-8.0 mm long, dark
reddish brown, and slender-bodied. The anren-
nae are monililorm and the pronotum has three
distinct longitudinal lurrows, The four species
in the U,S, occur under the bark of ash, beech,
elm or pine.
Common Species
Omoglymmius americanus (Laporte) (Fig. 166)-
6-8 mm; shiny dark reddish brown; elytral grooves
with large punctìJres; eastern l/2 U.S.
188 Subclass Pterygota
@
Ð
Anthribidae Ð
Ð
ø.
ü
Figrrrc 166 Ottro¡¡l,t,tnnrius antericanus tRhyso- @
d ìd rr')
(Ð
(1.-i1is. 2lJ;
Family Carabidae G ror¡nd
Beetles and Tiger -Beeiles
(Ð
This larnily of predaceous beetles is one of the ü
largest in the order with nearly 40,000 species
worldwide and about 3,100 in the U.S. and
Canada. The tiger beetles are considered a sub-
L
family by carabid taxonomists alrhough many :u,
texts classify them as a separate family, the
Cicindelidae, partly due to custom. Another \¡l'
group, the genus Omophron, has sometimes
been placed in a separate family, Omophro- U
nidae.
Tiger beetles are long-legged and run and i)
fly rapidly. They frequenrly fly low to the
ground for varying distances. The adults are
often colorful, sometimes iridescent, and
always have distinct, sharp mandibles. The
"S"-shaped larva has a hump with curved
hooks on the fifth abdominal segment. Larvae
are predaceous and occur in vertical tunnels
beneath the ground. The larva's head protrudes ,
from the circular tunnel opening to grasp a
passing insect. Sandy beaches with scattered
vegetation, stream banks, and sandy trails are
common habitats for adults and larvae. Some
also occur along forest edges, on clay soils, and
rn rotten stumps. The genus Ornu¡; contains
black, wingless, nocturnal species thart occur in
the Far West.
 Ground beetles are usually dark, shiny,
and somewhat llattened although some are
brilliant and/or iridescent. The legs are long
and often the mandibles are well developed.
The elytra of many are longitudinallv grooved
and the trochanters of sonle are large and
kiclncy'-shapcd lt4tlst spccics arc noctì.lrnal and
occur ur.tclcr clcbris, wtlocl , and rocks dtrring the
clay. Aclults arc trsualÌy scavcngcrs and larvae
are predaccous Sonre adults ieed on insects,
slugs, snails, or climb trees in search of cater-
pillars. One group eats seeds, particularly of
grasses. The elongate larvae have ten distinct
segments and taper toward each end. The head
and mandibles are large, and the ninth ab-
dominal segment has a pair of cerci and an anal
tube. Adults of the bombardier beetles (genus
Brachinus, having the tips ol the elytra trun-
cate), have an unusual and audible chemical
defense mechanism. A chemical reaction occurs
in glands near the tip of the abdomen; the
temperatìrrc rcaclles 100 "C and t he chemical is
sprayed in a water vapor cloud at the aggressor'
Common Species
Tiger Beetles ' segments brownish yellow; each elytron with 2 yel'
Cicindela oregona LeConte-12-14 mm; green or
blue; white or dull yellow markingsi long hairs near
inner edge of eyes; labrum short; punctate elytra,
edges sometimes faintly toothed; western North
America (edges of lakes and streams).
Cicindela punctulata Olivier-10-13 mm; brownish
black or slightly green with bronze sheen; row of
small greenish or bluish depressions on elytra; small
white markings present or absent along elytral mar-
gins; North America except Far West (attracted to
lights; emits applelike odor)'
Cicindela repanda Dejean (Fig. l67A)-12-14 mm;
bronze or brownish with greenish cast; white mark-
ings; short labrum; North America east of Rocky
Mts., British Columbia (riverbanks)'
Cicindela .texgutta[o Fabricius-12-14 mm; brilliant
green, blue-green, or violet; usually 9 spots on
elytra, somctir¡es 6-10 or spots absent; elytra punc-
tate; easrern North America (meadows, paths in
open woods).
Figure 167 Tiger beetles (Carabidae). A, Cicindela
repanda; B, Cicindela lranquebørica'
Cicindela tranquebarica Herbst (Fig' l678)-12'
l4 mm; brownlsh, greenish, dark blue, or black;
obtique white markings; throughout North America'
Ground Beetles
Bembidion patruele Dejean (Fie. l6tìA)-3-5 mm;
black with greenish luster; head, pronotum' and legs
reddish brown; elytra iridescent with orange or dull
yellow patches; most of U.S. except southernmost
states, Canada.
B e m b i d i o n q u a d r i m ac u I a t u m (Linnaeus)-2-4 mm ;
black with bronze tinge; legs and basal antennal
low spots; east of Rocky Mts. south to TX.
Calathus ruficoltis Dejean-8-10 mm; pale red; pro-
notum brighter redi elytra finely striated; Pacific
coast states.
Calosoma calidum (Fabricius) (Fig. l688)' Fiery
hunter-20-27 mm; black, sometimes green mar'
gins; elytra deeply grooved, 6 rows red or gold punc-
tures ; throughout North America.
Calosoma scrutator Fabricius. Fiery searcher-
25-35 mm; black with violet tinge; head and protho'
rax with gold, green, or red margins; elytra irides-
cent greeñ or blue-green, purple margins, grooved
and punctate; throughout U.S., southern Canada
(sometimes in trees).
Chloenius sericeus Forster-12-17 mnt; bright green;
antennae and legs pale brownish yellow' antennae
darker torvard tip; fine yellowish pubescence on
elytra; throughout North America (lake and stream
margins).
Order Coleoptera
 ü
t
Ð
Ð
(Ð

ü
I

t¿
@
O
€Ð

tl
ú.

(;

Figure 168 Ground beetles (Carabidae). A, Bembidion patruele; B, fiery hunter, Calosorna
caliclum; C, Golerita janus; D, Harpalus pensylvanicus; E, Lebia arriventris; F, Omophron
tesscl I a t u nt, C, S tenolophus com ma.
Chlaenius tricolor Dejean-10-13 mm; head and
pronotum green; antennae and legs brownish
orange; elytra bluish black, deep and narrow
grooves, punctures; covered with line hairs;
throughout U.S., southern Canada,
Galerita jønus Fabricius (Fig. l68C)-16-23 mm;
pronotum, legs, and base of antennaê reddish
brown; elytra bluish black; short pale yc, rw hairs
on bodl'l centrll anil castern lJ li, C)nt;u ) (ccnres
to lìghr<t
[{arpalus caligittost,r (I;abricrLrs) iì:rg, Ì:r ¡¡\)--.]l-
26 mnr; black; antcnnae and tarsi reddlsll brou.n;
ely'tra deeplv gioorcrJ; throughour North Atnerica
flarpalus pens.t'lvanicu.ç De Geer (Fig. l63D)-12-
l5 mm;shiny black; convex; reddish llrowrì ventral-
ly; elytra deeply grooved with few to many punc-
tures; throughout North America (especially cotton
lields, pastures).
190 Subclass Pterygota
er
:J
Lebia atriventrrs Say (Fig. l68E)-6-10 mm; head
,ü
and pronotum reddish yellow; elytra dark blue,
dense punctures, shallow grooves; throughout North
America (climbs trees). l. grandis Hentz similar but
larger with deep striations, eastern U.S. and south- {_
ern Canada.
Omophron tessellatum Say (Fig. l68F)-6-7 mm;
pale brownish ycllow; green rnetallir: rnarkinl¡s on
ìread, ¡ronc'lLrrn, cll'tra: centrai U.Si.:rncl Clarra,la (in ,J
rn0,sis:lr-',1 tìtítr \'3ter, llotÌr \\,iì"ft ì:\'(r sÍìnd t.o force
oLrt,'-1i". 1o lr!rhÌ:ì r
.i
Pusit¡ttchts rie¡tr t.;:;us [jabnciLrs,-25-:() lnrn; flat;
shiny c,r clull black; pronotuûì and eìytra with blue
nralgins; cll iia smooth, ridge near trase; east of
Rocky lv'{ts. u
Scarites subterraneus Fabricius-14-21 mm; shiny
black; mandibles large; head with 2 large depres-
sions or parallel lines; elytra grooved and separated
by convex ridges; U.S. except Northwest, south to
TX, southern Ontario.
Ç
{
l*-,

,\t
li.
.:-\ .

,:,
'À:
i::t
tjj
Stenolophus comtno (Fabricius) (Fig. l68G)-6-7
n-rm; head blackl pronotum ancl el¡'tra reddish
broun or 1cììoriish hrorvnìblack patch on pron.tì"rrn
and black \trpc on rnnel mar¿ilr ol c:tch clltron,
elytral stript'bìunt unteriorlyl ccntral ancl souther¡r
U.S (flies to lrghts) S leconlei Chaudoir, seeclcorn
beetle, similar, 5-6 mm, dark color patch on elytra
pointed anteriorly.
Family Haliplidae-
Crawling Water Beetles
These small (2-6 mm long), broadly oval beetles
are common in weedy areas of ponds and
streams. They are brownish or reddish yellow
with dark spots. The extremely long, flat and
broad hind coxae cover much of the abdomen
(Fie. 1698). Adults and larvae feed on algae.
Some adults in the genus Peltodytes have two
black spots on the posterior margin of the pro-
notum whereas those in the genus Haliplus lack
these two spots.
Figure ló9 Crawling water beetles t Ialiplidae).
A,, Haliplus lriopsi.s;8, v,-'nlral view o a l'aìiplid
shosin! cnl,rr¡:,:d :ô\ilc c\, co ',ì
Common Species
Haliplus i¡nmaculicollis Harrrs-'l 5'.ì (' nrrn; red-
dish yellorv rvith black spots; 2 imptessions on
posterior margin of pronotum; deep grooves on pro-
sternum; most of North America.
Haliplus triopsis Say (Fig. l69A)-3-4 mm; dull
yellow; anterior margin of pronotum with black
patch; black spots on elytra; east of Rocky Mts.,
NM.
Pe It s (Say)--3-4 mm ; dull
o rl.v t e.ç cl u o d ec i m p u nc I a t u
yeìlow; l2 hlack spots on elytra; eastern l/2 North
¡\rncrrca [' adenÍuluç LeC-onte sinrilar but with
blrrck bctrvcerr e1'es P ¡tedunrulutus Biatchley pale
1'ellow and black, legs yellow except h jnd femora en-
tirely black, NY to IA and south to CA and TX.
Family Dytiscidae-
Predaceous Diving Beetles
These oval, predaceous beetles occur in most
aquatic habitats. Many are yellowish brown to
nearly black and they often have dark yellow
patterns or the elytral and pronotal margins are
yellowish brown. Dytiscids resemble the Hy-
drophilidae (water scavenger beetles) but are
more streamlined and have short maxillary
palpi, long liliform antennae, and move their
hind legs simultaneously rather than alternately
when swimming. Dytiscids migrate from pond
to pond by flying and sometimes appear at
lights. Air bubbles for breathing are stored be-
neath the elytra. Larvae, sometimes called wa-
ter tigers, have a large head and long mandi-
bles, two tarsal claws, eight abdominal seg-
ments that form a body tapering at both ends,
and two tufted cerci on the tubular tip of the
abdomen. Larvae are predaceous on aquatic in-
vertebrates, fish, and tadpoles.
Common Species
Acitius semisulcatus Aubé (Fig. l70A)-12-14 mm;
broadly oval; brownish yellow above, black below;
dark "M"-shaped marking on top of head; 2 cross-
lines on pronotum; sides and tip of abdomen yellow-
ish; most of U.S.
¡1,eohuç (ii;int?Erolus (Crorch) (lrig l70ll)-7.5-
Ç li ¡lnr, rr, ¡ii and lìronottì nl r,'clrli';ir vellow; pro-
f .)tuÌìr ì,', Lrìr .' blrcÌ crori<lì¿Lrì(i.; ,:l¡ tr a b'ownish
Ir ì,.r., ,'i .-lt .r irlr j-,1 irl¡t 'n lL'nrlj ,-drn¿rl stripes;
tltt orl¡¡ì1,r¡1 ì'. r. t[ ¡\nler ic¡-,
(') btl;!t,¡ /rrnltrrolalus (Say)--lt)-33 ntrn; brown with
grecr.rish tinge; pronotum and elytra with yellow
margins; pronotum sometimes with 5 grooves (9);
most of U.S.
Order Coleoptera

Figure
.l70 Predaceous diving beetles (Dytiscidae).
A, Acilius setni.çulcatus; 8, Agahus disinÍegratu.s,
C, leg oi a d¡,trscid.
Dytist'tt.s fas< ivcntris Say-25-28 mm; greenish
black; pronotunl ancl clyrra rvith yellow margins; ba-
sal 2/3 each elvtron with I 0 deep grooves ( ç ); east-
ern I /2 North America.
Hydroporus undulatus Say-4.0-4,5 mm; very con-
vex; pale yellorv to yellowish brown; pronotum with
anterior and posrerior margins black; elytra blackish
and with irregular yellowish patches; hind coxal cav-
ities rouching; easrern I /2 U.S., southeastern
Canada.
Family Gyrinidae-
Whirligig Beetles
Mcmbcrs oi rhis aquatic iamily arc oval, flat-
tenctj, and nlack or clark metallic green. Adults
are readily recognized by nvo characteristics;
192 Subclass Pterygota
(l) each compound eye is widely separated
into an upper half (above the water) and lower
half (below rhe warer) (Fig. lTlB); and
(2) they swim together in circular movements
on the water's surface, Adults also swin¡ under-
water and fly at night. Adults ancl larvae are
predaceous. The genus Gyrinus contains small
species from 3.5-8.0 mm long.
l,¡t
l7l
Ý
Figure Whirligig beetles (Cyrinidae). A, Di-
neulus americanus; B, divided compound eye of a
t"
gyrinid.
Common Species
ü
Dineu[us arnericanus (Fabricius) (Fig. lTlA)-10-
l2 mm; shiny black with bronze sheen; outer margin
\-j
of elytra curved inward in a weak "S" near apex;
eastern U.S., southeastern Canada. \J
f
Ditteutus assimilis (Kirby)-10-l I mm; shiny black t
with bronze sheen; legs brownish yellow; most of
North America.
Dineutus c¡l¡atus (Forsberg)-12,14 mm; black; pro- I
notum and elytra with diffuse, bronzed, curved
stripe on sides; legs dark brown; eastern U.S. south
to FL, southwest to OK, southeastern Canada (small
streams).
Dineutus discolor Rubd-l l-13 mm; black on top;
brown, straw-colored, or rreddish below; distinctly (*
narrowed toward head producing a triangular shape;
eastern l/2U.5., southern Canada. (
Gyrinus borealis Aubé-6.5-7.5 mm; shinl, black;
el;-tral margins bronzed; undersidc evenly dark
brown to black; most of U.S., Canada. G. maculi-
venîris LeConte similar, underside with light and
dark blotches.
al
SUBORDER POLYPHAGA
Family Hydrophilidae-
Water Scavenger Beetles
Most members of this famill' are aquatic
although one sublamily is prinrarily terrestrial.
They lrcqucnt qLriet rvater rvith abundant vege-
tation and nlovc their hind legs alternately
when swinrrnrng. lv{any species lly to lights at
night. Hydrophilids are typically black or
brown, or occasionally dull green or yellowish.
They have long, 4-segmented maxillary palpi
and short, clubbed antennae, A few common
and large species contain a long, backward-
pointing spine on the metasternum. Adults feed
chiefly on dead vegetation or are omnivorous,
but the larvae are predators. Larvae are yellow-
ish brown or gray, elongated, generally tapered
at both ends, and have a single tarsal claw.

Comnron Spccies
Berosus stri¡itus (Say)-4-5 mm, conve\; greenish
yello*'; head black; pronotum u'ith 2 black mcdran
stripes and 2 spots; el¡'tra with scattered indistinct
spots; elytra rvith Iongitudinal grooves containing
fine punctures, ridges between grooves llat and
coarsely punctured; most of U.S. except Southeast, Figure 172 Water scavenger beetles (Hydro-
southern Canada. B. panlherinus LeCo¡te similar A, Hydrophilus triangulø¡¡s and a flat-
philidae).
tened hind leg; B, Sphaeridium scarabaeoides.
but pronotum with paired spots and elytra with
l0 distinct spots.
Tropisternus lateralis (Fabricius)-8.5-9.0 mm; con-
Enochrus ochraceus (Melsheimer)-3.5-4.0 mm; vex; shiny greenish or bronze-black; margins of
shiny pale yellowish brown; pale spot in front of head, prothorax, and elytra yellow; legs yellowish,
each eye; pronotum and elytra with pale margins, femora black at base; underside black; throughout
many punctures; eastern U.S., southeastern Canada. North America.
Hydrophilus triangularis Say (Fig. l72A)-34-
37 mm; shiny black with greenish tinge; abdomen
with triangular yellowish or pale reddish lateral Family Silphidae- Carrion Beetles
markings; prominent central ridge on ventral part of These beetles are moderately large, usually
thorax; throughout North America (flies to lights). black with orange, red, or yellow markings,
and have clubbed antennae. Adults and larvae
Sphaeridi u m sca ra baeoides (Linnaeus) (Fig. I 728)-

5.5-7.0 mm; shiny black; elytra with faint reddish
spot near base and apical I /4 laint yellorvish;
throughout North America (terrestrial; in cow
manure).
feed on decomposing animal matter and the as-
sociated insect larvae. Species of the genus
Nicrophoru.s excavate beneath dead rodents
and snakes and the young develop on the par-
tially buried carrion. Nicrophorus' beetles are

Order Coleoptera 93
 trt
I
elongatcd, usually red and bla,:k, and have
short, trunc¿rtccJ elltra Silphu bectles, the other
!ìomrnon gr'nus, arc broadl¡ oral, I'lattcncd,
Nicrophorus lotnentosus Weber-15-20 mm; shiny
black; elytra short with 2 orange-red bands; pro-
nottrm with dense ycllow pubescence; eastern l/2
t
and the pronotunl is very large
U.S., sot¡theastern Canada.
0
Silpha americana Linnaeus (Fig. l73B)-15-25 mm;
Common Species pronotum yellow with black or brown central patch;
elytra brownish with 3 indistinct, irregularly
e
Nicrophorus americ'anus (Olivier)-27-35 mm; shiny
black; elytra short, 2 broad orange-red bands on branched, raised ribs on each elytron; east of Rocky
Mts.
cì
each; club ol anlennac, top of head, and pronotum
Ã:
orange-red; east ol Rocky Mts, N, marginatus
Fabricius similar, 20-21 mm, pronotum black with Silpha lapponlca Herbst-9-13 mm; dull black; head
lighter margins, most of U.S., but primarily western
I /2, southern Canada.
and prothorax usually with dense yellowish pubes-
cence; Rocky Mts. to Midwest. d
Nicrophorus r,¡rbicollis Say (Fig. l73A)-20-25 mm; Silpha ramosa Say-12-18 mm; velvety black, each @
shiny blackish; orange-red or yellowish patch on ba- elytron with 3 irregularly branched longitudinal
sal l/3 and apical l/4 of elytra; last 3 antennal seg- ridges; pronotum finely punctate; western North @
ments reddish brown; eastern l/2 North America America.
south to FL.
Family Scaphidiidae-
e
Shining Fungus Beetles @
These small (2-7 mm long), oval, convex beetles
have a large pronotum and are usually shiny It-
black; a few have red markings. The truncated ('
elytra expose the pointed tip of the abdomen.
The beetles feed on fungi and occur in decaying
leaves and wood and under loose bark.
TT
\d
Common Species
Eubaeocera apicalis LeConte-1,4 mm; shiny black; (J
antennae, legs, and tips of elytra and abdomen dark
brown; each elytron with deep line along base and ,

inner margin; eastern l/2U.5.

Scaphidium quadriguttatum Say (Fig. l73C)-

3.-5-4.5 mm; shiny black; each elytron with reddish
or ¡'elìorvish spot in basal l¡'3 anC a¡rex, or only
basirÌ s¡r¡r1 ,'¡r * itlrort spots; earstcnt 1/:l'J.S.

{ F t¡-,"'
j{i' I' ìilr ilrylinie! ;r'r. .'
ì¡. Flt :: r.¡ ijì ,1{j f l;i) S;

\ This lanrilv ir: i'er), larg,: ',v tl: <:v'er 26,000

/ specii:s u,orldwide and about 2,900 in North
America. The very short elytra, slender body,
Figure 173 A and B, carrion beetles (Silphidae): and the habit of turning the tip of ttre abdomen
A, Nic'rophorus orbicollis;8, Silpha umericana C, a
up and over the body when running usually are
shining fungus beetle, Scapir idiurn quodriguttaturì1,
reliable identification characteristir:s for rove
beetles. The crossed mandibles are prominent
and some species have hairy bodies. Rove bee-

194 Subclass Pterygota

 ì
I

tles are predators and scavengers frequenting
carrion, dung, fungi, and decomposing plant
material; a lew are parasites of other insects. A
number of species live in bird and mammal
nests and others occur as guests or predators
in ant and termite nests. Species in the genus
Diaulola occur in tide pools along the Califor-
nia coastline.
Common Species
Pelecomalium testqceum Mannerheim (Fig. lTaA)-
3.5-5.0 mm; head and abdomen black; pronotum,
elytra, and sides brownish ycllow; punctures sparse,
contain short setae; Far West.
Philonthus t'.)'tniltcnni.s (F:abrì,:ius)-- )-ì-s trlrr'l;
shini'bllick;!'lvtr.ì rnctlrllic birL' grcerl ,,¡ fil tlle:
iine puncttrrcs ticti:c on cly'trll , : lì.ìt sc (rr ahcl:>i¡rcn;
throughout North Anler ica,
Philonthus lomatus Erichson (Fig. l74B)-6.5-
8.0 mm; shiny black with reddish tinge, although
punctures and elytra occasionally brownish yellow;
legs yellowish; most of U.S., southern Canada.
Scopaeus exignus Erichson (Fig. lTaC)--2-3 ¡rm;
black with reddish tinge; antennae itnd legs pale
yellow; pronotum dark yellowish; ab,Jomen black-
ish, paler at tip; eastern U.S. S. concavus Hatch
similar but has prominent "U"-shaped depression
on 5th abdominal sternite (o); western U.S.
Staphylinus cinnamopterus Gravenhorst-12-
14 mm; shiny dark brown; elytra brownish red; pro-
notum and elytra with coarse, dense l)unctures and
few hairs; abdomen black with reddish tip; most of
Ll.S e\c.tlt Pacilic Northq est, southeastern
Canrrl:,
Stu¡t,!t'it,/ iriri(r//o.çl/s ((ìr''rltr lo:st)--l!)-l:6 mm;
brown ri ith reddish brown spcts; tlc,t ;um of abdo-

I"igirre 174 Rove beetles (Staphylinidae). A, Pelecomalium testaceum; B, Philonthus

lomatus; C, Scopaeus exiguus; D, Slenus comma;E, Tachyporus jocosus'

Order Coleoptera 19q

 i.,
men pale broÈn and spotted;elytra w'ith d()nse punc
€l
tures;eastern U.S , southeastern Canada.
6ir
Staphylinus tna.rillosus (Linnaeus). Hairy rove
beetle-l0-21 mnr; shiny black; el1'tra with band ol
,Ð
yellowish gra)/ or yellowish brown hairs; abdominal
segments 2---l or 2-.1 with dense cover of yellowish Ð
gray or ycllou'ish brown hairs; throughout North
Anlcrica. q|
S(enu.s t'o¡n¡nr¿ L.eClonte (Fig. l74D)-3.5-5 0 nlm; í.rj
shiny black; tarsi ancl most of antennae dark brown;
eycs llrge; recl or orartge spot ncar middlc of each
elytron; throuBhout North America Ç)
Tqchinus,firnhriatus Gravenhorst-7-9 nlm; broad; Ø
head and pronotum shiny black; elytra reddish
brown with dark tips, as broad as long; most of
North America.
(t
Tachyporus jocosus Say (Fig. l74E)-3-4 mm; shiny @
black with reddish tinge; pronotum, legs, and elytra
reddish yellow or yellowish brown; antennae dull w
yellow with dark tips;throughout North America.
Ý.
\i-

Family Pselaphidae- Figure 175 A, a shortwinged mold beetle,

Shortwinged Mold Beetles
-fhis iamill ol short-rvinged beetles resembles
thc rove bcetlcs (Staphylinid¿ìe) but the mem-
be rs have a widcr abdomen and a narrower tho-
rax and head. All are 5.5 mm or less ih length.
The species occur under stones, loose bark,
moss, forest litter, and in ant nests. They feed
on mold, mites, and sometimes ant larvae.
Those in ant nests secrete substances attractive
to ants.
Common Species
Cylindrarctus longipalpis (LeConte)-2 mm; red-
dish brown with paler abdominal margins; palpi
long with enlarged llattened tips; stiff hairs; most of
North America.
Tmesiphorus costalis (Pselaphidae); B and C, hister
beerles (Histeridae): B, Platysoma carolinum;
C, Sa ¡tr i n us pe n nsy lva n icus.
Family H isteridae- Hister Beetles
This family contains small (0.5-10.0 mm long),
typically shiny black or sometimes bronze or
green beetles. The pronotum is very large,
sometimes partially covering the small head,
and the shortened, truncated elytra leave one or
two abdominal segments exposed. Most species
are predaceous on other insects but many are
scavengers and occur in carrion and dung.
Some are extremely flattened and occur under
loose bark; others infest ant nests.

Tmesiphorus costalis LeConte (Fig, l75A)-3.0- Common Species u

3.5 mm; shiny dark brown to black; short yellowish Hister abbrevløfus Fabricius-3.5-5.5 mm; shiny
hairs pressed against body; clubbed antennae over black; 2 grooves parallel pronotal margin, outer (
l/2body length (o); most of U,S. (sometimes in ant groove shorter; many deep grooves on elytra; most
nests). of U.S

196 Suþclass Pterygota

Hololepto quadriclenÍotu (Fabrìcius)-7-10 mnl;
shiny black; rectangulat shapr.d; el1'tra riith onlv l
corrr¡rlctc gì()o\c irrtd I incornplclc grottrc Ilcar ottter
mlrgin; abijonrcn coarscly punctate; southern U.S

Plat.ysonto carolinutn (Paykuli) (Fig. l75B)-

3-4 mm; very'tìattened; shiny black; 5 shallow
grooves on elytra, inner 2 onl¡' on apical l/2; most
of U.S (under bark)

Saprinus ¡tennst,lvanicrr.s (Paykull) (Fig. l75C)-

4-5 mm; metallic green or bronze-green; apical
l/2 elytra with coarse punctures, elytral groove
closest to ìnner margrn is arched;east of Rocky Mts.
Euspilotus assintilis (Paykull) similar but is black,
eastern l/2 U.S , southeastern Canada.
Spilodiscus hiplagiatus (LeConte)-5.0-6.5 mm;
black rvitlr 2 red elytral patches; eastern U S., south-
eastern Ca nada

Family Lucanidae-Stag Beetles

Stag beetles have long mandibles and lamellate
antennae whose terminal segments cannot be
closed as can those of the similar scarab beetles
(Scarabaeidae). Lucanids occur chiefly in
wooded areas and especially in decaying logs.
Some adults fly to lights. The larvae, which are
similar to scarab white grubs, leed on decaying Figure 176 A and B, stag beetles (Lucanidae):
wood.
A, giant stag beetle, Lucanus elaphus; B, Pseu'
dolucanus capreolus. C, a bess beetle, Popilius dis'
junctus (Passalidae).
Common Species
Dorcus parallelus Say l5-26 mm; dark brown to
blackish; head and pronotum shiny; mandibles with Family Passalidae- Bess Beetles
large central tooth; elytra deeply grooved and punc- The passalids are shiny, blackish beetles with
tured; eastern l/2 U.S., southeastern Canada. deeply grooved elytra and a horn on the head.
These beetles are gregarious and both adults
Lucanus elaphus Fabricius (Fig. l76A). Giant stag
beetle-28-40 mm, excluding mandibles; shiny red- and larvae occur together in galleries inside de'
dish brown; antennae and legs blackish; head wider caying logs. One species, Popilius disiunctus
than pronotum; mandibles extremely large, (llliger) (Fie. l76C), is 30-35 mm long and oc-
pronged, about 12-15 mm long (d); NC and VA curs in the eastern l/2 of the U.S.; the t\ryo re-
west to IL, IN, and OK.
maining North American species (genus Pøs-
Pla tycerus depressusLeConte- I 3 mm ; black ; west solus) are limited to southern Texas.
ern U.S., southern Canada (in dead aspen)

Pseudolucanus capreolus (Linnaeus) (Fig. l768)- Family Scarabaeidae-

22-35 mm; shiny, dark reddish brown; mandibles
with I tooth on inner edge (çr); head equal or
broader than pronotum; elytra smooth; eastern l/2
U.S., southeastern Canada.
Scarab Beetles
Scarab beetles form a very large f'amily (20,000
species worldwide and about 1,380 in North

Order Colooptera
 ú

America) of robust, convex beetles with lamel- notum granular, projects anleriorly over head in
e
late antennae (Fig. ló0D). The body sizes range scooplike fashion (ct); elytra grooved; throughout (Ð
North America.
from 2-70 mm in length in North America,
some are metallic, and others ¿re very hairy
ventrally. Scarabs are generally nocturnal and
Subfamily Aphodiinae-Aphodian Dung Bee- I
tles; antennae 9-segmented, 2 spurs at tip of
frequently attracted to lights. This family may tibia; under dung and debris. Ð
be divided into two feeding groups: (l) both
larvae and adults feed on carrion, dung, skin, Aphodius distinc'tus (Müller) (Fig. l77C)-5.0-
@
and leathers; (2) adults feed on leaves and 6.5 mm; shiny; head and pronotum black; elytra yel- (B
flowers and larvae feed on roots, sap, and de- lowish with.black spots and fine grooves; through'
caying wood. Many species in the latter group out North Ameríca.
(¿
are important agricultural pests. The stout lar-
vae have a curved body that is often wrinkled
and the thoracic legs are well
Aphodius fimetarius (Linnaeus)-6-8 mm; shiny
black; anterior corners pronotum reddish yellow; g
developed elytra reddish; head with 3 tubercles (d); through-
(Fie. l78D). out North America. ::e
The genus Phyllophaga contains about
l3l species o1' .lune (or May) beetles. The adults Subfamily Geotrupinae-Earthboring Dung ql
fly noisily at night and feed on leaves ol many Beetles; I 1-segmented antennae with 3-seg-
kinds of trees. The larvae, called *'hite grubs mented clubs; front tibiae very broad and scal' 6¡
(Fig. l78D), cause damage to plant roots and loped or toothed; under dung.
are very common in gardens and under sod' Eucanlhus lazarus (Fabricius)-6-12 mnr; light or u;
The skin beetles, subfamily Troginae, have dark brown; pronotum with large central swelling;
been classified in some texts as a family,
elytra with large deep grooves; east of Rocky Mts', t'
Trogidae. ^2.
Geotrupes splendidus (Fabricius) (Fig. l77D)- ' ,.!'Ð
Common Species l2-18 mm; broadly oval; metallic grecn, purple, or
bronze; hcacl strrface rough, ccntral tutrert:le; ç:astern ü
ì /l U S , sorrtlri'as,(er ¡l (-anacla
Subfanrilt Scr¡rnbl¡einac'-Durtg Becll s ancl )
1'urnblcLrtrgs; lrincl lcgs illser-t fl('scl t(, tip ol' !ilrbl'urnilr 'l io¡¡inrtc-'_Skirl lìcc tli:s: clors¿tl sur-
abclomcn thalt tt-r rnicldle legs, llind tibra rvith f ace r cr'5' r'or.¡gh, coltvcx, gra','ish b'o,,'n ' in and
I spur at tip; lound under dung and carrion; near old and dri carrion, skin, leatht:rs, nests;
tumblebugs form a piece of dung into a ball, rnost spccics in genus lrox.
roll it rvith hind legs to a suitablc site, bury it, Trox ¡nonuc'h¿r.ç Herbst (Fig. l77E)-12-16 mm;
then lay eggs in it. dark brorvn to blackish; head with 2 tubercles; pro-
notum ridged and with tubercles; elytra 'uith large,
round, widely separated tubercles in rows; most of
Canthon pilularius (I-innaeus) (Fig l77A)- I l-
l9 mm; black wirh bluish, greenish or coppery tinge; U.S. 'J
pronotum and elytra w'ith dense granules; most ol ft
US. Subfamily Melolonthinae-June (or lvlay) Bee'
tles, Chalers; tarsal claws of hind legs equal in l'¡
Copri,s .[ricutor Fabricius (Fig. l77B)-13-18 mm; size and toothed, or tips of hind tibiae lack
black; lop ol hc-acl \\ith horn ((') or tuberclc (? ); spurs. Hopiiø spp. usually 6-9 mm long, some.
pronotrrn coar',clr and dcnse'ly Jrurìctatc, nredian
groove prescnti I grc'roves on each ely'tron; eastern rvhat tlattened, silvery scaìes velttrirlly, hind
legs with I claw; Serica spp. shaped like small
l/2 North America.
June beetles (Phyllophaga) but elytral grooves
U
Onthophagus hecate Panzer-5-9 nlm; black with
purplish tinge and scattered, short gray hairs; pro-
evenly spaced in former. U
q,r'
198 Subclass Pterygota
(:t
 e
I
Figure 177 Scarab beetles (Sca'rabaeidae). A, a tumblebug, Canthon pilularius B-D"Jun¡¡
beãrles: B, Copris fricotor; ò, Aphodius tlistinctus; D, Geotrupes splendidus. E, a skin
lle, Trox monachus.
Diptotaxis lenebrosa Fall-7.5-10 mm; black; elytra
with small markings connected in netlike pattern;
western U.S., southwestern Canada.
Macrodactylus subspinosus (Fabricius) (Fig' l78A)'
Rose chafer-8-10 mm;slender; reddish brown with
dull yellou'hairs or scales; legs very Iong, tarsi as
long as lemora and tibiae combined; throughout
U.S
Phyttophaga ephilida (Say)-14-18 mm; shiny
brownish yellow, head and pronotum darker; front
of head not notched when viewed from above; chief-
ly southern Ll.S.
Phyttophaga fervida (Fabricius) (Fig. l78B)-19-
20 mm; shiny, dark reddish brown to reddish black;
front of head notched when viewed from above; pro-
notum widest at middle, punctured; elytra weakly
bee-
punctate,
-P. surface roughens posteriorly; most of U.S.
errans(LeConte) similar, l5-19 mm, Far West.
Phyttophaga lusca (Froelich) (Fie. l78C)-17'
24-mm; shiny,'dark reddish brown to reddish black;
front of head broadty but shallowly notched \ryhen
viewed from above; pronotum widest posteriorly;
throughout U.S., southern Canada.
P hy tt o p ha sa ru goso (Melsheimer)- I 8-23 mm' shiny
,.d¿irñ brõwn or reddish black; body shape similar
to P. Jusca; front of head deeply notched rthen
viewed from above; pronotum widest at middle;
Rocky Mts, eastward.
Polyphylto decimlineata (Say)' Tenlined June bee-
tt.-is-¡S mm; brown with yellowish and white ",
scales; 2 white stripes on head, 3 on pronotum, 5 on
each elytron; Rocky Mts., Southwest' P. crinits
LeConte similar, long erect hairs on head and pro'
notum, Pacific coast states'
Order Coleoplera 199

!
Figure 178 Scarab beetles (Scarabaeidae). A, rose chafer, Macrodaclylus subspinosus;
B-D, .f une beetles: B, Phlttophaga fervidø; C, Phyllophaga fusca; D, a white grub
(Ph.vtlophaea sp.); E, Japanese beerle, Popillia japonica;F, bunrble flower beetle, Euphorio
inda
Seri('e teri(eu Illigcr-8-10 ntnl; <lull blltcli or pur-
plish brown, r'crr- ir iclcscentl tnos( ol'U.S., stluthcrn
Canada (contnron at liglits). S. anlhracina l-eConte
similar, 7-8 mm, not iridescent, Far West
Serica vespertinø (Cyllenhal)-8-ll mm; shiny yel-
lowish brown to dark brown; each deep groove of
elytra with 2 rows of punctures; eastern North
America.
Subfamily Rutelinae-Shining Leaf Chalers;
hind legs closer to middle legs than to tip of ab-
domen, outer claw of hind tarsi larger than in-
ner claw, 2 spurs on tip of hind tibia; usually
shining.
Anomalq unclulata lvlelsheimer-8.0-9.5 nm; con-
vex and oval; shiny brownish yellow; center of pro-
200 Subclass Pterygota
'.:É¡
l:i@
Ð
Ð
EÐ
.J
!!,',
'1,g
t*l
LJ
{
notum blackish rvith greenish bronze luster; elytra
with nrembranous margins; elytra color variable,
usually with crossband of dark brown oval dots near (
middle and in apical l/3, but could cover all elytra
or be nearly absent; eastern U.S.
Cotalpa lanigera (Linnaeus)-20-26 mm; body
shaped like Phyllophaga spp. (June beetles); metallic
dark yellow, whitish yellow, or greenish yellow; ven-
tral side dark with long dense hairs, eastern l/2U,5.
Pelidnota punctala Linnaeus-17-25 mm; shiny
light brown to brownish yellow; pronotum with
black spot on each side; elytra with 3 black spots on
each side; ventral surface blackish or dark green;
throughout U.S., southern Canada.
(' i
Popillia japonica Newman (Fig. l78E). Japanese
beetle-8-12 mm; shiny dark green; elytra brownish
 t
ý

orange with grecn margins: ventl al stlr ce bl¡ckish [:trp]tor irr lrrrla (l-innae us) rFig l78F). Bumble
\villì \\h11,'lìtìbe. llll sl)o1t lll \l(JL'\ of r1l r¡.'1¡ 1¡ l i¡r rlì\, i .. ', iì,l6 rtrrn: i¡rrtr,nisl-t r',¡ll,¡rr;'read and
r¡J' ;ri,,ì,,iìì | \\ ilì
- r i,ì, ' t ¡l- r' ìl .t _ 1 , ¡ll : t.l' \\''rì r'," lr r' . s( melinles
,t l. r, , .l-.C r,,'llO* ltU-
, 't | . I I ì \J
Sullfamilt l)¡nrrstinac--Elc¡rhant. I-lct,:Lrli.' l,t, l;1 ,,1, i r,rgr,, {ì al¡iL-i ls)--9-il r¡¡n; l¡ead and
ancl Ririnoce ros Beetle-s; prirrllrìl¡' Stlrrth arrci lìt() I )tìLr l'r,rn;re-black, \\'liìl dcn:ii yellow hairs;
SoLrthw'esti Ìrortt on head and' or pr(,-notulll (cr ) ,:l¡trrr dark orange-brown lo l)laclíiitt, sparse hairs,
orabsent(?). rvhitc rnarkings; tip ol abdorncn with while stripe on
5/rølegtr-s s¡rp (elcphant beetlcs) ar c brown, 38- cach side; eastern l/2 U.5., southe¿rslern Ca¡rada.
50 mm long, heacl rvith 3 horns, southerrl U S. ¡rorth T. affinis (Cory and Percheron) similar, found with
to NY and west lo TXi D-vnasles spp (Hercules bee- T piger, shiny,9-10 mm.
tles) arc 50-70 mrn, long horn ovcr head, D. tityu's
(Linnaeus) in East, D. grantr Horn in Southwest;
rhinoccros bcctlcs include Phileurus spp., aboì-lt Family Helodidae- Marsh Beetles
25 rnnr long, 2 honls on head, southern l i 2 U S These oval beetles are 2-4 mm long. Some have
X.t' I o t,t, (' t c \ inoccros bectle-
nt u ¡ t n\ s ( Dru r)' ) Rh
enlarged hind lemora for jumping. Adults oc-
.l u a ¡

2-5--ì0 mnr: ihin¡. clark rcddish brorin or

brorvnislr cur orì vegetation and decaying material in wet
black; r'cntraì strll'ace rrirh thick red-btorvn hairs; a reas
I curved horn on head (o) or tube rcle ( Q ); eastern
U.S. west to AZ. Comnron Species
Cyphon colloris Guérin-Méneville (Fig. l79A):'
Subfamily Cetoniinae-Flower Beetles; tarsal 3.0-5.5 mm; shiny black with fine pubescence; red'
claws equal size and untoothed, body flattened dish yellow markings on sides of elytra; eastc.n l/2
from above; Euphoria spp. hairy and resemble U.S.
bumble bees.
Colinus nitida (Linnaeus). Green June beetle-20-
23 mm; narrowed posteriorly; velvety green; head' Family Dascillidae-
tibiae, anci metasternum metallic green; pronotum Softbodied Plant Beetles
and elytra brownish yellow latcrally; eastern I /2 Members of this family are pubescent, soft-
U.S. C te.\eno Casey similar, 20-34 mm; coppery bodied beetles 3-14 mm long. The mandibles
green, TX to so C,A,

Figure 179 A, a marsh beetle, cyphon collaris (Helodidae); B, a softbodied plant beetle'
Dãscittus davidsoni (Dascillidae); ô, a pill beetle, Byrrhus americanus (Byrrhidae); D'
a
waterpenny beelle, Psep he n us herric k i (Psephenidae).

Order Coleoptera
 u
:,,Ö
are prominent in many species. Adults are occa- name refers to the flat, circular larvae. Larvae
sionally collectcd on vegetalior'ì near \À ater. occur under stones in rapidly flowing water O
(e.g., shorelines, riffles) while adults crawl over
Common Spccies rocks and vcgetation along streams. Ð
Da.scillu.ç dut'icl.so¡ti LeConte (l'ig. I 798)- l0-
l4 mnr; shiny broun or blackish; f ine gray pubcs- Common Species {Ð
cence on elytra, 2 irrcgular brown bands presenl or
sometimes absent;CA.
Psephenus herric'ki (DeKay) (Fig. l79D)-4-6 mm;
brownish black or dull black; head and thorax dark-
. ri}
est; line pubescence and punctures; eastern North
America. Remaining Psephenus species occur in {
Family Byrrhidae- P¡ll Beetles Pacilic coast states.
These beetles are convex, black, and 5-10 mm a
long. The head is concealed from above. When
disturbed these hard-bodied beetles retract their Fam i ly Ptilodactylidae ç,
These small (4-6 mm long), brownish beetles
legs and antennae, forming a compact "pill."
Habitats include along sandy shorelines, be- usually have the head hidden from above. The (t
neath logs and stones, and among grass roots. antennae bear a slender process on most seg-
qÐ
Adults and larvae are plant feeders. ments. Ptilodactylids are found on vegetation
near water,
.,.1r@
Common Species
Byrrhus americanus LeConte (Fig. 179C)-8.5-
9.5 mm; broadly oval, very convex; black; dense
Common Species
Ptilodactyla serricollis (Say) (Fie. l80A)-4-6 mm;
L
gray pubescence; elytra with 3-4 black lines; north- brown to blackish; pubescent; eastern l/2 U.S. ,:-
ern l/2 U.S. except Far West, most of Canada,
\i
Family Dryopidae- E)
Family Psephenidae- Longtoed Water Beetles iJ
Waterpenny Beetles Beetles in this primarily aquatic family are
Only six species of these plant-leeding water l-8 mm long, gray or brown, and have long legs
beetles occur in the U.S. The common family with large tarsal claws. The head, with its very

i
!

r
I
''.
Figurc 180 A, Plilodoctyla serricollis (Ptilodactylidae); Il', a longtoed water beetle, Helicltus
lithophilus, and antennae (Dryopidae); C, a riffle
beetle, Stenelmis quadrima':ulata 1J
(Elmidae); D, a variegated mud-loving beetle, Neoheterocerus pallidas (Heteroceridae).
(j
202 Subc ass Pterygota
short antennae, is usually concealed in the pro-
thorax. Adults occur on stones, debris, and
stream bottoms in rapidly moving water such as
riffles and shorelines. The aquatic larvae are
flattened; a lew are terrestrial.
Common Species
Helichus lithophilus (Germar) (Fig. l80B)-5-6 mm;
dark reddish brown with bronze luster; dense cover-
ing of fine pubescence; tip of abdomen pale red be-
neath; east of Rocky Mts.
Family Elmidae-'Riffle Be :tles
Thcsc trcctlcs arc oval to e I lindri, ¡1, -8 nlm
j
long, anci the cl5'tra may' be smoo h ol higìrl¡-
ridged. The legs are long with large tarsal clarvs.
Most species occur on rocks and deb'ris in rilfles
of streams. A few are in ponds ol are terres-
trial.
Common Species
Stenelmis quadrintaculala Horn (Fig. l80C)-
2.'7-3.5 mm;dark reddish brown to black;pronotum
often grayish; 2 oblong yellowish white patches near
base and apical l/3 of elytra; u'hite, waxy pubes-
cence on body;eastern U.S.
Family Heterocer¡dae-
Variegated M udloving Beetles
Members of this family are flattened above,
l-8 mm long, and often variegated with yellow-
ish bands or spots. The flat mandibles extend
forward and the outer margins of the lront and
middle tibiae have a row of distinct spines.
These beetles inhabit burrows along shorelines
and often fly to lights. Adults feed on zoo-
plankton or are omnivorous.
Common Species
Ncohetcrocerus prrllidtts (Say) (Fig. l80D)-6-7 mm;
hlack: covcrcd u ith brorvnish and vcìlotish hairs,
latter iornling 3 incìistinct crossbands on clytra;
nrost ol [.] S
Family Rhipiceridae-
Cedar Beetles
Cedar beetles, brown insects with orange anten-
nae, resemble June beetles (Scar¿rbaeidae). The
larvae parasitize cicadas. The five North Amer'
ican species are in the genus Sandalus.
Fami ly Buprestidae- Flatheaded
or Metallic Wood Borers
Buprestids are a large family (about 15,000 spe'
cies worldwide and 720 species irr North Amer'
ica) ol harcl-bodicd, oftcn rnetallic rlr brightly
colore-<l [.cetles The bo<11' u:,u;tlll' ltar; ¿t charac-
-l
tcri<1ir' rha¡rc (Fig. l8l) his llrrnily r:sembles
thc click bc'ctles (Elatc'ridaet but rrlelnìrers lack
thc pointcd posterior corners of the pronotum
and the movable prothorax of'tlre click beetle.
Adults frequent flowers, leaves, and tree truirks
and limbs that occur in full sunlight (e.9., the
southern edge of a woods). Dead or dying trees
ancl especially freshly cut wood attract these
beetles. Larvae, known as flatheaded wood
borers, typically have an expanded and flat'
tened anterior area (Fig. lSlC). The winding,
grass-filled galleries under bark, and oval holes
in wood are signs of these larvae, many of
which are serious pests of orchards and forests.
A few species produce twig galls and some are
leaf miners (genus BrachYs).
A few of the common genera are identi-
lied as follows: Agrilus species have long, nar'
row bodies and the first segments of the hind
tarsi are as long as the three following segments
combined; Chrysobothris species have the
coxae of the hind legs flattened and distinctly
widened at the bases, the prosternum is broadly
diamond-shaped behind the procoxae, and the
lateral lobes of the third segments of the hind
tarsi are equal in length to the second segments;
Dicerca species have the tips ol the elytra
elongatecl and cliverging.
Order Coleoptera

Figure l8l Flatheaded or metallic wood borers (Buprestidae). A., Acmaeodera pulchella;
B, rednecked cane borer, Agrilus ruficollis: C, flatheaded apple tree borer, Chrysobolhris
femorata adult and larva.
Common Species
Acmaeodera pulchella (Herbst) (Fig. l8lA)-5.5-
12.0 mm; cylindrical; shiny, dark bronze-brown;
pronotum with orange or yellow spot near posterior
corners; elytra black with orange- ¡eìlorv nrarkings;
many short brou'n hairs; most oi US , southcrn
Canada
Agrilus anxius Cory . Bronze brrch borer-6-13 nrm;
dark green-bronze; front of head greenish or cop-
per-bronze; front of pronotum coppery; apex of ely-
tra granular; most of U,S. and Canada (on bj¡ch).
Agrilus hilineatus (Weber). Twolined chestnut
borer-6-12 mm; greenish black; dense yellow pu-
bescence on sides of pronotum and elytra, forms 2
longitudinal stripes on elytra; east of Rocky Mts.
Agrilus ruficollis (Fabricius) (Fig. l8l B). Rednecked
cane borer-5-8 mnr; bluish black; head and thorax
coppery red;east ol RockY Mts.
Anlhaxia aeneogclster Castelnau and Cory-
4-5 mm; flat; dull bronze-black dorsally, brassy
green ventrally; Rocky Mts. westward.
Buprestis apricans Herbst. Turpentine borer-24-
26 mm; grayish bronze with greenish ¡netallic luster;
each elytron with I rows large punctures; NC to TX
(on pines).
Buprestis aurulenla Linnaeus. Golden buprestid-
l4-19 mm; iridescent grcen or bluish green; elytral
margins and central strìp goldcn or coppery; Rocky
Mts. westward
204 Subclass Pterygota
Uj
qf
e
Ð
it
U
;
q,
@
Chalcophora virginiensis (Drury)-20-38 mm; shiny
,g
bronze-black; punctures and ventral side brassy;
pronotum and elytra roughly sculptured; irregular I
lighter areas on elytra; eastern l/2U.5. and Canada.
C liberta (Germar) similar, l9-25 mm, bright cop-
pery or brassy, blackish brown markings.
t
t
Chrysobothris femorata (Olivier) (Fig. l{llC). Flat-
headed appletree borer-7-16 mm; dark bronze,
sometimes brassy, coppery, or greenish luster; usual- u
ly 2 irregular crossbands on elytra; coarsely and
densely punctate; throughout North America, ü
Dicerca divaricala (Say)-15-21 mm; brown or gray Ç/,
with brassy, coppery, or greenish bronze luster; apex
of elytra prolonged and diverging; throughout North
America.
Melanophila Julvoguttala (Harris). Henrlock bor-
er-9-12 mm; shiny black with brassy luster; apical
l/2 elytra with 6-8 small orange-yellow spots in a
circle; body rough and punctated; northern l/2U.5.
south to AZ, southern l/2 Canada (on cut pine logs,
spruce, hemlock).
Family Elateridae-Click Beetles L
4
Click beetles are elongated, flattened insects
with a large, movable prothorax výhose poste-
rior corners are usually pointed. The body is
often greatly tapered posteriorly. The family is r
large containing over 7,000 species v/orldwide
and nearly 800 species in North Ame:rica. The
spinelike prosternal process (Fig. ltl2A) that
U
fits into a socket in the mesosternum and the
loosely joined prothorax and mesothorax en-
able the bectle to flip over w'herl it is upside
down. Tlrc treetle arches its bod-v and then spr ne
quickly stritightr-trs it, f'orcing the spine back
into lhc sockcl artrl snapping lhc pronottr m ancl
elytral base against the supporting surlace; the
beetle ilips ovcr with an audible cIcking sound.
Adults feed on vegetation and are especially
common under loose bark of decaying trees.
Larvae, called wireworms, are cylindrical
and elongated, usually reddish brown, and have
a hard and shiny cuticle. Wireworms occur
chiefly in the soil and many are crop pests due
to their feeding on roots, seeds, and stems'
Some species inhabit rotting wood or occur be-
neath bark and feed on other insects.

Common Species
Agriotes nett(u,t (Say). Wheat r,r irervortn-7-9 rnm;
yello*ìsh brt,rrn trr dark bro*n; pronottlrn broader
than long; tle nse co¿rrse punctt¡rcs in deep cll tral
grooves; posterior corners ol- pronolum and sides of
elytra dull ycllow; shorr dull yellow pubescence; east
of Rocky M ts

Alaus oculatus (l-innaeus). Eyed click beetle-25-

45 mm; shiny black with sparse gray-white scales;
pronotum with 2 large oval eyespots margined with
gray-white sca¡es; east of Rocky Mts (in decaying
logs). /. melanops LeConte similar, 25-36 mm'
without scalcs on pronotum, Far West. A myops
(Fabricius) also similar, 24-38 mm, sparse pubes-
cence, small pronotal eyespots with indistinct mar-
gins, chiefly southern U.S. Figure 182 Click beetles (Elateri<Jae)' A, spine on
prosternum of click beetles (also on buprestids);
Ampedus collaris (Say)-8-9 mm; shiny black; pro- B, tobacco wireworm, Conoderus vespertinus' Lawa
thorax bright red; antennae and legs dark brown; and adult; C, Ctenicera inflata; D, Melonolus com'
eastern North America. munts.

Conoderus vespertinus (Fabricius) (Fig. l82B). To-
bacco wireworm-7-10 mm; yellowish with dark
reddish brown or blackish markings as illustrated;
eastern |/ZU.S.
Ctenicera inflata (Say) (Fie. l82C)-8- I I mm;
bronze-black; relatively broad, convex; Iegs and
sides sometirnes reddish; dense yelÌou'ish pubescence
pressed against body; 3rd antennal segment ) 2x
length of 2nd and longer than 4th; most of U.S.'
southern Canada.
Ctenicero lobata tarsalrs (Melsheimer)-9-l? mm;
shiny black; elytra dull yellow, lateral and inner
margins with narrow black line; 2nd antennal seg-
meni very small, 3rd triangular and larger than 4th;
eastern U S. C. pruinina (Horn), Creat Basin wire-
worm, similar but all black, elytra der:ply grooved,
Pacific Northwest, Creat Basin.
Limonius californicus (Mannerheim). Sugarbeet
wireworm-7-10 mm; antennae and femora dark
brown; elytra reddish brown; dense white or yellow-

Order Coleoptera :
ish pubescence; dorsu¡n coarsely punctured; western
North America.
Melanotus (otnnlttnrs (Gy llenhal) (Fig. l 82D)-
I l-15 mm; rcddish bro*,n; sparsc pubescence; finely
punctate; 3rd anrennal segment twrce Icngth of 2nd
and together as long as or longer rhan 4th; eastern
l/2 North Arnerica (vcry comnlon; llies ro lights).
M. sitnilis (Kirby) similar shapc, l3-17 mm,
brown to blackish, nrost of North America.
dark
Family Lampyridae- Firef lies
The broad, llat pronotum generally covers the
head from above in this family. The elytra are
relatively solt. Light is produced in organs
located in the yellow or greenish areas near the
end ol the abdomen. This biolumenescence is
caused bv a¡l cnzvnte, luciferasc, reacting with a
substance, lucilerin. The color and llashing lre-
quency of the light vary by species and are as-
sociated with mating. Most of the individuals
flying at night are males; the females of most
species fly very little and remain in the vegeta-
tion while flashing. Larvae are also able to
glow; they and the short-winged or wingless
females are called glowworms. Not all species
produce light. Adults and larvae are p 'daceous
on insect larr ae, slugs, ancl snlils.
Common Spccics
Ell¡'t'hnia corru\ca (l-innaeus)- l() I 1 nlrn; black or
brorvnish bl¡ck I rcdrlish i'ello',i crrrvccl ¡ atch nc.ar
but not reachrng sidc ntargins of ¡tronolrrrn; clr rra
with brorin or -u-cllo*ish prrbesccncc and..-4 ridgcs;
no light organs; most ol U S (diurnal)
L.uc'irlota atrt (l:abricius) (Fig. l83A)-8-l I nrm;
black; sides ol pronotum yellowish with red-orange
marking abovc; ell'tra granular, 4lou,ridgcs in basal
2/3;eastern North Arncrica (in trees;weak light¡.
tl{ic'rophotrts dnputtus l.eClonrc-10-15 rnrn; pro-
r.totunr ancl clVtra gravrsh bro$n ancj abdortrcn pink-
ish (cr); Ç l:rrr.aìikc., pirrkish, f'latrcned, rinvclyrra;
C'4, C)R, ( O (Llrv erass ol loorhilìs)
Photittttç si'inttllqns (Sai)-_5-8 rlrn; dark brou,¡;
pronolurn reddrsh *,ith dull ycllo* nrargìn and black
central spot; elytra outer margin and central stripe
206 Subclass Pterygota
yellow; Ç very short-winged; eastern U.S. south-
west to KS and TX. P. pyralis (Linnaeus) similar, (Ð
l0-14 mm, elytra slightly wrinkled, Ç long-winged.
Photuris pcnnsylvanicr.rs (De Geer) (Fig. l83B)-l l- c
l5 mm; head yellowish; pronorum yellowish with
reddish central area and dark middorsal stripe; ely-
tra dark brown with yellowish stripes and mafgins;
o
eastern U.S. southwest to KS and TX. 6,
Family Cantharidae-
e
Soldier Beetles O
in this family generally have dark yel-
Beetles
low, orange, or red elongated bodies. The ely- g
tra are pliable or leatherlike and the eyes bulge
outward. The beetles are common on flowering
plants. Adults and larvae are chiefly preda-
e
ceous on other insects although some adults (Ð
leed on nectar and pollen or are omnivorous.
@
Common Species
Cantharis bilineatus Say-6-8 mm; antennae black
ú.
except base, 2 oblique black pronotal bands, elytra ('
black except side margins, and legs black; remainder
of body reddish yellow; eastern l/2 U.S,
'L/
Canlharis dit,isa LeConte-6-8 nrm; b,lack; thorax
yellow u'ith 2 large black spots rn central area; CA to
ì]ritrh ('olrrnrbilr.
tJ
(-ltuulro'¿rttttltus l'¡enn-çvlt o¡¡itu, (t)e Geer)
(Itig l8-ì( ¡ -t¡- i2 nìnì; pronoluln altrl elvtla dull
1'eìlou -orarrgc i,:th a dark orirl arca ttn ¡tostr:rior l/2
ol t'ach el¡ tron; rcst of bod¡' blackislr; cast of Rocky
lvf rs,, AZ (on goldenrod and Queen r\:nne's lace in
late surlrler)
Podabrus |omentosus (Say) (Fie. l83Dl-9.12 mm;
elytra black with grayish pubescence; head, antennal
base, prothorax, and legs reddish yellttw; most of
U.S. (on giant ragweed along streams).
,j
Tr-v¡:therus latipennis (Germar)-6-7 mm; elytra
short, coverirry l/2 or less of body; hind wings ex-
tend to tip of abdomen (hind wings short in similar
rove beetles [Staphylinidae]); dorsal surface black;
ely,tral ti¡rs, pronotal margins, and ventral side yel- ;
loivish; easrern I /2lJ.S. (on catnip).
ü
\-ì

Figure 183 A and B, fireflies tLampyridae)'. A, Lucidoto atra;8, Photuris pennsylvanicus.

Cãnd D, soldier beetles (Canrharidae): C, Chauliognathus pennsylvanicus} D, Podubru's
tomenlosus

Family Lycidae-
Netwinged Beetles
Members of this family usually have soft,
ridged, netlike wings and a large pronotum that
often covers most of the head. A few western
species (genus Lycus) have a short snout'
Adults feed on decaying vegetation and larvae
are predaceous. Many species are brightly col-
ored and occur in wooded areas witþ dense un-
dergrowth,

Common Species Figure 184 A netwinged beetle,

0- latum (Lycidae).
of
rh
rn Family Dermestidae-
Canada. Dermestid Beetles
Calopteron terminale (Say)-8-17 mm; orange-yel-
This group consists of convex'
low; posterior l/3 of elytra with purplish black elongate-oval beetles 2-12 mm long' The
band; most of U.S. nae are clubbed and the body is hairy or
Larvae are light brown and covered
Dictyopterus aurora (Herbst)-6-1 I mm; black; pro- hairs (Fig. l85A). Most species are sca
notum with orange-red sides; most ol North Amer-
lca.
the larvae feed on leather, furs, ntuseum specl-
mens (including insects), woolen and silk
ucts, rugs, stored foods, and carrion.

Order Goleo

Figure 185 Dermestid beetles (Dermestidae).
larva and adult; B, larder beetle, Dermestes lardarius.
Common Species
Anthrenus scrophulariae (Linnaeus) (Fie. l85A),
Carpet beetle-2.0-3.5 mm; broadly oval, convex;
black; white scales cover sides ol pronotum and
form zígzag irregular crossbands; dark red or dull
yellow stripe along inner margins of elytra; through-
out North America (adults on flowcrs),
Allagenus megüloma (Fabricius). Black carpet bee-
tle-3.5-5.0 mnr; oblong; convex; head and pro-
notum black; elytra reddish brown ro black; pubes-
cence sparse; throughout North America.
Dermestes caninus Germar-7.0-S.s mm) utact;
pronotum with dense yellow and white pubescence;
gray, reddish brown, and black pubescence on ely-
tra; throughout North America (used to clean flesh
from bones of decaying animals).
Dermestes lardarius Linnaeus (Fig. l85B). Larder
beetle-6.0-7.5 mm; elongare-oblong; black; basal
l/2 elylra with dense yellowish, light brown, or gray
hairs, 2 black patches at base, 6 black spots near
middle of basal I /2; throughout North America (on
stored meat, cheese, insect collections).
Trogoderma ornata Say-2.0-2.5 mm; oblong-
ovate; shiny black with scattered yellow pubescence;
black elytra with red patches at base and smaller
patches apically; throughout North America (also in
stored cereals, seeds, nuts).
208 Subclass Pterygota
c
c
@
e
A, carpet beetle, Anthrenus scrophulariøe,
Family Ptinidae-Spider Beetles
Spider beetles are I -5 mm long, the pronotum is
very narrow, and the legs are quite long. The
head is partially or completely hidden from
above. These beetles often resemble spiders.
They inhabit dried plant and animal materials I
such as stored grain, carcasses, and museum
specimens.
Common Species ;
Ptinus /ar (Linnaeus) (Fig. 186A). WhitemarkçC
spider beetle-2,5-4.5 mm; dull reddish yellow Or
pale brown (o,), or dark brown (9); bodyoval 19j
or elongate-oval (6); pronotum with pale yellowish
markings; elytra with irregular patches of white
Family Anobiidae- Deathwatch
and Drugstore Beetles
Beetles in this family are l-9 mm long, the pro-
notum extends hoodlike over the head to con.
ceal it, and the body is usually cylindrical but
can range to nearly spherical. The last three
segments of the antennae are usually length-
ened and enlarged although a few species have
serrate or pectinate antennae. Adults and
live and feed on dry plant materials such as
dead twigs and logs, furniture, seeds, cereals,
tobacco, and museum specimens. The name
"deathwatch" is derived from the ticking noise
a few woodboring species make, once said by
some to foretell a death.
Common Species
Lasioderma serricorne (Fabricius), Cigarette bee-
tle-2-3 mm; elongate-oval, convex; dull reddish
yellow or brownish red; antennae serrate, 2nd and
3rd segments shorter than lst, llth segment oval;
throughout North America (indoors).
Ptilinus ruficornis Say-3.0-4.5 mm; elongate, cy-
lindrical; black or dark reddish brown; antennae and
legs reddish yellow; antennae lan-shaped ( ct),
branch ol 4th segment 6-7 t.imes length of segment;
eastern U.S. (on dead branches of oak and maple).
Stegobium paniceum (Linnaeus) (Fig. l868). Drug-
store beetle-2.5-3.5 mm; oblong, convex; reddish
brown with yellowish pubescence; sides of pronotum
serrated anteriorly; elytra finely grooved and punc-
tured; throughout North America (indoors).
Figure 186 A, whitemarked spider beetle, Pfinus
/ur (Ptinidae); B, drugstore beetle, Stegobium
paniceum (Anobiidae).
Family Bostrichidae-
False Powderpost Beetles
These cylindrical and elongated beetles are
ally 3-20 mm long and the head of most
is bent downward so it is almost
above. A few species primarily in the'
states have a readily visible head and large
dibles. The pronotum in this family has
tubercles anteriorly and the tips of the elytra
bent downward in many species.
bore into living trees, dead branches,
seasoned lumber. One species in the western
states, Scobiciq declivis (LeConte) sometimes
bores into lead telephone cables (it does not eat
the lead) causing a short circuit from the
moisture that enters the holes
Conrmon Species
Amphicerus bicaudatus (Say) (Fig. l87A).
twig borer-6.0-l 1.5 mm; reddish brown to
ish black; pronotum round, many tubercles on
rior l/2; elytra slope downward at apex, 2,
curve inward near apex of elytra; eastern U
to CO, NM (burrow into fruit tree twigs). z4
nu /us (Pallas) similar, I l-13 mm, Southwest,
Melalgus confertus (LeContc) (Fig. l87B)-
I5 mm; black; elytra brown; head not concealed by
pronotum, mandibles large; pronotum narrO"iveSt at
base; Far West (in orchards)
Family Lyct idae-
Powderpost Beetles
Lyctid beetles are 2-7 mm
somewhat flattened, and brown or
head is visible from above and the
club is 2-segmented. These beetles bore
dried wood and wood products such as furni.
ture, beams, and hardwood floors, reducing,
them to powder
Common Species
Lyctus linearis (Goeze)-25-30 mm; reddish
elytra with I row of punctures; throughogt
America; L. cavicollis LeConte similar, )
punctures on elytra, Pacific coast states
Order Coleoptera 2Og
 commonly occur under bark and feed on
insects and possibly fungi.

Common Species
Calitys scaöra (Thunberg) (Fig. l88A)-9-10 mm;
reddish brown; head about l,/4 greatest width of
pronotum; front coxal cavities closed; Far West.

Temnochila virescens (Fabricius)-10-13 mm; elon- @

gated and flattened; metallic green or blue;
West. &
Tenebroides corticalis (Melsheimer)-7-8 mm;
black; shape similar to T. mauritonicus; ïth
nal segment smaller than 9th; eastern l/2U,5
Tenebroides mauritanicus (Linnaeus) (Fig. l88B).
Cadelle-6-ll mm; flattened; shiny dark brown to
black; pronotum not grooved; throughout
America (in grain, flour).

Figure 187 A and B, false powderpo beetles

(Bostrichidae): A, applc twig borer, Am¡tlt ccru; bi-
caurlatus; I), .\lclulgtts confcrttrs. ( , sc-r.rthi rn lyctus
bcct le, 1- v t' tr r s ¡t Iu n ic rt l/i s' ( l- yct rcla,: )
c
;Fr
Lyctus planicotlis [.cConte (Fig. l87C). Southern
lyctus beetle-4-6 mm; blackish with short yellow- Figure 188 Trogositid beetles (Trogositidae).
iÐ
white hairs; pronotum as long as wide; each elytron A, Cali tys scabra; B, Tenebroides maurilqnicus,
with I I lidges that contain hairs, double row of
punctures in grooves; most of U.S., chielly South.
:t
Family Cleridae- ,:<)
Family Trogositidae (Ostom idae)- Checkered Beetles 1,i:

The beetles in this family are often brightly col- LJ

Trogositid Beetles ored with orange, red, or blue, and the body is
This group has two dilferent forms: one form very hairy. The last segments of the palpi are
(subfamily Ostominae) is oval, the head is
dilated and there are membranous lobes be-
much narrorvcr than the pronotum, and the ely-
neath the tarsi. Most adults and larvae are
tra have long, crect hairs; the other form (sub- predaceous on other insects, especially wood-
family Tenebroidinae) is elongate, the head is
about equal to the width of the pronotum, and
inhabiting beetles such as bark beetles
(Scolytidae). Many clerids occur on flowers and ,

the pronotum is widely separated from the ely-

foliage. Species in the genus Necrobiø, known I
tral base. Adults are generally metallic green,
as ham beetles, occur on spoiled meat, fÏslr, and
blue, or dark brown to black. Adults and larvae
cheese, feeding on insect scavengers that are
present.
210 Subclass Pterygota
 ò
I

Figure 189 Checkered beetles (Cleridae). A, Enoclerus nigripes; B, E. sphegeus; C, Phyllo-

baenus pallipennis; D, Thanasimus dubius.

Common Species Family Melyridae-

Enoclerus nigripes (Say) (Fie. lE9A)-5-7 mm; dull
red or reddish brown; apical 2/3 elytra hlack with 2
whitish crossbands; eastern U.S. (in fores ).
Enoclerus sphegeus (Fabricius) (Fig. l89Iì)-9-
l0 mm; facc yellorr'; pronotum arrC ely'tri bro* nish
black, elytra with 2 apical yellow or u'h:tish cross-
bands; abdomen red; western l/2 U.S. (in forests,
May-June).
Softwinged Flower Beetles
Members of this family have prorninent front
çoxae, the antennae insert in front of the head
above the mandibles, and the body may have a
soft appearance from the numerous; erect hairs.
Many species have a fleshy lotre betvveen the
tarsal claws and some have oran¡1e, eversible ,;
sacs along the sides of the abdonten that arç .

Necrobia rufipes (De Geer). Redlegged ham bee¡le-

3.5-6.0; shiny metallic blue or green; basal segments
of antennae and legs reddish brown; most of North
America (on drying carrion, bones, fish, ham,
cheese). N. ruficollis (Fabricius), redshouldered ham
beetle, similar, 4-5 mm, head and posterior 3/4 ely'
tra metallic blue or green, pronotum and base of ely-
tra reddish brown.

Phyllobaenus pallipennis (Say) (Fig. l89C)-3.5-

5.0 mm; bronze-black; antennae, elytra and legs red with 2 oblique markings; elytra blue-black;
brownish yellow; elytra with apex, side margins and ern I /2 U.S.
central crossband faintly dark brown; pronotum
widest at middle; most of U.S.
Collops quodrimoculaløs (Fabricius) (Fig. I
4-6 mm; pronotum and elytra reddisit yellow'
Thanasimus dubius (Fabricius) (Fig. l89D)-7.5-
elytron with 2 large blue-black patches; rest of
9.0 mm; head, pronotum, and basal area of elytra
brownish black; east of Rocky Mts.
reddish brown; remainder of elytra black with 2 ir-
regular bands of light pubescence; eastern I /2 U.S.
Collops vittatus (Say)-4-5 mm; black;
and sides of each elytron reddish'yellow or
Trichodes nutalli (Kirby)-8-11 mm; dark blue,
greenish blue, or purple; elytra with 3 reddish yellow
orange; remainder of elytra dark blue;
may have central dark area; hairy and
crossbands; east of Rocky Mts., ID, British Colum-
most of U.S. and Canada.
bia. T. ornatus Say similar, 6-7 mm, 3 yellow irregu-
lar crossbands on elytra.

Order Coleoptera
Malochius aeneus (Linnaeus) (Fie. l908)-5-6 mm;
front of head, anterior corners pronotu r, and all
but base and cenrral strip oi elyira reddìsl bro"vn to
orange; rerlaìnclel dark nl,:taìlii grcer : eait ol
Rockl'!lts., ()R, \\ ,\, tsrìtislr (-olumbia
Figure 190 Softwinged flower beetles (Melyridae)
A,, CoIIops quodrirnuculalus; B, Malachius aeneus.
Family N itidulidae-Sap Beetles
Sap beetles are l.5-12 mm long, somewhat flat-
tened, the last three abdominal segments are
clubbed, and the elytra are sometimes short-
ened leaving the tip of the abdomen exposed.
Adults and larvae feed primarily on decaying
plant materials, including damaged and fer-
menting fruit and vegetables, sap, and fungi
beneath bark. A few are predaceous on other
insects. Glischrochilus species (Fig. l9lC) are
referred to as picnic beetles because they are at-
tracted to food and beverages at picnics.
212 Subclass Pterygota
,iÐ
,.',@
ì'@
H¡'@
r'l;* l
rÌï@
Figurel9l Sap beetles (Nitidulidae). A,
beetle, Carpophilus hemiplerus; B, Conotelus
obscurus; C, Glischrochilus fasciatus; D, Omotitø
Common Species
Corpophilus dimidiatus (Fabricius), Corn sap
tle-2.5-5.0 mm; brownish yellow to black; squarish
pronotum; orange spot on each elytron; most of
U.S, except Rocky Mts. and northçentral states (in
corn lields).
Carpophilus hemipterus (Linnaeus) (Fig. l9lA).
Driedfruit beetle-2.5-4.5 mm; black; elytra with
dull brownish yellow bands; most of U.S., especially
South.
Conotelus obscurus Erichson (Fig. l9lB)-2.5"
5.0 mm; very slender; short wings; antennae (except
club) and legs brownish yellow; east of Rocky Mts: 1:
(in flowers). C. stenoides Murray similar, elytra
wide with fine grooves and coarse punctures, south-
ern U.S. (on corn).
Glischrochilus fasciotus (Olivier) (Fig. l9
5-7 mm; shiny black; elytra with four yellow or
dish patches; throughout North America-. i '
Glischrochilus sanguinolentus (Olivier)-3.5-
6.5 mm; shiny black; elytra red with 2 black spots,
apex black; eastern l/2 North America. A. quad-
risignatus (Say) similar, reddish brown, elytra with
4 pale yellow spots.
Nitidulq bipunctata (Linnaeus)-5-6 mm; black;
reddish spot on apical l/2 of each elytron; fine pu-
bescence; throughout North America (on bones and
skins of dead animals).
elytr dark brown; very pubescen¿ and
punctate; east of Rocky Mts, (under stones,
Uleiota dubius (Fabricius) (Fig. l92C)-4-6
very flattened; brownish black or pale brown,
margins paler; pronotal margin serrated;
U.S., southern Canada (under bark).

Omosita colon (Linnaeus) (Fig. l9lD)-2-3 mm;

brownish black; margin of thorax ycllow; each ely-
tron with 3-4 yellow spots near base and large patch
on apical l/2 that contains a dark spot; east of
Rocky Mts.

Family Cucujidae-
Flat Bark Beetles
The unusual flatness of most of these beetles is
a good field identification characteristic. Cucu-
jids are 2-12 mm long. Most adults and larvae
live under bark where they prey on bark beetles,
other small insects, and mites; others are para-
sitic or infest stored grain and similar foods.

Common Species
Cathartus quadricollis (Guérin-Méneville). Square-
necked grain beetle-2.5-3,5 mm; reddish brown;
pronotum longer than wide, rectangular to square,
margins not toothed; throughouß North\America (in
stored foods).

Catogenus rufus Fabriçius-5-l I nim; dark reddish Figure'192 Flat bark beetles (Cucujidae).
brown; crossgroove on head behind eyes; pronotum toothed grain beetle,
with depressed central area; eastern U.S, B, Telephanus velox; C,

Cucujus clavipes Fabricius-10-14 mm; entirely red

Family Cryptophag¡dae-
Beetles ì
or dull red; most of North America (under bark), :
r:
Cryptophagid
Oryzoephilus surinamensis (Linnaeus) (Fig. l92A)- These beetles are l-5 mm long, yellowish
Sawtoothed grain beetle-2.5 mm; dark brown to
reddish brown; pronotum with 6 teeth on each side or black, and usually covered with silky
and 3 ridges on dorsum; throughout North America cence. The body is oval and convex and thç
(in stored foods). O. mercator (Fauvel), merchant tennal club is 3-segmented. Cryptophagids
grain beetle, similar but area directly behind eye beneath leaves and wood chips, in
is 1l/2 vertiçal diameter of eye whereas it is ) l/2 and on fungi and flowers.
in O. surinamers,s, on cereals but not grain.

Telephanus velox Haldeman (Fie. l928)-3.5-

5.0 mm; yellowish brown; head and apical l/3 of

Order Co leoptera
Common Species club. The tarsi appear 4- or 5-segmented. '

Anchicera ephippiata Zi¡rtmernrann (Fig. l93A)- Adults and larvae occur in decaying wood,
l-2 mm; shinv black head and pronotum; elytra red- under bark, on fungi, or in debris, and they
dish yellow wìth 2 dark patches so¡netimes extending feed on fungi.
into wide band; most ol'U.S.

Cryptophagus qcutangulus Gyllenhal-2.5 mm; flat- Common Species

tened; pale brownish yellow, head and thorax
darker; lateral margin of pronotum bowed inward
on apical l/2 leaving a projection at middle and on
anterior margin; throughout North America (some-
times in stored food).
Ischyrus quadripunctalas (Olivier) (Fig' l94A)-
7-8 mm; head black; pronotum Yellowi h red or yel-
low with 4 black spots; elytra reddish yellow or yel-
low with black patches as illustrated; eastern U.S' ,
Megalodacne åeros (Say) (Fig. l9aB)-18-21 mm¡
shiny black; elytra with 2 red patches at base and 2
on apical l/3; eastern l/2 lJ.S. M. lasciata
(Fabricius) si ilar,9-15 mm.

Triplax festiva Lacordaire-5.0-6.5 mm; shiny

black; pronotum, scutellum, and central crossband
of elytra reddish yellow; eastern l/2 U.5.,
South.

Figure 193 A, a cryptophagid beetle, Anchicerct

ephippiata (Cryptophagidae); B, clover stem borer,
Lø n gu ria m oza rd i (Languriidae).
\,

Family Languriidae-
Languriid Beetles
These elongated beetles'are 5-10 mm long and
black or blue-black with red or orange mark-
ings. Adults are pollen and leaf feeder and the
larvae are stem borers.

Comnron Specit's
Languria nto:.ttrcli L¡trcìllc (Fig l93B). Clc rer sIcm
borer-5.0-7.5 nim; shiny blue-black; hc,rd, ¡lro-
notum, and ventral side red; east ol lìocky )"4ts ' AZ
(on clover, alfalfa).

Family Erotylidae-
Pleasing Fungus Beetles
Erotylids are 3-20 mm long and generally oval Figure 194 Pleasing fungus beetles (Erotylidae).
with black, reddish, and yellowish markings. A, Ischyrus quadripunctatus; B, Megalodacne
heros; C, Tritoma humerolis.
The antennae have a large 3- or 4-segmented

214 Subclass Pterygota

Tritoma hunteralis Fabricius (Fig. l9aC)-3-4 mm;
shiny blackì cl)tra rvith reddish yellow patches near
base; eastern I.l S. f, sanguinipcttnis (Sat') sirnilar,
4-5 mm, el1'rra red.
Family Phalacridae-
Shining Fungus Beetles
These brownish beetles are very convex, shiny,
and l-3 mm in length. The body is broadly oval
to nearly spherical. Phalacrids are common on
goldenrod, daisy, Queen Anne's lace, and
many other f,lowers.
Common Species
Phalacrus simplex LeConte (Fig. l95A)-2.0-
2.3 mm; red-brown or blackish; antennae and legs
pale; antennae insert under shelf between eyes so
bases not visible; central and western U,S,
Stilbus opicalis (Melsheimer)-1.9-2.4 mm; oval;
shiny black except tipsof elytra abruptly reddish;
eastern l/2U.5.
Figure 195 Shining fungus beetles (Phalacridae)
A,, Phalacrus simplex; B, Stilbus nitidus.
Stilbus nitidus (Melsheimer) (Fie. l95B)-l
1.5 mm; very sh y orange-brown; eastern l/2U,5.
North America. Some lady beetles are similar
to the leaf beetles (Chrysomelidae) but differ
having three distinct tarsal segments instead
appearing to have four as do leaf
Adults and larvae are predaceous,
found feeding on aphids and sometime¡
insects, mealybugs, and mites. Two
species in the genus Epilac:hna are plant
and garden pests. Lady beetle adults
in the winter and some are found in.
gregations under debris or on
elevations. Larvae (Fig. l96F) are
beetles are described below. The genus Scym-
n¿¿s contains tiny pubescent beetles, usually
I -3 mm long, and marked with red or yeliow.
Common Species
lady beetle-4-5 mm; very convex, spherical; shiny
ì ì ¡
black; I round red spot in center ot'r:itclt elyr-ron; ab-
clorrrinal scgrnr'nls recl; most ol Ll.S. anc Calrada.
Coccinella tranSversoguîluta richardsonì Brown
(Fig. l96C). Transverse lady beetle--6.0-7.5 mm;
head black, white spot on eaçh side ¡rear eyes; pro-
notum black with spot on anterior corners; elytra red
or yellow with black band and spots as shown;
j:
throughout North America.
Order Coleoptera
 ir

I
I

o
Ð
{)

Figure 196 Lady beetles (Coccinellidae). A, twospotted lady beetle, Adalia bipunctota; B,
twice-stabbed lady beetle, Chilocorus stigma; C, transverse lady beetle, Coccinella trans-
versoguttolq richardsoni; D, Coleomegilla maculata fuscilabris; E, Cyclonedq munda; F,
convergent lady beetle, Hippodamia convergens, adult and larva; G, H. porenlhesis;H, ollo
abdominalis.

216 Subclass Pterygota

Coleomegitla ntaculqta fuscilabris Mulsant
(Fie. l96D)-5-7 mm; elongate oval; head black;
pronotum and elytra pinkish red or sometimes yel-
lowish with irregular black spots; throughout North
America,
Cycloneda munda (Say) (Fig. l96E)-4-5 mm; very
convex; head black with white on front (O) or 2
white spots (9): pronotr¡rn black bc' Cered with
white, anlerior nrargin rvith 3 s'hile d¿r 'hcs towarcl
posterior nl¡trg,in; elytra reddrsh rcllou , r lccl, *'ith-
out spots; tlrroughottt U.S , chii lìy \\'c,tL'rn slates.
C. sanguineu (Linnaeus) si¡nilar, r:asl ot' ìlocky Mts.,
Az.
Epitachna t'ørivestis Mulsant. Mexican bean bee-
tle-6-7 mm; brownish yellow; each elltron with 3
basal, 3 central and 2 apical spots; most of U'S.'
southeastern Canada (on beans; larva spiny and
plump). E, borealis (Fabricius), squash beetle, simi-
lar, 7-10 mm, spots larger, only I apical elytral spot'
eastern l/3 U.S.
H ippoda m ia co n verge ns -Méneville
C uérin
(Fig. l96F). Convergent lady beetle-4-6 mm; head
anã thorax black with pale whitish margins; 2 whit-
ish bars converge on prothorax; elytra reddish
orange usually with 6 spots on each elytron, spots
sometimes fewer in number or absent; throughout
North America.
Hippodamia
'mm; parenthesis (Sav) (Fig' l96C)-
4-j head mostly black; pronotunlr black with
white irregular border; elytra yellowish red. with
black apical markings forming 2 parenthesesi
throughout North America.
Hippodamia tredecimpunctata tibialis (Say)' Thir-
t.énrpott.d lady beetle-4.5-5.2 mm; elongate oval;
head black; pronotum black with pale yellowish
border; elytra orange or reddish with l3 black spotsi
throughout North America.
Hyperaspis unclulata (Say)-2.0-2.5 mm; shiny
black; head yellorv (6); pronotum ycllorv Iaterally;
each elytron with irregular, yellorvish oval spot near
center, and narrow marginal band; most of U'S',
southern Canada' H. signata (Olivier) similar,
2.5-3,0 mm, elytra lack marginal bands, head and
pronotum all black ( I ); eastern I /2 U 'S.
Ollo abdominalis (Say) (Fie. l96H)-4-5 mm; grav
to pale yellow with black spots as shown; throughout
North America (especially western states)'
Paranae¡nia vittigera (Mannerheirn)-4-5
black; pronotum reddish with 2 large black
elytra reddish with 3 broad longitudinal
West.
Family E ndomych
Handsome Fungus eetlês
Thcse sm¿rll (chiefly 3-8 mm long), oval beetles,
are shirr¡- ancl often brighl oratllle c'r reddish
rvitli biack rnarkings. N'llrn¡'res;ernble lerdy bee-
tles (Coccinclli<1ae) but dif'ler þry haring the'
head readily visible from above and the anterior
corners of the pronotum generally extende
forward. The tarsi may appear 3- or 4'seg:
mented. M ost specles occur on fungi in decaY'
ing wood o r lruit and under bark; they feed
fungi.
Common Species
Ap ho ris ta vlllala (Fabricius ) (Fie. l9?A)-5
brownish red or orange; flat pronotum with black
edges , sometimes 2 indistinct brownish spots; elytra
with long centr I and 2 shorter lateral black
most of U.S., outhern Canada
Endomychus biguttatus SaY (Fig. 197
5.0 mm; head, Pronotum and legs shiny
reddish or orange with 2 smaller black spots
middle and 2 uch larger patches toward apex;
of U.S. and Canada.
Family Byturidae-
Fruitworm Beetles
The small (3.5-4.5'mm), oval, hairy beetles
this family are orange or pale brown. in
The second and third tarsal segments are
ventrally. There are only five species in the U
Byturellus grisescens (Jayne) may be swept
from oaks in California and Oregon, Byturus
rubi Barber (Fig. l9?C), the eastern raspberry
fruitworm, is brownish yellow with dense'
yellow pubescence. The adults of thís
and B. bakeri Barber, the western rasp
fruitworm, occur on flowers of
blackberries and the larvae fecd on the
Order
 are shiny and cylindrical; they are s
called false wireworms and resemble click bee-
tle larvae (Elateridae).
Members of the genus Eleodes (Fig. 198)
o
are large black beetles that angle their abdo-
mens upward when disturbed and may emit a
o
foul-smelling black fluid. There are about 100 @
species of this western genus, many of which
are quite common in semiarid and desert areas. €
The genus Diaperis contains colorful, spotted
species that resemble lady beetles (
lidae).

Figure 197 A and B, handsome fungus beetles (En-

domychidae): A, Aphorista v¡ttala; B, Endomychus
bigultatus. C, eastern raspberry fruitworm, Byturus
rubi (Byturidae).
\
Family Tenebrionidae-
Darkling Beetles
The tenebrionids are a large family of beetles
with about 15,000 species worldwide and 1,400
species in North America. The many common
species tend to be dull black or brown and often
resemble ground beetles (Carabidae). Darkling
beetles have a 5-5-4 segmentation, the eyes are
usually notched rather than entirely round, and
the I I -segmented antennae are generally mon- Figure 198 Eleodes spp.of darkling beetles
iliform or filiform. The body varies from (Tenebrionidae).
broadly oval to elongate, and smooth to very
rough. These beetles are more common in the
Common Species
arid western states, occurring under rocks, de-
Alobales pennsylvønicø (De Geer) (Fig. l99A)-
bris, and loose bark. Larvae of many species 20-23 mm; black; posterior corners of pronotum
pointed; most of U.S. (under bark).

218 Subclass Pterygota

t
r
Bolitotherus ('ornulus (Panzer) (F it¡ 1998)- I0-
l2 mm; dark brown to duÌl black; bod¡ surfacc er-
tremcly rottgh;protloturn with 2 horns (f ) or tubcr-
cles ( Q ); clytra ',vith 4 rorvs large tuber:lcs; easter n
l/2 U.S. (in old iungi on logs).

Neomida bicornis (Fabricius)-3-4 mm; elongate-

oval; dorsal side metallic bluish green; pÍonotum oc-
casionally brownish; front of head u'ith 2 short
horns and top with 2 longer horns (ct); throughout
North America (on old fungi).

Phloeocles pttstulosus (LeConte) (Fie' l99C)-15-

23 mm; flattenecl; grayish black with highly rough-
enerl and hardened dorsum; elytra sometirnes rvhit-
ish at base and apex; CA'

Plaf ydema excavatum (Say) (Fig. l99D)-4' 5-

5.5 mm; broadly oval; shiny black; antennae and
legs dark reddish brown; 2 horns between eyes (O)
with concave area between horns, or head with tu-
bercles (Ç ); elytra deeply grooved; most of U'S'
(under bark). P. ruficorne (Sturm) similar, without
horn, dull black, antennae pale reddish yellow, east-
ern l/3 U.S. P. etlipticum (Fabricius) without horn,
2 dull red spots at base of elytra, eastern 1/2U.5.

Tenebrio mol¡tor Linnaeus (Fig' l99E)' Yellow

mealworm-13-16 mm; shiny blackish or dark red-
dish brown; deeply grooved, intervals between
grooves dcrrsely punctured; thror-rghotrt North
America (in storcd grains, meal, llour)', T obscurus
Fabricius, clark nlealworm, similar, dulì'color, inter-
vals between elytral grooves granular' Neatus
tenebroides (Palisot de Beauvois) similar, eastern
l/2U.5. (under bark).
Tribolium confusum Jacquelin duval (Fig' l99F)'
Confused flour beetle-4.5-5.0 mm; flattened; red-
dish enla club; F
thro meri food,
mus T. , red
flou ante tly at Figure 199 Darkling beetles

apex.
A, Alobates PennsYlvanica; B, Bolitotherus
nutus; C, Phloeodes Pustulosus; D, Plalydema
cavatum; E, yellow mealworm, Tenebrio
IJloma punctulal0 LeConte-7.0-8.5 mm; pale red-
F, confused flour beetle, Tribolium confusum,
dish brown; last segment antennae rounded at tip;
eastern U.S., chiefly South (under pine bark)' U' im-
pressa Melsheimer similar, ll-12 mm, head-with
ãeep curved impression, eastern U.S. (under oak and
beech bark).

Order Coleo¡rtera
Family Alleculidae-
Combclawed Beetles
These brownish or black, elongated and convex
beetles resemble darkling beetles (Tene-
brionidae) but have a row of small, blunt teeth
on each tarsal claw which darkling beetles lack
(Fig. l65A). Alleculids are 4-12 mm long and
the body often is glossy due to its pubescence.
Adults occur on llowering plants, ftrngi, and
under bark, and many probably are pollen
feeders. Larvae resemble the false wireworms
(Tenebrionidae).
Common Species
Hymenorus nþer (Melsheimer)-5-7 mm; black; an-
terior part of elytra brown to brownish black; legs
brown to yellow brown; elytral grooves disappear
posteriorly; eastern U.S. south to FL, west to MN
and TX, southern Canada.
Figure 200 A, a combclawed beetle, lsotttra
sericea (Alleculidae); B, a longjointed be:tle,
I h ro ma c ra ae n ea (Lagr lidae).
220 Subclass Pterygota
Isomira sericea (Say) (Fig. 2004)-5-6 mm; pale
brownish yellow; dense line pubescence; 4th segment
of maxillary palpi long and slender; eastern l/2U,5. ,
Family Lagriidae-
Longjointed Beetles
These slender beetles are l0-15 mm long and
a dark metallic color. The 5-5-4 tarsal segmen-
tation and enlarged apical segment of the an'
tennae help characterize this family. ¡\dults oc-
cur and feed on leaves or sometimes are found
under bark.
Common Species
Arthromacra aenea (Say) (Fie. 2008)-9-14 mm;
metallic green, coppery blue, or dark brown; anten'
nae reddish brown; vçntral side dark bronze; eastern
U.S,, southeastern Canada.
Family Salpingidae-
Narrowwaistecl Bark Beetles
The pronotum abruptly narrows posteriorly in
many species of this family and the general
body shape and black color resçmble the
ground beetles (Carabidae). Body length.varies
from 3-30 mm. Adults occur under bark, rocks,
litter, and on vegetation; adults and laryae are
predaceous.
Common Species
Pytho niger Kirby (Fie. 20lA)-5.5-6.0 rnm; black¡
pronotum flattened; head and pronotum with longi'
tudinal depression; elytra grooved, many punctures,
base with 2 depressions; northern l/2U.5', Canada.
Famil y Mycetophagidae-
Hairy Fungus Beetles
These small (1.5-6.0 mm long), hairy beetles are
flattened and broadly oval. They are brown to
black and may be marked with red ctr orange.
Members appcar on shelf ftrngi and under
nroldy balk a¡rd vegetation, app,lr,:rtrly fr:eding 'L)
on f unfli .
/r-
 Common Species
Cis Juscipes Mellie (Fig. 20lC)-2.0-5.5 mm;
to dark reddish brown; eastern l/2U.5'

Family Pyrochroidae-
Firecolored Beetles
These flattened beetles have soft elytra, the
heacl is narrowed between the large eyes, and
many have prominent serrate or pectinate an'
tennae. The family name is derived from the
black color with contrasting red or yellow
nota. Adults occur on vegetation
daceous larvae live under bark.

Common Species
Dendroides cyanipennis Latreille (Fig
l3 mm; head, antennae, and elYtra blackish; rest
body reddish yellow; most of U.S.' so'
Canada.

Figure 201 A,
nlger (Salpingi
Mycetophagus
minute treefung

Common Species
Mycetophagus punctatus Say (Fig' 20.18)-4'0'
S.-S mm; blãckisir with ieddish yellow markings;
an-
tennae red-brown becoming blackish at apex; east-
ern l/2 U.S.

Familv Ciidae-
Minute Treef ungus Beetles
The cylindrical body of these brownish or black
beetles is 0.5-6.0 mm long. The heac is not visi-
ble from above and the antennae h¿ ve a 3-seg-
mented clrrb. Thcse beetles resemblr the Scoly-
tidae and Bostrichidae and,lci:ur unrler bark, in
decaying wood, or in dry, hard frrngi' Ciids
feed on lungi.
Figure 202 A, a firecolored beetle,
cyonipennis (PYrochroidae); B' w
N acerdes me la nu ra (Oedemeridae).

Order Coleoptera
Neopyrochroa flabellata' (Fabricius)-15-17 mm;
head, basal portion of antennae, thorax, and legs
reddish yellow; elytra black; most of U.S., sourhern
Canada (in woods).

Family Oedemeridae-
Oedemerid Beetles @
Oedemerids arc 5-20 mm long, have soft elytra,
and the tarsal segmentation is 5-5-4. These &
slender beetles resemble the more common blis-
ter beetles (Meloidae) but differ partly because
the next to last tarsal segment is expanded and
very hairy beneath. Adults occur on flowers or
foliage and are attracted to lights. Adults are
pollen and nectar feeders; larvae live and feed
in damp decaying wood (especially conifers).

Common Species
Nacerdes melanura (Linnaeus) (Fig. 202tìr, Wharl
borer-8-12 rnrr; dull yellow; elvtra deep rurpìe at
apex, u'ith 8 clcvated longitudintrl line:ii lr.gs and
ventral surlacc' blackish; throughoul U S, southern
Canada (in lunrberyards, woodsheds).

Figure 203 False darkling beetles (Melandryidae).

Family Melandryidae- False A, Eustrophinus bicolor; B, Osphyo variqns;
Darkling Beetles or C, Penthe obliquata ü
Melandryid Bark Beetles \

These dark-colored beetles are 3-20 mm long Penthe obliquata (Fabricius) (Fig. 203C)-l1,5-
and often slightly flattened and. pubescent. 14.0 mm; velvety black; cutellum with long yellow-
orange hairs; eastern l/2U.5. P. pimelio (Fabricius)
They have a 5-5-4 tarsal segmentation and usu- sim ilar, scutellum black
ally two dents near the posterior margin of the
pronotum. Adults and larvae occur under dry
bark and in dry wood and fungi; a lew adults
frequent flowcrs and leavcs.

Common Species
Eustrophinus bicolor (Fabricius) . (Fig. 2034)-
4.5-6,0 mm; convex; shiny black; antennal base and
apex, legs, and ventral side reddish brown; each l/2
posterior margin of pronotum distinctly 'rS"- The arched body, slightly shortened elytra, and
shaped; most of U.S. sharply tapered, pointed abdomen readily iden-'
tify members of this family. They are commonl
Osphya varians (LeConte) (Fig. 2038)-5-8 mm; on flowers and jump or tumble when disturbed.
black; margin and center of thorax reddish yellow;
.

sparse, fine, gray prostrate hairs; most of U.S. The adults and most larvae are plant feeders.

222 Subclass Pterygota

.l

Ficure 204 Tumbling flower beetles (Mordellidae). A, Clipa oculota; B, Mordella

má rgina ta ; C, M ordet I istena p' t s tu Ia ta'

Common SPecies larvae are predators ofgrasshopper eggs or

Clipa octopunctata (Fabricius)--6-7 nr n; black or parasites ol bee eggs and l:rrvat:. The active,
gráyish black; pronotum with irregulr r yellorvish parasitic larvae gain access to be': nests by at'
pubescence on margins; elytra with 8 irregular spots taching themselves to foraging bt:es; they then
ôf yellowish pubescence; easter¡ì U'S' west to KS
and more seden'
develop into several distinct
and TX. (hypermetamorphosis
tary forms
Glipa oculata (Say) (Fig' 20aA)-5-7. mm; head' contain a blistering substance,
thorax, and abdomen blackish with yellow and gray This material is extracted from
markings; antennae, tibiae, and tarsi dull red; east- southern EuroPe, the SPanishflY,
ern l/2 U.S. drug. Some species will secrete blistering
rials or oily substances (latter in genus
Mordella atrata Melsheimer-3-6 *t) ¿ult btutt;
brownish pubescence; margins of sternites. gray; as a defensive action.
."r,.rn l/i U.S' M. albosuturalls Liljeblad similar'
3-5 mm, base and inner margin of each elytron
with
Comm n Species
silvery pubescence; western l/2 U'S'
Epicau ta atbida (Say)-la'20 mm; yellowish
grayish; 2 dark longitudinal lines on pronotum;
Mordetla marginata Melsheimer (Fie' 2048)- segment of antennae as long as following 4
3.0-4.5 mm; black or dark gray; silvery pubescence ments; western l/2 U.S
scattered as short lines in various arrangements on
pronotum and elytra; Rocky Mts' westward' Epicauta maculato (Say)' Spotted blister
tle-10-14 mm; black; gray pubescence; mal¡y
Mordellistena ) Gig' 20aC)- spots on elytra; Far West'
2-3 mm; black lverY Pubescent
spots across el band; through-
Epicauta pennsYlvanica
out North America. Black blister beetle-7-15
sparse pubescence; most of U.S.'
(on goldenrod, garden flowers)
Family Meloidae- Blister Beetles
The blister beetles are elongated beetles with a Epicau ta vittata (Fabricius) (Fig' 2058)
prothorax that is often narrower than the head blister beetle-12-18 mm; du yellow above
and sometimes necklike. The wings and body
are soft and frequently the tip of the abdomen
is exposed. Adults are plant feeders and many
Order ColeoPtera
 :,ii,"
iffffi
'i
wi
s¿l.t as long as or long,gr than the
tibiae) occur on flowers and foliage, li
onthe dafewfrequentsanddunes.
f', O
Common Species
A nthicus cery'inus LaFerté-Sénecteie-Z.4-2.7 mm; Ð
reddish Lrrown; antennae and legs dull yellow; elytra
with blackish rnarkings on apical l,z2 which enclose
a broad pale yellow cross; throughout North Amer-
0
lca. Êù.

Notoxus monodon Fabricius (Fig. 206A)-2.5-

4.0 mm; dull brownish yellow; pronotal horn pres-
ent; dense grayish hairs; elytra with black crossband
on apical l/2, 2 black spots near base; head and
sides of pronotum sometimes blackish; throughout
North America, N, conslricto Casy similar, pronotal i
horn forked, CA (in orchards).
ir

Figure 205 Blister beetles (Meloidae). A, Epicauta

pennsylvanica; B, E. vittata; C, Meloe anguslicollis.

Lytta cyanipennis (LeConte)-13-18 mm¡ metallic

green or bluish; Rocky Mts. westward. L.' nuttattii
(Say), Nuttall blister beetle, similar, l6-28 mm.

Lytta immaculata (Say)-25-27 mm; black; gray or

yellowish pubescence; Midwest to Rocky Mts.

Meloe angust¡collis Say (Fig. 205C)-12-15 mm;

head and pronotum blue-black; elytra and abdomen
blackish with violet tones; elytra short, wrinkled,
overlap, and point outward; northern U.S., Canada.

Family Anthicidae-
Antlike Flower Beetles
Anthicids are 2-6 mm long and somewhat ant-
like in appearance. The greatly lowered head
with its necklike posterior region and the 5-5-4
tarsal segmentation help distinguish the family.
The pronotum of some extends hornlike over
the head (i.e., genus Noloxus with the tarsus
shorter than the tibiae, and genus Mecynotar-
Figure 206 A, an antlike flower beetle, Nofoxlæ
monodon (Anthicidae); B, a pedilid beetle, Pedilus
lugubris (Pedilidae).
Family Pedilidae- Pedilid Breetles
Members of this family resemble the An-
thicidae but are more elongate (4-15 mm long)

224 Subclass Pterygota

and cylindrical. The head narrows ,)osteriorly
into a neck and the prc'notum s o;al or
rounded ,\dtrlts lrcquent l-l trvers tlr l-c,liagc
and are attractcci to Iights. lvlcst oI thc r;f'ccics
occur in thc western states.
Common Species
Pedilus lugubris Say (Fie. 2068)-6-8 mrn; black ex-
cept lst 2 antennal segments and front of head red-
dish brown; most of U.S., central Canada.
Family Cerambycidae-
Longhorned Beetles or
Roundheaded Wood Borers
Cerambycicls are usually medium to large,
elongated and cylindrical beetles with antennae
over half the body length (the antennae of some
species are much longer than the body). The
eyes are notched (Fig. 2074) and many species
are brightly colored. Some smaller cerambycids
resemble the leaf beetles (Chrysomelidae) but
the leaf beetles are smaller (usually less than
12 mm) and the antennae are less than half the
body length. The longhorned beetles are a large
family with about 20,000 species worldwide and
over 1,200 in the U.S.
Most adults occur and feed gn flowers
and foliage during the day; some eàt bark or
stems. Numerous species are nocturnal, resting
under bark or on trees during the day, and are
often attracted to lights at night. Adults prefer
to oviposit on solid wood of weakened, dying,
dead, or freshly cut or fallen trees. Larvae are
cylindrical, whitish, and have a rounded head-
thorax area as contrasted with the flatheaded
wood borers (Buprestidae). Cerambycid larvae
bore into and feed on wood. Only a few of the
common species are described below.
Common Species
Subfamily Prioninae-pronotum with distinct
margin on sides and often margin is toothed; up
to76+ mm long.
Orthosoma hrunneum (Forster)-22-48 nrnt;
light rcddi,,h l¡rown; deep im¡lresr;ions bot\veen an-
rcrìn¡r: .ì I rc,rth on lateral nrargins of ¡tronotum;
('ast' r'n l; .. . si)utheastcrn (.an¿r ia
Prionus culi.lornicus N4otschulsk¡. (lalif o¡ nia pri-
onus--.10-60 rnm; shiny; dark reclclish brown; 3
sharp teeth on lateral margins of pronotum; Far
west. '
'
Prionus laticollis (Drury) (Fig. 2074). Broadnecked i
root borer-22-47 mm; sñiny black; antennae, j
reaches apical l/2 of elytra (o) or l/2 or less bodÿ-:i
length ( Ç ); prgnotum as wide as base of elytra, prô-:i
notum with 3 lateral teeth; eastern l/2 North Amer-'1
ica. P. imbricornrs, tilehorned prionus, similar, dark
reddish brown.
Subfamily Lamiinae-last segment of maxillary
palpi pointed at tip.
Monochomus maculosus Haldeman (Fig. 2078).
Spotted pine sawyer-14-27 mm; dark reddish
brown to blackish; bluish gray markings on',iytrþi
antennae up to 2 l/2xbody length; pronotum $,ith
large tooth on each side; throughout U.S.
Far West), southwest Canada.
Monochamus litillator Fabricius. Southern
sawyer-20-30 mm; brown; elytra mottled
gray, brown and black pubescence; pronotum
large tooth on each side; apex of each elytron
spine; most of U.S., southern Canada.
Saperda calcarata Say (Fig. 207C). Poplar
er-25-3 I mm; dense covering of gray and yellow
pubescence; front of head and scutellum yellow;
pronotum with 3 ycllowish longitudinal stripes;
coarse punctures; tip of each elytron spined¡
throughout North America.
Saperda candida Fabricius. Roundhe¿ded
borer-15-20 mm; light brown with 2
stripes from head to elytral apex; ventral side
eastern U.S., Canada.
Tetraopes tetrophlhalmus (Forster). Red
beetle-9-I4 mm; red-orange; pronotum witb
black spots; elytra with 6 black spots; eastern'
North America (on milkweed). T.
Schönherr similar, 2 wide crossbands on
southern U.S. T. Jemoratus LeConte
l7 mm, antennae ringed with gray, 4 elytral
throughout U.S.
OrderGol
 o
@
&,

fiI

Figur e 207 Longhorned beetlc; or roundheaded rvood borers (Ceran-ibi,cidire) . .\"

broadnecked root borer, Prionu: laticollis, and a notched eye; B, spotled ¡rinc sa+vyr:r,
Monoc'hamus maculosus; C, poplar borer, Saperda calcarsta: D, Strøngalia Jamelica;8,
T.vpocerus velutinus; F, Callidiultt anlcnnutLotl hesperurn; C, Iocust borer, Megacyllene
robin¡ae, H, redheaded ash borer, Neoclytus acumtnatus.

226 Subclass Pterygota

Subiamil¡' l.eptrrrinae-apex of last segmerlt of
maxillary palpi bÌunt, elytra ot'ten gleatly nar-
rowed posteriorly, eyes oval or only slightiy
notched.
Enaphalodes rufulus (Haldeman). Red oak bor-
er-22-28 mm; reddish brown, covered with brown-
ish orange pubescence; pronotum usually with 2
small tubercles near center; apex of each elytron
with 2 short spines; eastern U.S. west to MN and
TX.
Strangalia famelica (Newman) (Fie. 20'7D)-12'
l4 mm; very slender; brownish yellow with yellowish
pubescence; pronotum with 2 black stripes; black
spots on elytra;eastern U.S,
Strophiona /rrc¡ø (L,eConte)-10-13 r¡m; yellow or
golden; 3 *icle crossbands on el¡'tra; Far \\/est (near
oaks ).
Typocerus velutinus (Olivier) (Fig. 207E)-8-14 mm;
head, antennae, and pronotum black; pronotum
with yellow pubescence; elytra reddish brown with
yellow crossbands; eastern U.S.
Subfamily Cerambycinae-apex of last seg-
ment of maxillary palpi blunt, elytra not greatly
narrowed posteriorly, eyes distinctly notched
and partly surround base of antennae.
Callidium anlennatum hesperum Casey
(Fig. 207F)-9-14 mm; metallic blue or þlue-black;
rounded thorax with 2 depressions; lbgs black,
femora large; throughout U.S. (early spring)'
Megacyllene robiniae (Forster) (Fig' 207G). Locust
borer-12-18 mm; black; head, pronotum, and ely-
tra with yellow crossbands as shown; Rocky Mts.
eastward (on goldenrod in fall).
Neoclytus acuminatus (Fabricius) (Fie' 207H)' Red-
headed ash borer-6-18 mm; head, thorax, and
middle and hind legs reddish; elytra light brown with
darker apex, ,l crossbands of fine yellow hairs; east-
ern U.S., southeastern Canada.
Family Chrysomelidae-
Leaf Beetles
Leaf beetles are a large family of very common
and often brightly colored insects. There are
over 25,000 species worldwide and about 1,460
species in North America. The body shape of
many resembles lady beetles (Coccinellidae) but
the tarsal segmentation of lady beetles is 3-3-3
whereas that of leaf beetles appears as 444,, :.
Leaf beetles also resemble some longhcrned :
beeties (Cerambycidae) but the former are usu--
ally smaller (l-12 mm long in the U.S.) and the
antennae are nearly always less than half
body length.
Adults feed on flowers and foliage;
larvae eat foliage (includes leaf mining)
others bore into stems and roots. Larvae ten$
to be soft-bodied and brightly colored. Some
bear spines or are partially covered with their
own excrement. Many chrysomelids are impor-
tant agricultural pests. Only a few of the com-
mon species are described below.
Common Species
Subfamily Cassidinae-Tortoise Beetle$;
broadly oval to circular, head usually concealed
from above, some resemble small turtles or lady
beetles, many brilliantly colored, some
tened, often on morning glorY'
Agriconola bivittata (Say) (Fig. 2084)-4.5-6.0 mm;
pronotum pale yellow with large brdrvn triangular
area; each elytron pale Yellow with 5
longitudinal stripes; most of U'S.
Chelymorpha cassidea (Fabricius) (Fig.
Argus tortoise beetle-8.0-l I .5 mm; reddish or
low; pronotum with crossrow of 4 black dots and
larger ones posteriorly; elytra with 13
most of U.S
Deloyala suttata (Olivier) (Fie. 208C).
toise beetle-5-6 mm; margins of pronotum and
tra translucent pale yellow cxcept at anterior corners
and apex of elytra; elytra blackish with irregular yel'
low patches; most of U.S.
.
Metriona bicolor (Fabricius) (Fig' 208D). Golden
tortoise beetle-S-6 mm; brilliant brassy gold when
alive but dull reddish yellow when dead; margins of
pronotum spread out and flattened; elytra with
round, depressed spots; eastern U.S.
Order Coleoptera
 o
C
Ð
@
ø'

Figure 208 Lcaf beetles (Chrysomelidae). A, Agriconota bivittuta; B, argus tortoise beef le,
Chel¡'rnorpha cassidea; C, mottled tortoise beetle, Delo.yakt guttata; D, golden tortoise bee-
tle, iÝ[etriona bicolor; E, odontota dorsalis; F, pachybrac,års sp.;G, cryptocephølus sp.;
H, Babia quadriguttata; l, Neochlamisus gibbosa.

228 Subclass Pterygota

Subfamily Hispinae-Leafmining Leaf Beetles;
4-7 mm, mouth directed downward or poste-
riorly rather than forward, elytra often dis-
tinctly ridged and with punctures between
ridges; larvae are leaf miners.
Microrhopala vittata (Fabricius)-5-7 mm; head and
pronotum reddish; elytra blackish or brown, with 2
reddish longitudinal stripes, 8 rows of punctures;
eastern U.S. (on goldenrod).
Odontota dorsalis (Thunberg) (Fig, 2088)-6.0-
6.5 mm; orange-red with head and middorsal elytral
stripe black; elytra ridged; eastern l/2 U.S. (on lo-
cust trees). O. scapularis (Olivier) similar shape,
pronotum with 2 lateral stripes, anterior corners ely-
tra reddish, AZ eastward.
Subfamilies Chlamisinae, Clytrinae, and Cryp'
tocephalinae-Casebearing Leaf Beetles; pro-
notum covers head up to eyes, elytra expose last
abdominal segment, larvae carrl protective
case often of own excrement. Mem >ers of ge-
nus Pachybrachis (Fig. 20tìF) have pronotum
with distinct posterior marg in anc non-scal-
loped lateral margins; those in genus Cryp-
tocephalus (Fig.208G) have pronotum without
distinct posterior margin and lateral margins
are weakly scalloped,
Babia quadriguttata (Olivier) (Fig' 208H)-3.0-
5.5 mm; shiny black; 2 reddish spots at base and
apex of elytra; antennae short and weakly serrated;
throughout U.S.
Figu re 209 Leaf beetles (Chrysomelidae). A, threespotted flea beetle, Disonycha triqngu'
laris ,'8, potato flea beetle, Epitrix cucumeris; C, Systena blanda
Neochlamisus gibbosa (Fabricius)
2.5-4.5 mm; bronze-brown, elytra
ish; pronotal surface rough, large 2-pronged
cle in center; elytra with many tubercles; pulls
and rolls off plants if disturbed; eastern U,S.
ti
l,'
Subfamily Donaciinae-metallic green,:
coppery, or black, long antennae, adults
larvae on aquatic or shoreline plants like
lilies or sedges; common genus $
(Fie.2l0A).
Subfamily Alticinae-Flea Beetles; dark blue,
green, or black, jumping behavior, hind femora
enlarged, adults eat many tiny holes in leaves
Chaetocnems pulicariø Melsheimer. Corn flea
tle-I.5-2.5 mm; shiny black with bronze or
green luster; basal 34 antennal segments
tibiae and tarsi brownish yellow; most of U.S:
corn, grasses).
Disonycha triangula r¡s (Say) (Fig. 209A).'lhreespot'
ted flea beetle-4.5-7.0 m; shiny blu or blue-
black; pronotum yellow with I black dots; most of
U.S. D ranthomelas (Dalman), sf ilì:,ch fle¡ beetle,
sirnilar but pronotum without dots.
Epitrix cucumeris (Harris) (Fie. 209E1). Potato flea
beetle--1.5-2.5 mm; shiny black; hairy; antennae
and legs brownish orange; throughotrt U.S. E. hlr'
tipennis (Melsheimer), tobacco flea beetle, similar
shape, dull reddish yellow, sometimes darker band
across middle of elytra
Order Coleoptera

Systena blondo Melsheimer (Fig.
209C)-3.0-
4.5 mm; head and prothorax reddish brown; elytra
pale yellow, brown, or blackish, and with 2 lighter
stripes; throughout U.S,
Subfamily Criocerinae-head narrowed poste-
riorly to form neck, elytral punctures in rows.
Crioceris asparagi (Linnaeus) (Fig. 2l0B). Aspara-
gus beetle-ó-7 rnm; lÌattened; shiny; head dark me-
tallic blue; pronotum red, 2 bluish spots; elytra blu-
ish green with ycllow patchesi nrost of U.S. (on
asparagus). C riuoclecimpunctata (Linnaeus), spot-
ted asparagus beetle, similar shape, head and pro-
notum reddish brown, each orange-red elytron with
6 black spots, eastern U.S.
Lema trilineala (Olivier). Threelined potato bee-
tle-6.0-7.5 mm; reddish yellow; 2 small black spots
on pronotum; elytra with 3 longitudinal black
stripes; pronotum distinctly constricted in middle;
eastern U.S. (on potatoes).
Subfamily Chrysomelinae-broadly oval, con-
vex, head inserted into pronotutn up to eyes,
brightly colored. Phyllodecta species are small,
metallic blue or purple, feed on willow and
poplar.
Calligrapha phìladelphica (Linnaeus) (Fig. 2l0C)-
8-9 mm; very convex; head and pronorum dark olive
green, metallic; antennae and legs dar( reddish
brown; elytra yellowish white with dark malkings as
shown;eastern l/2 U.S.
Chrysomela scripta (Fabricius) (Fig. 2l0D)-7.0-
9.5 mm; prothorax blackish with orange lateral
margins; each yellowish elytron with 7 purplish
spots, pattern variable; most of North America (on
willow, poplar).
Leptinotarsø decemlineata (Say) (Fig. 2l0E). Col-
orado potato beetle-5.5-ll mm; dull yellow; pro-
notum spotted; elytra with l0
longitudinal black
lines as shown; throughout U.S., southern Canada
(on potatoes; larva humped, brick red).
230 Subclass Pterygota
i,
t¡
i:'
i.t
,i
n{
äÉ
Subfamily Galerucinae-soft elytra, pronotum, f:.i
I:
usually with distinct lateral margins.
Acalymma trivittatum (Mannerheim) (Fig. 2l0F).
o
Western striped cucumber beetle-4-ti mm; pro-
notum orange-yellow; elytra pale yellow with 3 black
i*,O
longitudinal stripes; base of antennae and femora
pale yellow; Far West. A. vittatum (Fabricius),
r.o
striped cucumber beetle, similar, antennae all black, ¡',(Ð
eastern U.S. west to CO and NM, eastern Canada.
Ceratr¡ma trifurcala (Forster)-4.0-8.5 mm; yellow
or dull red; head black or blue-black; elytra with
blackish border and 3 pairs large blackish spots,
spots sometimes connected or nearly absent; east of
Rocky Mts
Diabrotica longicornis (Say). Northern corn
worm-5-6 mm; pale green or greenish yellow;
sometimes reddish brown tinge on head and thorax;
east of Rocky Mts.
i
Diabrotica undecimpunctata howardi Barberl
(Fig. 2l0C). Spotted cucumber beetle, southern corfir
rootworm-6-7 mm; greenish yellow; head black;l
elytra with 12 black spots; bases of antennae and
legs pale yellow; most of U.S. (primarily eastern).
D. u n d e c i m p u n c t a t a u n d ec i m pu ncfølc lvf annerheim,
western spotted cucumber beetle, similar, antennae,
legs, and body all black, Far West.
Subfamily Eumolpinae-convex, oblong, 3rd:
tarsal segment bilobed, metallic or yellowish
and spotted
Chrysochus auratus(Fabricius)-8-ll mm;
l/2U.5.
cent blue-green with coppery tinge; eastern
(on milkweed, dogbane). C. cobaltinu,s LeConte
similar, 9-10 mm, darker metallic blue, Far West.
Colaspis brunneq (Fabricius) (Fig, 2l0H). Grape
colaspis-4-6 mm; dull brownish yellow; outer seg-
ments antennae often black; legs pale_yellowish;
most of U.S. (on grapevines, cotton).
Paria fragarjae Wilcox (Fig. 2l0I). Strawberry
rootworm-3-4 mm; color and markings varia-
ble; head reddish yellow; pronotum black ,or
yellow; elytra black, yellow, .or yellow with 4
(sometimes 6) black spersi :
America.
Figure 210 Leaf beetles (Chrysomelidae). A, Donqcia sp.; B, asparagus beetle, Criocerìs
asiaragi; C, Cattigrapha philadetphico; D, Chrysomela scripla,' E, Côlorado potato beetle,
Làptinotarsa decemlineata,' F, western striped cucumber beetle, Acalymmq trivittatum;
C, spotted cucumber beetle or southern corn rootworm, Diabrotica undecimpunctots how'
ordi;H,grapecolaspis,Colaspisbrunnea; I,strawberryrootworm' Pariafragariae.

Order Coleopt€ra
Family Bruchidae-Seed Beetles
These oval or egg-shaped beetles are generally
less than 5 mm long and their elytra do not
cover the tip of the abdomen. The head is ex-
tended into a short, broad snout and the anten-
nae may be clubbed, serrate, or pectinate.
Adults occur on flowers, leaves, and seed pods,
and both adults and larvae feed inside stored
seeds such as peas and beans.
Common Species
Acanthoscelides obtectus (Say) (Fig. 2llA)-2.5-
4.0 mm; blackish with brownish gray pubescence;
antennae black with red-brown base and rip; elytra
with short crossbands ol brown pubescence;
throughout North America.
Bruchus pisorum (Linnaeus). Pea weevil-4-5 mm;
black with dense reddish brown and whitish pubes-
cence; pronotum with small whitish triangle at mid-
dle of posterior margin; elytra with grayish, yellow-
ish, and whitish patches, irregular white crossband
on posterior I /2 ol elyta; whirish patch at tip of ab-
domen; most of North America (on peas in spring).
Family Anthribidae-
Fungus Weevils
Fungus weevils are generally less than¡I0 mm
long and have a short, broad, and flat beak.
The antennae are clubbed (but not elbowed) or
sometimes filiform. The.tip of the âbdomen is
Figure 2l I A, a seed beetle, Acanthoscelides
Ettpariu.s marmoreus (Anthribidae); C, oak timberworm, Arrhenodes
232 Subclass Pterygota
not exposed as with the similar seed beetles
(Bruchidae). Anthribids occur beneath old
bark, on dead branches, and sometimes in
weedy fields. Wood, fungi, seeds, and pollen
serve as food for larvae and probably adults to
some extent
Common Species
Araecerus fasciculatus (De Geer). Coffee bean wee-
vil-2.5-4.5 mm; dark brown with yellow or brown
pubescence; antennae and legs reddish brown;
through out most of North America (in seeds,
ries, dried lruit).
Euparius marmoreus (Olivier) (Fie. 2llB)-3.5-
8.5 mm; yellowish brown and pubescent; dense pale
brown and grayish yellow pubescence on snout, an.
terior l/2 pronotum, and a patch in center
posterior part of elytral patch with black spots;
ringed with gray and dark brown; easternl/2
America.
Family Brentidae- Brentid Weevils
These elongated and cylindrical weevils are l0-
30 mm long and have a snout that extends for"
ward and does not curve downward. Adults oc-
cur under bark or in damaged wood and feed
on fungi, sap, or insects. Five of the six North
American species are confined to Florida,
Texas, and California, and the sixth species in-
habits the eastern U.S.
obtectus (Bruchidae); B, a fungus weevil,
(Bintidae).
minutus
Common Species black turpentine beetle, similar, 5-10
dish brown to black, eastern l/3 and
,

Arrhenodes minutus (Drury) (Fig, 2l lC). Oak tim-

berworm-7- l7 mm; dark reddish brown; elytra
with narrow and elongate yellowish spots; beak Ips grandicol/u (EichhofÐ (Fig. 2l2A).
ine engraver-2.8-4.7 mm; dark reddish
longer than head and very slender ( Ç ) or as long as
(d); eastern U.S,
black; tip of elytra deeply co cave, punctured,
head and broad
5 teeth on each side; eastern l/2 North Ameriça
pines). /. calligraphus (Germar) similar, 3'5'6.5
elytral concavity with 6 teeth on each side,
Family Scolytidae- Bark Beetles
Bark beetles are generally l-3 mm long (some Scolytus rugulosus (Ratzeburg) (Fig' 2l2B).
reach 9 mm in length), very cylindrical, and hole borer-l .1-2.9 mm:' grayish black; elytra
dish brown at apex; most of U.S., primarily
brown or black in color. The antennae are (chiefly on fruit trees). S. multistr¡qtus
clubbed and elbowed (Fie. l60C) and the head smaller European elm bark beetle, similar,
may be lowered and not visible from above' ventral side abdomen concave and with a
The prothorax is very large and often the poste- American elm.
rior abdominal segments are curved downward
X-vleborus sa.uesensi (Ratzeburg)-- 1.5-2.5 mm; pale
to form a broad concavity (Fie. 2.1A). Adults brorvn to black; scutellum conical ( ()); tip of elytra
and larvae live under tree bark and either sLecpll' slopcd, rvith rows of ¡roinrcd gra.rules and 2
(l) mirte thc wood's surfat:c atrd ti-e undcrside Irrl)crclcs ìn tcnter; humpbacked atlpearance (Ct);
ol the bark (bark or engra\cr bectl, s), (2) bore nìo\1 (rl L,i S . southcrn (lanacla
into and leed on the wood, cr (3) core cleeply
into the wood to cultivate and l'eerd orr iungi
which will grow on the tunnel wallr; (ambrosia
or timber beetles). The larvae of surlace feeders
chew a maze of characteristic tunnels (galleries)
on the wood and bark, thus creating an en-
graved appearance. The new adults emerge
through tiny holes they have chèwed in the
bark. Bark beetles as a group are olten consid-
ered to be the most destructive forest insects'
The Dutch elm disease of American elms is
transmitted by Scolytus multistriatus (Mar-
sham).

Common Species
Dendroctonus frontalis Zimmerman. Southern pine
beelle-2.2-4,2 mm; dark brown to black; front of
head with 2 tr¡bercles separated by a groove; elytra
over 2x length of pronotum, not concave at tlps'
long hairs on sides and tips; southern U'S' west to
ÃZ (on pine). D. brevicomis LeConte similar, head
tubercles smaller, area near front margin of pro-
notum with narrow transverse elevation (Q ) or de-
pression (cr ), Far West.

Dendroctonus valens LeConte. Red turpentine bee-

tle-5.5-9.0 mm; light reddish brown to dark brown;
most of U S. except southeastern and Culf coast
states (on ¡rines primarily). D terebrøns (OÌivier),
Figure 212 Ba rk beetles (Scolytidae). A, southern
pine engraver, Ips grandicollis; l) hothole borer,
Scolytus rugulosus.

Order ColeoPtora
Family Curculionidae-Weevils
or Snout Beetles
Most weevils have a well-developed, down-
ward-curved snout. The antennae of most spe-
cies are elbowed and clubbed and the first seg-
ment often fits into a groove in the side of the
snout (Fig. 2l3A). This family is the largest in
the order with over 40,000 species worldwide
and 2,500 species in North America. The great
majority occur and feed on all parts of plants
and many species are important pests. There
are about 42 subfamilies of weevils but only a
few will be described below.

Common Species

Subfamily Cyladinae
Cylas formicarius elegantulus (Summers)
(Fig. 2l3B). Swcetpotato weevil-5-6 mm; reddish €;,
brown; elytra hluish black; anrlike in appcarance;
antennae srraighr; pronotum grearly constricted in
posterior I /3; southern U.S. (larvae in sweet
@
potatoes).

Subfamily Apioninae-1.0-4.5 mm; antennae

not elbowed; abdomen deeply grooved and
broadly oval giving a pear-shaped body; gray or
black;commonly on legumes. ,,

Subfamily Rhynchitinae-Toothnosed Snout

Beetles
Rhynchites bicolor (Fabricius). Rose curculio-
6 mm; red; snout and ventral sides of body black;
elytra very broad; most of U.S, (in rose flowers).

Subfamily Ithycerinae
Ithycerus noveborocensis (Forster). New York
Weevil-12-18 mm; shiny black; gray and brown
pubescence; scutellum yellowish; eastern l/2 North
America (on beech, hickory, oak).

Subfamily Thylacitinae-beak stout, squarish
and widened toward tip; lst antennal segment
passes below eye when pressed against head.
Figure 213 Weevils or snout beetles (Curcu-
lionidae). A, lateral view of the head; B, sweetpotato
weevil, Cylas formicarius elegantulus; C, a white-
fringed beetle, Graphognathus leucoloma; D, straw-
berry root weevil, Otiorhynchus ovatus; E, rice l ;

weevil, Sitophilus oryzae; F, maize billbug, Spåe-

nophorus maidis.

234 Subclass Pterygota

Graphognathus leucoloma (Boheman) (Fig' 2l3C)'
A whitefringed beetle-9'12 mm; blackish; dense
covering of gray or gray-brown scales; pronotum
with 2 whitish longitudinal stripes; elytra with long
hairs and whitish stripe on each lateral margin;
southern U.S.
Panlomorus cervinus (Boheman). Fuller rose wee-
vil-7-9 mm; pale brown; elytra with whitish and
oblique stripe on sides; without hind wings; Far
West.
Subfamily Otiorhynchinae-beak as in sub-
family Thylacitinae; I st antennal segment
passes above middle of eye when pressed
against head.
Otiorhynchus ovatus (Línnaeus) (Fig. 2l3D). Straw'
m; shiny black; antennae
sparse yellowish Pubes-
round and ridged; elYtra
nd rough surface; most of
U.S. (on strawberries).
and
Subfamily Rhynchophorinae-Billbugs
Grain Weevils; antennae insert near eyes, lst
segment extends behind eye, l/2 or more of an-
tennal club enlarged and shinY.
Rhodobaenus tredecimpuncfalus (llliger). Cockle-
bur weevil-7-ll mm; red; pronotum with 5 black
spots, elytra with 8; primarily eastern l/q U.S'
Sitophitus granarius (Linnaeus). Granary wee'
vil-3-4 mm; shiny reddish brown to blackish; pro-
notum with sparse elonlate punctures; elytra deeply
grooved, single rows of punctures; throughout
North America (in stored grain, cereals).
Sitophitus oryzae (Linnaeus) (Fig' 2l3E)' Rice wee-
vil-2.1-2.8 mm; dull brown; pronotum with dense
rounded punctures; elytra with 4 reddish spots'
deeply grooved, double rows deep punctures;
thróughout North America (in stored grain, under
bark, on leaves).
Sphenophorus maidis Chittenden (Fig. 2l 3F). Maize
UlllUug-tl-15 mm; shiny reddish brown to black-
ish; grayish yellow crusty stripes be veen black
stripes; southern U S. (on corn).
Sphenr.tphoru.ç ¿e¿c Walsh-6,5-9.0 ntn ; shiny red-
dish brown to blackish; body sÌ-,ape like S. rraidis,'
pronotum with 3 raised longitudinal bands, central
band wiCest in middle and lateral bands wavy
between bands coarsely punctured; elytra with
rows alternating coarse and fine punctures; eastern
l/2 U.S. (on grasses, corn). S. parvulus
bluegrass billbug, similar shape, 5-8 mm,
gray crust, pronotum without raised bands. , .
Subfamily Cryptorhynchinae-snout held
rest in groove under prosternum; genus cono'
trachelus contains spp. with rough and bumpy
elytra and elyt ra much broader than Pronotum.
Conotrachelus nenuphar (Herbst) (Fig' 2laA).
curculio-4.5-6.5 mm; dark brown; brownish
pubescence; whitish yellow hairs form 2 short
lar bands on pronotum and çrossband on
l/3 of elytra; elytra with broken ridges and
cles; east of Rocky Mts. (on fruit trees,
Subfamily Baridinae-each side of
expanded and visible from above as a bump
tween corners of elytra and pronotum. ,
Trichobaris tinotata (Say) (Fig. 2l4B): Potato
borer-3-4 mm; black; dense white scales gr
trate pu bescence on body except for head,
corners pronotum, and scutellum;
grooves; east of RockY Mts., AZ' CA.
Subfamily Cleoninae-spp. in common
Lixus are elongated and cylindrical, 10-15
curved beak nearlY length of prothorax,
on plants near water
Lixus concavus SaY (Fig 2l4C). Rhubarb
lio-10.0-13.5 mm; black; antennae and tarsi
dish brown; sparse, short grayish pubescence;
orange- brown pollen on fresh sPecimens;
with la rge depression which extends onto
most of U.S. (on rhubarb, curled dock).
Subfamily Hyperinae-on clover and.
all T spp. in genus HYPerø.
Hypera postica (Gyllenhal) (Fig. 2laD). '
weevil-3-5 mm; brown, graY-brown, or
black; dark brown middorsal stripe on elytra'
irregular spots; throughout U.S.
Hypera pun ctata (Fabricius) (Fig. 2l4E)' Clover
weevìl-5.0- 8.5 mm; black; dense t¡rown' Yellow-
bro\\'n, and gray scales give mottled and sti¡rpled ap-
pcararìcc; snottt short; throughout U.lì.
Order Coleoptera
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Figure 214 Weevils or sncut beetles (Curculionidae). A, plum curculio, Conotrachelus

nenuphar; B, potato stalk borer, T¡ichobaris trinotata; C, rhubarb curculio; Lixus concavus;
D, alfalfa weevil, Hypera poslica) E, clover leaf weevil, H. punctqta; F, boll weevil, An-
thonomus grandis; C, an acorn weevil, Curculio sulcatulus.

Subfamily Anthonominae- )100 spp. in ge- Curculio sulcatulus (Casey) (Fig. 2laG)-6-8 mm; ;
nus Anthonornas, snout long. dark or reddish brown; thin covering of short gray- 1

Anthonomus grandis Boheman (Fig. 2laF). Boll
weevil-4,0-7,5 mm; reddish brown to blackish; pale
yellow scalelike hairs irregularly cover body; south-
ern U.S. west to TX (on cotton).
Anlhonomus signatus Say. Strawberry weevil-
2-3 mm; black; antennae and legs reddish brown;
scutellum whitish; elytra reddish brown, 2 black
patches on posterior I /2; beak longer than head and
pronotum; pronot.um coarsely punctured; east of
Rocky Mts.
Subfamily Curculioninae-Acorn and Nut
Weevils; very long and slender snout, all North
American spp, in genus Cørculio, larvae in
acorns, pecans, hickory nuts, other nuts.
brown scales and hairs; eastern l/2U.5. (on oaks).
C. caryøe (Horn), pecan weevil, similar, 7-9 mm,
dull brown with yellowish hairs, outer l/4 of femora
with distinct rectangular tooth projecting at right
angle (on hickory, pecan).
GENERAL REFERENCES
Arnett, R.H,, Jr. 1967. Presenr and Future System-
atics of the Coleoptera in North America.
Ann. Ent, Soc. Amer. 60:162-170.
1968. The Beetles of the United Stotes (A
Manual for ldentification). Amer. Ent. Insti-
tute, AnnArbor, Mich. I,ll2pp.
Baker, \Ã/ L. 1972. Eastern Forest Insects. U.S.
Dept. Agric. For. Serv. Misc, Pulb. 1175.
6a2 pp.

236 Subclass Pterygota